Pics of the Day

An Afar woman (Cushitic).

A Saudi Arab man (Semitic).

A Somali woman (Cushitic).

An Ashkenazi Jew man (Semitic).

An Oromo boy (Cushitic).

An Amhara woman (Agaw Cushitic origin).

Guide to the Universal Basic Genetic Model

The Universal Basic Genetic Model (UBGM) is a genetic model that allows one to quickly determine an individual’s ancestral composition. First introduced on Genome analysis of modern Semitic individuals, it simply consists of the main, least-admixed ancient West Eurasian, East Eurasian, South Eurasian, North African and Sub-Saharan African genome components that are known to exist.

These ancestral elements are, again, as follows:

Least-admixed West Eurasian reference samples:

IRN_Ganj_Dareh_N (Iran Neolithic component)

Levant_Natufian_EpiP (Levantine Natufian component)

AZE_Caucasus_lowlands_LN (Caucasus Hunter-Gatherer component)

TUR_Marmara_Barcin_N (Anatolian Neolithic component)

WHG (Western Hunter-Gatherer component)

Yamnaya_RUS_Samara (European Steppe component)

Least-admixed Sub-Saharan African reference samples:

CMR_Shum_Laka (ancient Pygmy component)

COG_Kindoki_230BP (ancient Niger-Congo component)

KEN_Kakapel_300BP (ancient Nilo-Saharan component)

MWI_Chencherere (ancient East African Hunter-Gatherer component)

ZAF_2000BP (ancient Khoisan component)

Least-admixed Iberomaurusian reference sample:

MAR_Taforalt (Iberomaurusian component)

Least-admixed East Eurasian reference sample:

CHN_Huatuyan_500BP (ancient East Asian component)

Least-admixed South Eurasian reference sample:

LAO_Hoabinhian (Ancient Ancestral South Indian component)

Let us now take a look at each of these ancestral elements, and recap how they relate to the Cushitic peoples and other Afro-Asiatic-speaking populations.

Lebanese Arabs (Semitic). Lebanese, Syrians and other Semitic peoples of the northern Levant and Mesopotamia also carry the Levantine Natufian component at significant frequencies, albeit lower than their kinsmen in the southern Levant and Arabian peninsula. This is due to admixture between ancient Semites in the north and newcomers from the Caucasus, Iranian plateau and Mediterranean, outsiders who introduced the Caucasus Hunter-Gatherer, Iran Neolithic and Anatolian Neolithic genome elements into the ancestral Semitic gene pool.

West Eurasian components

Iran Neolithic component (IRN_Ganj_Dareh)

The Iran Neolithic component is a West Eurasian ancestral element belonging to early farmers of the Zagros Mountains in Iran. It is closely related to the Caucasus Hunter-Gatherer component (discussed below), but differs from it by containing some deeply embedded admixture associated with the Mbuti pygmies of central Africa (cf. Lazaridis et al. (2018)).

Ancient DNA analysis has revealed that the Iran Neolithic element was a core ancestral component borne by the makers of the Indus Valley Civilization (IVC). Iran Neolithic ancestry has been estimated to comprise between 45%-82% of the Indus Valley people’s ancestral makeup, with the rest mainly consisting of the Ancient Ancestral South Indian component (11%-50%) (Narasimhan et al. (2019)).

With regard to modern populations, the Iran Neolithic component attains its highest average frequencies among the Balochi, Makrani and Brahui populations of South/Central Asia:

Global peak frequencies of the Iran Neolithic ancestral component. This West Eurasian genome element attains its highest percentages among Balochi individuals of South/Central Asia (66.2% maximum).

Genetic distance of the Iran Neolithic component to modern populations. The Balochi, Makrani and Brahui have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the Iran Neolithic component reaches a frequency peak among Semitic-speaking Assyrians, Sephardic Jews and Ashkenazi Jews (17.3% among Assyrians). It is generally absent among Afro-Asiatic-speaking populations in Africa.

A Balochi man (Indo-Iranian). The Iran Neolithic ancestral component peaks in frequency among Balochi, Brahui, Makrani and other populations of South/Central Asia.

An Iranian Bandari man (Indo-Iranian). Among Iranians, the Iran Neolithic component peaks in frequency among Bandari individuals.

A Brahui man. Although the Brahui of Pakistan/Afghanistan/Iran today speak a Dravidian language, they predominantly carry the Iran Neolithic genome component rather than the Ancient Ancestral South Indian component which prevails among other Dravidian-speaking populations in the Indian peninsula. This could be because the Brahui originally spoke a non-Dravidian language and later adopted a Dravidian tongue. Alternatively, the Brahui might be relics of the original Dravidian speakers. As such, other Dravidian-speaking populations in South Asia might be language shifters i.e., peoples who originally spoke non-Dravidian languages and later adopted Dravidian tongues.

Levantine Natufian component (Levant_Natufian_EpiP)

The Levantine Natufian component is a West Eurasian ancestral element belonging to the Epipaleolithic Natufian hunter-gatherers of the Levant. It has often been linked with the ancient dispersal of the Afro-Asiatic languages and the Y-DNA haplogroup E.

Ancient DNA analysis has revealed that Natufian ancestry was a core genome component borne by the makers of the Pre-Pottery Neolithic (PPN) industry of the Levant. The PPN culture bearers were, in turn, ancestral to the ancient Canaanites of Israel/Palestine and other early Semitic societies of the northern Levant and Mesopotamia, including the Bronze Age peoples of Jordan, Alalakh and Sidon. In Africa, Levantine Natufian ancestry has also been associated with the spread of pastoralism and the Y-DNA haplogroup T via the Middle Neolithic herders of Skhirat, Morocco (for details on this, see Genetic affinities of the Middle Neolithic pastoralists of Skhirat, Morocco and First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants).

With regard to modern populations, the Levantine Natufian component attains its highest average frequencies among the Semitic-speaking Mahra and other Yemenis, Socotri, Saudi and Arab bedouin populations of the Middle East:

Global peak frequencies of the Levantine Natufian ancestral component. This West Eurasian genome element attains its highest percentages among Yemenite Mahra individuals of the Arabian peninsula (80.2% maximum).

Genetic distance of the Levantine Natufian component to modern populations. The Yemenite Mahra and other “pure” Semitic groups have the closest affinity to this ancestral element.

The Levantine Natufian element constitutes just over half of the predominant non-African ancestry (>70%), which Cushitic peoples bear.

A Socotri girl (Semitic). The Levantine Natufian ancestral component peaks in frequency among the Mahra/Yemeni, Socotri, Saudi and bedouin Arab populations of the Middle East. Ancient DNA analysis of Socotri individuals from the medieval period indicates that they almost entirely bore Levantine Natufian ancestry, with minor Caucasus Hunter-Gatherer/Iran Neolithic admixture and no Sub-Saharan African admixture. This confirms that the Socotri are among the “purest” living descendants of the Levantine Natufians.

A Saudi Arab bedouin man (Semitic). After the Yemenite Mahra, the Levantine Natufian genome component reaches its next highest frequencies among the Arab bedouin populations of the Middle East. The Natufian genome element is also present at varying frequencies in all other Semitic-speaking populations, representing the root ancestry of the Semitic peoples.

Syrian Arab boys (Semitic). Syrians, Lebanese and other Semitic peoples of the northern Levant and Mesopotamia also carry the Levantine Natufian component at significant frequencies, albeit lower than their kinsmen in the southern Levant and Arabian peninsula. This is due to admixture between ancient Semites in the north and newcomers from the Caucasus, Iranian plateau and Mediterranean, outsiders who introduced the Caucasus Hunter-Gatherer, Iran Neolithic and Anatolian Neolithic genome elements into the ancestral Semitic gene pool.

An Assyrian man (Semitic). Assyrians, a Semitic-speaking population inhabiting Mesopotamia, are closely related to Mesopotamian/Levantine Arabs. They bear the same overall ancestry as Iraqi, Syrian, Lebanese, Jordanian and Palestinian individuals, including the Levantine Natufian genome component. However, Assyrians have received more gene flow from outsiders from the Caucasus and Iranian plateau, whereas Levantine Arabs have had greater contacts with foreigners from the Mediterranean.

A Mizrahi Jew man (Semitic). Mizrahi Jews are ethnic Jews who never left the Middle East. As such, they are the root ancestral stock of Ashkenazi Jews and Sephardic Jews (i.e., Mizrahi Jews who intermixed with southern Europeans). Mizrahi Jews have retained a great genetic resemblance to their Palestinian kinsmen. They bear the same overall genome components as Palestinians and at similar frequencies, including the Levantine Natufian component.

An Ashkenazi Jew man (Semitic). Genome analysis indicates that Ashkenazi Jews and Sephardic Jews share ancestral roots with Mizrahi Jews and other Semitic peoples, including the Levantine Natufian component. However, Ashkenazi and Sephardic Jewish individuals carry additional admixtures, acquired through contact between their (mostly male) Semitic ancestors and (mostly female) native southern Europeans.

A Sephardic Jew man (Semitic). Genome analysis indicates that Sephardic Jews are very closely related to Ashkenazi Jews. Sephardic Jewish individuals bear the same overall ancestral elements as Ashkenazi Jews and at comparable frequencies, including the Levantine Natufian component of the earliest Semitic peoples.

An Egyptian man (Egyptian). Archaeogenetic analysis indicates that dynastic Egyptians of the Old Kingdom almost entirely bore Levantine Natufian ancestry. The later dynastic Egyptians from the Old-Middle Kingdom and Middle Kingdom carried less Levantine Natufian ancestry, primarily due to admixture with outsiders bearing the Caucasus Hunter-Gatherer component. Modern Egyptians, both Coptic and Muslim, directly descend from the ancient Egyptians of the Old-Middle Kingdom period.

A Libyan man (Egyptian-Berber origin). Genome analysis indicates that Libyan individuals mostly carry the Levantine Natufian genome component. This ancestral element was derived from a mixture of ancient Egyptian, Middle Neolithic Skhirat, and recent peninsular Arab sources.

An Rbaya girl (Semitic). Genome analysis indicates that the Rbaya bedouins of Tunisia predominantly bear the Levantine Natufian genome component. Almost all of this Levantine Natufian ancestry traces back to a Levant Tell Qarassa source rather than an ancient Egyptian or Middle Neolithic Skhirat source. This confirms the Rbaya’s tradition claiming Arabian origins since all individuals of recent peninsular Arab descent similarly derive most of their Levantine Natufian ancestry from the Levant Tell Qarassa source.

A Douz man (Semitic). Genome analysis indicates that the Douz bedouins of Tunisia predominantly bear the Levantine Natufian genome component. For many Douz individuals, much of this ancestry traces back to a Skhirat source. Other Douz persons (like Rbaya individuals) derive much of their Natufian ancestry from a Levant Tell Qarassa source, confirming their tradition of recent peninsular Arab origin.

A Somali man (Cushitic). Genome analysis indicates that, on average, Northern Somalis born in the northeastern Puntland region of Somalia carry the most non-African ancestry among Afro-Asiatic speakers in the Horn. This non-African ancestry is comparable in frequency to that borne by North African individuals (cf. Fig. S1 in Osman and Jonasson (2022)). It also includes a notable Levantine Natufian component, which was largely inherited from the Middle Neolithic pastoralists of Skhirat, Morocco. These ancient Skhirat herders bore the Y-DNA haplogroup T and the mtDNA haplogroup M1, non-African clades which are signature uniparental lineages of the Afro-Asiatic-speaking populations in the Horn and Nile Valley (up to 80% of Somalis still carry the haplogroup T, a global peak; cf Plaster (2011)).

An Oromo woman (Cushitic). Cushitic peoples, both ancient and modern, bear a predominant non-African ancestry (over 70% on average), of which the Levantine Natufian component constitutes a major element. Most of the rest of the non-African ancestry carried by Cushitic individuals consists of Eurasiatic components (i.e. European Steppe and ancient East Asian elements).

Afar women (Cushitic). The Afar are a Cushitic-speaking population inhabiting areas in Djibouti, Ethiopia and Eritrea. Genome analysis indicates that they bear the same overall ancestral composition as other Cushitic-descended peoples, including the Levantine Natufian component.

A Beja man (Cushitic). The Beja are today the northernmost Cushitic peoples, inhabiting areas in Eritrea, Sudan and Egypt. Genome analysis indicates that they share the same overall ancestral makeup as other Cushitic individuals, including the Levantine Natufian component.

An Amhara woman (Agaw Cushitic origin). Although the Amhara and Tigray-Tigrinya today speak Semitic languages, they are genetically, physically and culturally most similar to the Cushitic-speaking Agaws, who live right beside them in the Ethiopian and Eritrean highlands. Genome analysis indicates that, like Agaws and other Cushitic speakers, the Amhara and other Abyssinians/Habesha predominantly bear non-African ancestry (over 70% on average), of which the Levantine Natufian component constitutes a major element. This suggests that Amhara and Tigray-Tigrinya originally spoke Cushitic Agaw languages and adopted their present Semitic tongues from Sabaean outsiders. Consequently, the Amharic and Tigrinya languages today both have an Agaw substratum, which was retained from the Cushitic languages that the Amhara’s and Tigray-Tigrinya’s Agaw ancestors originally spoke. (N.B. Although Abyssinians/Habesha are of Cushitic Agaw origin, they are genetically differentiated from the Agaw proper by having greater East African hunter-gatherer and peninsular Arab influences. For the Amhara, these admixtures are estimated at 16% apiece by Hodgson et al. (2014) (cf. Table 2 and Table S6).)

An Ethiopian Jew woman (Agaw Cushitic origin). Like Abyssinians, Ethiopian Jews (Beta Israel) share their core ancestry with Cushitic-speaking Agaws, but also have some extra admixtures derived from contact with both East African hunter-gatherers and Semitic peoples. The forager admixture of Ethiopian Jews and Abyssinians is, in particular, reflected in their high altitude adaptation alleles and in their elevated frequencies of the Y-DNA haplogroup A, an archaic African paternal lineage (17%-20% of Amhara, up to 36% of Tigrinya, and as high as 41% of Ethiopian Jews bear the A clade; cf. Wood et al. (2005), Appendix A; Scozzari et al. (2014), Table S7; Cruciani et al. (2002), Table 2; Gebremeskel (2018), Table 3.7; also see Alkorta-Aranburu et al. 2012).

An Iraqw woman (Cushitic). Along with the Alagwa, Gorowa and Burunge, the Iraqw are the last remaining South Cushites in the Great Lakes region. Although they have intermixed with and assimilated their Bantu/Nilotic/hunter-gatherer neighbors, the Iraqw proper have managed to remain physically and genetically distinct. Genome analysis indicates that Iraqw individuals predominantly bear non-African ancestry (~63%). Like other Cushitic peoples, this non-African heritage consists of a mixture of the Levantine Natufian, European Steppe and ancient East Asian components.

A Nubian boy (Cushitic origin). Although Nubians of the Nile Valley today speak a Nilo-Saharan language, genome analysis indicates that they predominantly bear non-African ancestry (over 70% on average), of which the Levantine Natufian genome component constitutes a major element. As such, many modern Nubians appear to be direct descendants of the “Red Nubians” (i.e. the Egyptian-related inhabitants of ancient Nubia). This, in turn, suggests that they originally spoke an Afro-Asiatic language, and adopted their present Nilo-Saharan tongue from adjacent Nilotic peoples (i.e. descendants of the Nilo-Saharan-related “Black Nubians”).

A Sudanese “Arab” man (Nubian origin). Like Cushitic speakers and Nubians, Sudanese “Arabs” also predominantly bear non-African ancestry (over 70% on average), of which the Levantine Natufian component is a major element. However, proximal genome analysis indicates that, unlike peninsular Arabs, most of the Levantine Natufian ancestry which Sudanese “Arabs” carry was derived from Middle Neolithic Moroccan (Skhirat) and ancient Egyptian sources rather than the peninsular Arab-associated Levant Tell Qarassa source. This suggests that Sudanese “Arabs” are actually of Nubian origin and adopted their present Arabic language during the Middle Ages, with the spread of Islam.

A Benadiri Somali man (Azanian Cushitic origin). Like most Muslims in the Horn, the Benadiri/Bravanese of southern Somalia have traditions of descent from various Arabian forefathers. However, haplogroup and genome analyses of Benadiri/Bravanese individuals indicate that they are of Cushitic Somali origin, with secondary Persian, Indian and Bantu admixtures. As such, Benadiri/Bravanese individuals carry the Levantine Natufian component at elevated frequencies.

A Wolayta woman (North Omotic). Genome analysis indicates that the North Omotic-speaking Wolayta, Shekecho and Shinasha predominantly bear non-African ancestry (over 70% on average), of which the Levantine Natufian genome component constitutes a major element. As such, North Omotic speakers appear to directly descend from the original Omotic peoples, who spoke Afro-Asiatic languages. (*N.B. North Omotic speakers today bear the highest frequencies of the haplogroup J in the Horn region (~50%; cf. Plaster (2011)). This Y-DNA lineage is associated with the spread of Caucasus Hunter-Gatherer-related ancestry. As such, it is intrusive to the Afro-Asiatic speech area of the Middle East, the Horn of Africa and North Africa.)

A Toubou woman. Genome analysis indicates that the Toubou of Chad and Sudan carry significant non-African ancestry (~60% on average). This non-African ancestral heritage consists of the Levantine Natufian component, as well as European Steppe and ancient East Asian elements.

A Baggara/Shuwa “Arab” woman (Chadic origin). The Arabic-speaking Baggara of Chad and Sudan, who are known as Shuwa “Arabs” in Nigeria, carry the Levantine Natufian component at significant frequencies. However, genome analysis indicates that virtually all of their Natufian ancestry was derived from the Middle Neolithic pastoralists of Skhirat, Morocco rather than from a more direct Levantine/peninsular Arab source. Modern Chadic speakers, who primarily bear the Niger-Congo ancestral component, also carry a trace of the Baggara’s specific non-African ancestry. This suggests that the Baggara/Shuwa originally spoke a Chadic language(s) and adopted Arabic relatively recently, during the medieval Islamic period. As such, they represent a vestige of the original Chadic speakers, who transmitted their Afro-Asiatic language(s) to the formerly Niger-Congo-speaking ancestors of present-day Chadic speakers.

Caucasus Hunter-Gatherer component (AZE_Caucasus_lowlands_LN)

The Caucasus Hunter-Gatherer component is a West Eurasian ancestral element belonging to Neolithic period hunter-gatherers from the Caucasus Mountains. It is associated with the ancient dispersal of the Y-DNA haplogroup J.

With regard to modern populations, the Caucasus Hunter-Gatherer component attains its highest frequencies among Georgians and other communities inhabiting the Caucasus area:

Global peak frequencies of the Caucasus Hunter-Gatherer ancestral component. This West Eurasian genome element attains its highest percentages among Pontic Greek individuals of Trabzon, Turkey (89.2% maximum).

Genetic distance of the Caucasus Hunter-Gatherer component to modern populations. Armenians, eastern Greeks and Trabzon Turks have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the Caucasus Hunter-Gatherer component reaches a frequency peak among Assyrians (56.4%).

A Georgian man (South Caucasian). The Caucasus Hunter-Gatherer ancestral component peaks in frequency among Georgians and other populations of the Caucasus Mountains.

A Pontic Greek woman. Pontic Greeks from Trabzon, Turkey have been found to carry especially high frequencies of the Caucasus Hunter-Gatherer genome component, an ancestral element which constitutes most of their ancestry. This suggests that many Greek speakers in eastern areas descend from Hellenized locals, whereas Greek speakers in Achaea and other western areas directly descend from the ancient Mycenaeans.

An Anatolian Turk man. Among Turkish speakers, the Caucasus Hunter-Gatherer ancestral component reaches a frequency peak among Anatolian Turks from Trabzon (76.1% on average). Trabzon Turks also have very little of the ancient East Asian component (1.5%). Altogether, this suggests that Anatolian Turks from the eastern Trabzon area descend from local inhabitants (likely originally South Caucasian speakers), who later adopted the Turkish language.

An Azerbaijani man. Azerbaijanis (Azeris), another Turkic-speaking ethnic group, also bear the Caucasus Hunter-Gatherer component at elevated percentages. Frequencies of this ancestral element peak among Azeris of Turkey (averaging nearly 70%). Azerbaijanis of Dagestan carry the Caucasus Hunter-Gatherer component at lower frequencies, due to a higher Yamnaya-related Steppe influence in this area.

Anatolian Neolithic component (TUR_Marmara_Barcin_N)

The Anatolian Neolithic component is a West Eurasian ancestral element belonging to the first agriculturalists of Anatolia/Turkey. It is the defining ancestry of the Early European Farmers, and still comprises one of the core elements of European heritage. Ancient DNA analysis also indicates that the Early Neolithic occupants of Kahf Taht Al Ghar, an archaeological site in Morocco, entirely bore Anatolian Neolithic ancestry. These ancient cultivators appear to have been Early European Farmers who arrived from Iberia (cf. Simões (2023a)).

With regard to modern populations, the Anatolian Neolithic component attains its highest frequencies among Sardinians and other communities native to southern Europe:

Global peak frequencies of the Anatolian Neolithic ancestral component. This West Eurasian genome element attains its highest percentages among Sardinian individuals of southern Europe (80.8% maximum).

Genetic distance of the Anatolian Neolithic component to modern populations. Sardinians, Corsicans, western Greeks and other southern European groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the Anatolian Neolithic component reaches a frequency peak among northern Maghrebi populations, Ashkenazi Jews and Sephardic Jews (32.8% among Ashkenazi Jews of Belarus) i.e., Afro-Asiatic-speaking groups that received significant gene flow from southern Europeans.

A Sardinian woman (Romance). The Anatolian Neolithic ancestral component peaks in frequency among Sardinians and other populations native to southern Europe.

A Corsican girl (Romance). After Sardinians, the Anatolian Neolithic genome component reaches its next highest frequencies among Corsicans.

A Greek woman from Achaea, western Greece (Paleo-Balkan). Achaean Greeks have among the highest frequencies of the Anatolian Neolithic genome component. This suggests that they directly descend from the ancient Mycenaeans, the earliest attested Greek speakers, who archaeogenetic analysis has found mostly bore Anatolian Neolithic ancestry.

A Spanish man (Romance). After Sardinians and Corsicans, Spaniards and other Iberians have among the highest average frequencies of the Anatolian Neolithic component (~45%).

A Portuguese man (Romance). Portuguese carry the Anatolian Neolithic genome component at an average frequency similar to that of Spaniards (~45%).

An Albanian man (Paleo-Balkan). Albanians of the Balkans carry the Anatolian Neolithic component at a mean percentage comparable to that of southwestern Europeans (~45%).

A Northern Italian man (Romance). Northern Italians carry the Anatolian Neolithic genome component at an average frequency similar to that of Iberian and Albanian populations (~45%).

A Portuguese Brazilian man (Romance). Many Brazilians and other Latin Americans are of recent Portuguese or Spanish descent. As such, they have the same overall ancestral composition as their brethren in Iberia, including the signature Anatolian Neolithic genome component of southern Europe.

An Andalusian Spanish man (Romance). Genome analysis indicates that Spaniards from Andalusia share the same overall ancestral composition as other Iberians and southern Europeans, including the Anatolian Neolithic component.

A Pardo Brazilian woman (Romance). Pardos are Brazilians of mixed heritage. Genome analysis indicates that many Pardo individuals are predominantly of Portuguese descent (over 70% Portuguese ancestry). As such, they bear high frequencies of the signature Anatolian Neolithic component of southern Europe.

A Southern Italian man (Romance). Since the Imperial Rome period, when the former Roman Republic expanded its holdings beyond southern Europe to include territories in the Middle East (e.g. the province of Judaea), many parts of southern Italy and all of Sicily have served as contact areas where native southern Europeans have intermixed with Middle Eastern peoples. Consequently, Southern Italians and Sicilians today have significantly lower average frequencies of the Anatolian Neolithic component than their Northern Italian counterparts. Moreover, in Principal Component Analysis charts, Sicilians (peoples of European origin with substantial Middle Eastern admixture) cluster in between North Italian and Levantine individuals, near Maltese and Ashkenazi/Serphardic Jews (peoples of Middle Eastern origin with substantial European admixture).

Western Hunter-Gatherer component (WHG)

The Western Hunter-Gatherer component is a West Eurasian ancestral element belonging to Mesolithic period foragers of Europe. Although widely distributed throughout the continent, it is a vestigial ancestry which only exists today at low frequencies.

With regard to modern populations, the Western Hunter-Gatherer component attains its highest frequencies among Basques:

Global peak frequencies of the Western Hunter-Gatherer ancestral component. This West Eurasian genome element attains its highest percentages among Basque individuals of southern Europe (21.6% maximum).

Genetic distance of the Western Hunter-Gatherer component to modern populations. The Basque and certain western and eastern European groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the Western Hunter-Gatherer component reaches a frequency peak among Ashkenazi Jews (1.6% among Ashkenazi Jews of Ukraine), due to admixture with Europeans.

A Basque man (Basque). The Western Hunter-Gatherer ancestral component peaks in frequency among Basque individuals of southern Europe.

European Steppe component (Yamnaya_RUS_Samara)

The European Steppe component is a West Eurasian ancestral element belonging to nomads of the Pontic-Caspian Steppe in eastern Europe. It is associated with the ancient spread of the Indo-European languages and the Y-DNA haplogroup R1. Along with the Anatolian Neolithic component of the Early European Farmers and (to a lesser extent) the WHG component of the Mesolithic Western Hunter-Gatherers, the Steppe component forms the core ancestry borne by modern European individuals.

With regard to contemporary populations, the European Steppe component attains its highest frequencies among the Khanty, Udmurt, Selkup, Ket and other communities of northern and eastern Europe:

Global peak frequencies of the European Steppe ancestral component. This West Eurasian genome element attains its highest percentages among Khanty individuals of northern Europe (69.8% maximum).

Genetic distance of the European Steppe component to modern populations. The Akhvakh, Tajik, Tabasaran and other Indo-European and Northeast Caucasian-speaking groups of the Caucasus and Central Asia have the closest affinity to this ancestral element.

Among Semitic-speaking populations, the European Steppe component reaches a frequency peak among Ashkenazi Jews and Sephardic Jews (21.2% among Ashkenazi Jews of Belarus), due to admixture with Europeans.

On the Universal Basic Genetic Model, which presents a rough overview of a person’s ancestral composition, Cushitic individuals show a low European Steppe ancestry of under 10%. However, more precise distal genome analysis indicates that they actually carry significant European Steppe ancestry, averaging around 20% (see Detecting the Eurasiatic genome component in Cushitic peoples). Consequently, many Cushitic individuals have been observed to bear European Steppe-associated mtDNA haplogroups, such as the I clade and haplogroup H derivatives.

An Udmurt woman (Uralic). The European Steppe ancestral component peaks in frequency among the Udmurt, Khanty and other populations of northern and eastern Europe.

Khanty women (Uralic). Along with the Udmurt, the Khanty people of northern Russia bear the highest average frequencies of the European Steppe genome component.

A Russian man (Slavic). After Uralic populations, the European Steppe genome component reaches its next highest frequencies among Russians and other Slavic peoples of eastern Europe (~60% on average).

A Danish man (Germanic). Danes and other Germanic peoples of northern Europe bear the European Steppe genome component at a mean percentage comparable to that of Slavic peoples (~60%).

An Irish man (Celtic). The Irish and other Celtic peoples of Ireland and Great Britain carry the European Steppe genome component at an average frequency similar to that of Slavic and Germanic populations (~60%).

A Kalash girl (Indo-Iranian). In South/Central Asia, the European Steppe genome component is found at highest frequencies among the Kalash of Pakistan. Kalash individuals have been observed to bear as high as 50% of Yamnaya-related ancestry, a Steppe percentage typical for many areas in western Europe. This heritage is often reflected in the physiognomy of Kalash individuals, who are frequently pale-skinned and blond or red-haired, with light-colored eyes — all traits inherited from their European Steppe ancestors.

Tajik women (Indo-Iranian). After the Kalash, the European Steppe genome component reaches its next highest frequencies in South/Central Asia among the Persian-speaking Tajik of Afghanistan, Tajikstan and Uzbekistan. Tajik individuals in the southeastern Rushan district of Tajikstan have been observed to bear Steppe percentages averaging almost 50%.

A Pashtun man (Indo-Iranian). The Pashtun of northwestern Pakistan and southern and eastern Afghanistan are another Indo-European-speaking people, who also carry the European Steppe component at high percentages. Pashtun individuals have been observed to bear Yamnaya-related ancestry at frequencies of up to 40%.

A Zoroastrian woman (Indo-Iranian). Iranian peoples are characterized by a mixture of the Caucasus Hunter-Gatherer, Iran Neolithic, European Steppe, Anatolian Neolithic and Levantine Natufian genome components. Among Iranians, Zoroastrians bear the highest average percentage of the European Steppe ancestral element, a component which in certain individuals occurs at frequencies of almost 30%. Like other Iranians, Zoroastrians also have a very low frequency of the Ancient Ancestral South Indian component (1.1%).

Sub-Saharan African components

Ancient Pygmy component (CMR_Shum_Laka)

The ancient Pygmy component is a Sub-Saharan African ancestral element belonging to early Pygmy-related hunter-gatherers of Shum Laka, Cameroon.

With regard to modern populations, the ancient Pygmy component attains its highest frequencies among the Baka, Bakola, Biaka, Mbuti and other African Pygmy groups:

Global peak frequencies of the Pygmy ancestral component. This Sub-Saharan African genome element attains its highest percentages among Baka individuals of central Africa (100% maximum).

Genetic distance of the ancient Pygmy component to modern populations. The Bedzan, Bakola and other Pygmy groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the ancient Pygmy component reaches a frequency peak among Bedouin Arabs of the Levant (0.4% among BedouinA from the Negev desert). It is generally not carried by most Afro-Asiatic-speaking individuals.

A Baka man (Pygmy). The Pygmy ancestral component peaks in frequency among Baka, Bakola, Biaka, Mbuti and other Pygmies of central Africa.

Ancient Niger-Congo component (COG_Kindoki_230BP)

The ancient Niger-Congo component is a Sub-Saharan African ancestral element belonging to early agriculturalists of Kindoki, Congo. It represents the principal ancestry borne by Niger-Congo speakers in western Africa and their descendants, including African Americans and Afro-Caribbeans.

With regard to modern populations, the ancient Niger-Congo component attains its highest frequencies among the Esan, Mende, Yoruba and other Niger-Congo-speaking individuals in western Africa. This element is also borne at elevated percentages by the Hausa and other Chadic speakers, who today speak Afro-Asiatic languages, but appear to have originally spoken Niger-Congo languages:

Global peak frequencies of the Niger-Congo ancestral component. This Sub-Saharan African genome element attains its highest percentages among Esan individuals of western Africa (96.2% maximum).

Genetic distance of the ancient Niger-Congo component to modern populations. The Ngumba, Umbundu and other West African groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the ancient Niger-Congo component reaches a frequency peak among individuals in southern Morocco and other such contact zones, where Afro-Asiatic speakers have historically interacted with Niger-Congo speakers. In other parts of the Maghreb and elsewhere, most Afro-Asiatic speakers do not bear any Niger-Congo-related admixture (for further discussion on this, see Y-DNA haplogroup E1b1a (formerly E3a) appears to have originated in the Middle East).

An Esan man (Niger-Congo). The Niger-Congo ancestral component peaks in frequency among the Esan, Mende, Yoruba and other populations of western Africa, including the Hausa.

A Yoruba man (Niger-Congo). After the Esan and Mende, the ancient Niger-Congo genome component reaches its next highest frequencies among the Yoruba of Nigeria.

Igbo Jews (Niger-Congo). Genome analysis indicates that the Igbo people of Nigeria almost entirely carry the ancient Niger-Congo component. More controversially, it has been claimed that Igbo Jews (a subset of the Igbo population who practice Judaism) also largely bear this ancestral element, with no trace of Jewish-related Levantine ancestry.

Hausa men (Niger-Congo). The Hausa of the Sahel today speak a language which belongs to the Chadic branch of the Afro-Asiatic family. However, they bear a close physical resemblance to neighboring Niger-Congo speakers. Genome analysis also indicates that the Hausa predominantly bear the ancient Niger-Congo component. Altogether, this suggests that the Hausa originally spoke a Niger-Congo language and later adopted an Afro-Asiatic tongue.

A Fulani woman (Niger-Congo). Genome analysis indicates that the Fulani of the Sahel predominantly carry the ancient Niger-Congo component. Frequencies of this ancestral element vary depending on location, from a high of ~80% among the Fulani of Guinea to a lower 60% average among the Fulani of Ziniare, Burkina Faso. The rest of the Fulani’s ancestry consists of a mixture of North African Iberomaurusian, West Eurasian and East Eurasian components. Further genetic analysis suggests that the Fulani acquired their Iberomaurusian element as well as their cattle pastoralism tradition from the Early Neolithic herders of Takarkori, Libya.

A Bella man (Niger-Congo). The Bella, also known as Ikelan, were formerly slaves of the Tuareg Berbers. Serological and craniometric analysis indicates that they are of West African origin, descending from Niger-Congo speakers. As such, the Bella predominantly carry the ancient Niger-Congo genome component. Most Bella/Ikelan individuals today also have a bit of Berber admixture, derived from interaction with the Tuareg proper.

A Yao Bantu man. The Yao, Makhuwa, Nyanja and other southeast African Bantus are documented ancestors of the Bantu peoples of Somalia. Genome analysis indicates that they bear a predominant Sub-Saharan African ancestry (>95%), which consists of an almost equal apportionment of the ancient Niger-Congo component and ancient Nilo-Saharan component.

An African American man (Niger-Congo). The Niger-Congo ancestral component comprises most of the ancestry of African American individuals (>80% on average), due to their recent origins in West Africa.

A Jamaican man (Niger-Congo). The Niger-Congo ancestral component also comprises most of the ancestry of Afro-Carribean individuals (>80% on average), due to their recent origins in West Africa.

Haitians (Niger-Congo). Among African American/Afro-Caribbean individuals, Haitians carry the highest mean frequency of the ancient Niger-Congo component. They average ~90% of this Sub-Saharan African ancestral element, a percentage typical for populations in western Africa.

Ancient Nilo-Saharan component (KEN_Kakapel_300BP)

The ancient Nilo-Saharan component is a Sub-Saharan African ancestral element belonging to early modern inhabitants of Kakapel, Kenya. It comprises most of the ancestry carried by both Nilo-Saharan and Niger-Congo speakers in eastern Africa. As such, it appears to be the genomic analogue to the linguist Roger Blench’s Niger-Saharan macrophylum, which posits that “Niger-Congo and Nilo-Saharan are genetically related and should be regarded as a single macrophylum” (cf. Blench (2008)).

Admixture runs suggest that the ancient Nilo-Saharan component is related to the ancient Niger-Congo component. However, it differs from the latter by the fact that it lacks alleles derived from archaic African specimens, which are embedded within Niger-Congo ancestry (for more on this archaic African admixture, see here).

With regard to modern populations, the ancient Nilo-Saharan component attains its highest frequencies among the Luo Nilotes and Luhya Bantus:

Global peak frequencies of the Nilo-Saharan ancestral component. This Sub-Saharan African genome element attains its highest percentages among Luhya individuals of eastern Africa (99% maximum).

Genetic distance of the ancient Nilo-Saharan component to modern populations. The Luhya, Luo and other Bantu/Nilotic groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the ancient Nilo-Saharan component reaches a frequency peak among certain Sudanese “Arab” individuals (as high as ~50% among some Kababish). However, among other Sudanese “Arabs” and Afro-Asiatic speakers generally, it occurs at low-to-moderate frequencies.

A Luo man (Nilotic). The Nilo-Saharan ancestral component peaks in frequency among the Luo, Luhya and other Nilotic and Bantu populations of eastern Africa.

A Dinka man (Nilotic). Genome analysis indicates that the Dinka, Nuer, Shilluk and other northern Nilotes of the Nile Valley predominantly bear the ancient Nilo-Saharan ancestral component. However, these northern Nilotic individuals also possess significant Eurasiatic admixture (i.e., European Steppe and ancient East Asian components), which was likely derived from contacts with the Predynastic Egyptians.

A Tutsi man (Bantu). The Tutsi and Hutu, Bantu speakers of Nilotic origin, also predominantly carry the ancient Nilo-Saharan ancestral component. This shared heritage is reflected in the uniparental markers the Tutsi and Hutu bear, which mostly consist of the paternal E1b1a haplogroup and maternal L macrohaplogroup derivatives like other Bantus/Nilotes in the Great Lakes region (cf. Luis et al. (2004), Figure 1; Göbel et al. (2019)). (*N.B. Miao et al. (2024) analysed the genomes of Rwandan individuals and observed a close affinity between this sample and their Nilotic Luo sample from Kenya. Genetic distance analysis also indicates that the Tutsi trace much of their recent ancestry to the Nilotes of the Pastoral Iron Age, consistent with the Universal Basic Genetic Model.)

A Hima man (Bantu). The Hima of Uganda, another Bantu-speaking population of Nilotic origin closely related to the Tutsi and Hutu, also predominantly carry the ancient Nilo-Saharan ancestral component.

A Hutu man (Bantu). Like their Bantu kinsmen the Tutsi and Hima, the Hutu of Rwanda predominantly carry the ancient Nilo-Saharan genome component.

An Ogiek woman (Nilotic). The Ogiek, a Nilo-Saharan-speaking population of Kenya, mostly bear the ancient Nilo-Saharan component. Genetic distance analysis also indicates that, among all modern groups, the Ogiek are the people with whom the Tutsi and Hima Bantus share the closest affinity. This further confirms the Nilotic ancestral origin of the Tutsi-Hima.

A Maasai man (Nilotic). The Maasai of Kenya and Tanzania are closely related to the Ogiek Nilotes and Tutsi Bantus. As such, they predominantly bear the signature ancient Nilo-Saharan genome component of the Bantu and Nilotic peoples.

A Kalenjin man (Nilotic). The Kalenjin Nilotes of the Great Lakes region are closely related to the Maasai Nilotes. Genome analysis indicates that they too predominantly carry the ancient Nilo-Saharan genome component.

A Kikuyu man (Bantu). As with all Bantu/Nilotic peoples in eastern Africa, from the Great Lakes region to the Nile Valley, the ancient Nilo-Saharan genome component comprises most of the ancestry of the Kikuyu.

A Samburu man (Nilotic). The Samburu Nilotes of Kenya are closely related to the Maasai and Kalenjin Nilotes. As such, they predominantly carry the ancient Nilo-Saharan genome component.

A Datog woman (Nilotic). The Datog Nilotes of Tanzania are closely related to other Nilotic peoples in the Great Lakes region. Genome analysis indicates that they predominantly carry the ancient Nilo-Saharan genome component.

An Ari woman (South Omotic speaker). In contrast to the North Omotic-speaking groups, the Ari and other South Omotic-speaking individuals are both phenotypically and genetically distinct from other local Afro-Asiatic speakers. The South Omotic speakers are instead similar to Niger-Congo and Nilo-Saharan-speaking populations inhabiting the Great Lakes region. This suggests that the Ari and other South Omotic speakers are language shifters i.e., their forager ancestors originally spoke non-Afro-Asiatic tongues (Gopalan et al. (2019); Boattini et al. (2013)).

A Swahili man (Bantu). Although the modern Swahili Bantus claim Persian descent, genome analysis indicates that they predominantly carry the ancient Nilo-Saharan genome component. They also have low-to-moderate non-African admixture (~20% on average), most of which was derived from Cushitic peoples whom their Bantu ancestors absorbed.

A Malagasy man (Blasian). The Malagasy are a Blasian population inhabiting the island of Madagascar. Genome analysis indicates that they are of Bantu origin, with significant Austronesian (Indonesian) admixture. As such, the ancient Nilo-Saharan component of the Bantu/Nilotic peoples forms the core of the Malagasy’s ancestry.

Akhdam children (Afro-Arab). The Akhdam (Al-Akhdam) are an Afro-Arab population inhabiting southern Arabia. Genome analysis indicates that they are of Bantu origin, with significant Arabian admixture. As such, the ancient Nilo-Saharan component of the Bantu/Nilotic peoples forms the core of the Akhdam’s ancestry.

Siddi men (Bantu). The Siddi of the Indian peninsula, also known as Habshi, are descendants of slaves from the Great Lakes region in eastern Africa. Genome analysis indicates that they mostly carry Bantu-related ancestry, with minor Indian admixture from contact with locals. As such, Siddi individuals also predominantly bear the ancient Nilo-Saharan component of the Bantu/Nilotic peoples.

Ancient East African Hunter-Gatherer component

The ancient East African Hunter-Gatherer component is a Sub-Saharan African ancestral element belonging to early foragers of Chencherere, Malawi.

With regard to modern populations, the ancient East African Hunter-Gatherer component attains its highest frequencies among Hadza forager individuals:

Global peak frequencies of the East African Hunter-Gatherer ancestral component. This Sub-Saharan African genome element attains its highest percentages among Hadza individuals of eastern Africa (33.8% maximum).

Genetic distance of the ancient East African Hunter-Gatherer component to modern populations. Southeastern Bantu, Hadza, Sandawe, Ari and other forager groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the ancient East African Hunter-Gatherer component reaches a frequency peak among Iraqw individuals in Tanzania (~20%).

A Hadza man (Hadza). The East African hunter-gatherer ancestral component peaks in frequency among the Hadza foragers of eastern Africa.

A Tehla Sandawe woman. The Sandawe foragers of the Great Lakes region carry the East African Hunter-Gatherer component at significant frequencies. Genome analysis indicates that Sandawe individuals also bear substantial non-African admixture (~30%), which was derived from Cushitic peoples. This Cushitic admixture is highest among the Tehla, the pastoralist division of the Sandawe, and is reflected in their distinct physiognomy.

Ancient Khoisan component (ZAF_2000BP)

The ancient Khoisan component is a Sub-Saharan African ancestral element belonging to early foragers of southern Africa.

With regard to modern populations, the ancient Khoisan component attains its highest frequencies among Khoisan forager individuals:

Global peak frequencies of the Khoisan ancestral component. This Sub-Saharan African genome element attains its highest percentages among Khoisan individuals of southern Africa (100% maximum).

Genetic distance of the ancient Khoisan component to modern populations. The Khoisan have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the ancient Khoisan component reaches a frequency peak among Ashkenazi Jews (0.2% among Ashkenazi Jews of Lithuania). It is generally absent among other Afro-Asiatic-speaking populations. (For specifics, see Ancient Khoisan were dark-skinned and did not look Eurasiatic.)

A Khoisan man (Khoisan). The Khoisan ancestral component peaks in frequency among Khoisan individuals of southern Africa.

North African component

Iberomaurusian component (MAR_Taforalt)

The ancient Iberomaurusian component is a North African ancestral element belonging to Epipaleolithic hunter-gatherers of northern Africa, who are associated with the Iberomaurusian archaeological culture. It is closely related to the Levantine Natufian component, but differs from it by containing ~33% of Sub-Saharan African admixture (cf. Ferreira et al. (2021)).

With regard to modern populations, the Iberomaurusian component attains its highest frequencies among Berbers and other Maghrebi populations:

Global peak frequencies of the Iberomaurusian ancestral component. This North African genome element attains its highest percentages among Berbers and other Maghrebi individuals in northwestern Africa (54.6% maximum).

Genetic distance of the North African Iberomaurusian component to modern populations. Moroccan Berbers and other Maghrebi groups have the closest affinity to this ancestral element.

The Iberomaurusian ancestral component is also present at low frequencies among most other Afro-Asiatic speakers. Among Cushitic speakers, Sudanese “Arabs” and other pastoralist groups in the Horn of Africa, Nile Valley and Chad Basin, this genome element was mostly acquired via the Middle Neolithic pastoralists of Skhirat (who bore ~14% of Iberomaurusian-related admixture).

A Berber man (Berber). The Iberomaurusian ancestral component peaks in frequency among Berbers and other Maghrebi individuals of northwestern Africa.

A Tuareg Berber woman (Berber). After Berbers from Morocco, genome analysis indicates that Tuareg and Sahrawi individuals carry the Iberomaurusian ancestral component at highest average frequencies.

A Moroccan “Arab” woman (Berber origin). Although some communities and individuals in Northwest Africa are of recent peninsular Arab descent (such as the Rbaya of Tunisia as well as many Tunisian Douz), genome analysis suggests that most Arabic-speaking Maghrebis are actually of Berber origin. Their Berber-speaking ancestors seem to have begun adopting the Arabic language during the Middle Ages, a period of intensified contact with peninsular Arabs. Berber and Arabic speakers of the Maghreb are thus genetically closely related, sharing similar ancestry, of which the North African Iberomaurusian genome component represents a major element.

Southern Moroccans. Southern Morocco is traditionally a contact zone, where Afro-Asiatic speakers have interacted with Niger-Congo speakers. Consequently, Berber and Arabic-speaking Maghrebis in this area carry the Iberomaurusian genome component at lower than average frequencies and the ancient Niger-Congo genome component at higher than average frequencies.

East Eurasian component

Ancient East Asian component (CHN_Huatuyan_500BP)

The ancient East Asian component is an East Eurasian ancestral element belonging to early inhabitants of Huatuyan, China.

With regard to modern populations, the ancient East Asian component attains its highest frequencies among Gelao and Hmong individuals:

Global peak frequencies of the East Asian ancestral component. This East Eurasian genome element attains its highest percentages among Gelao individuals in eastern Asia (73.2% maximum).

Genetic distance of the ancient East Asian component to modern populations. The Hmong and other East Asian groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the ancient East Asian component reaches a frequency peak among Somalis and other Cushitic individuals in the Horn. It is also present at lower frequencies among most other Afro-Asiatic-speaking populations.

Along with the European Steppe component, this ancient East Asian element appears to have spread from South/Central Asia into Northeast Africa, thereby introducing Eurasiatic (i.e. European Steppe and ancient East Asian) ancestry to these areas.

(*N.B. Genome analysis using a broader Global25 database of over 7000 ancient samples indicates that the East Asian ancestry found in Northeast Africa is actually Upper Paleolithic European ancestry. This ancestry dates from a period when European and East Asian populations had not yet separated, explaining the East Asian affinities contained within it. For details, see Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa.)

A Hmong woman (East Asian). The East Asian ancestral component peaks in frequency among the Gelao, Hmong and other populations of eastern Asia.

South Eurasian component

Ancient Ancestral South Indian component (LAO_Hoabinhian)

The Ancient Ancestral South Indian component (AASI) is a South Eurasian ancestral element belonging to early inhabitants of the Hoabinhian cultural complex in southeast Asia.

With regard to modern populations, the Ancient Ancestral South Indian component attains its highest frequencies among the Onge and Jarawa of the Andaman Islands, as well as tribal and Dravidian-speaking groups in southern Asia. For this reason, the AASI element is also known as the Onge component (cf. Lazaridis et al. (2016), Extended Data Figure 5):

Global peak frequencies of the Ancient Ancestral South Indian component. This South Eurasian genome element attains its highest percentages among Asur individuals of southern Asia (100% maximum).

Genetic distance of the Ancient Ancestral South Indian component to modern populations. The Andamanese Jarawa and Onge groups have the closest affinity to this ancestral element.

Among Afro-Asiatic speakers, the Ancient Ancestral South Indian component reaches a frequency peak among EmiratiC individuals in the United Arab Emirates (10.9%). It is generally absent among other Afro-Asiatic-speaking populations.

A Jarawa boy (Andamanese). The Ancient Ancestral South Indian component peaks in frequency among the Jarawa and Onge Andamanese, as well as tribal groups and Dravidian speakers in southern Asia.

Odisha women (tribal Indians). After the Onge and Jarawa Andamanese, the Ancient Ancestral South Indian component reaches its next highest frequencies among the tribal populations and Dravidian speakers of the Indian peninsula.

A Dravidian man (Dravidian). The Ancient Ancestral South Indian component constitutes most of the ancestry borne by Dravidian speakers, a fact which is often physically evident.

A Sinhalese man. Although the Sinhalese of Sri Lanka today speak an Indo-European language, they predominantly carry the Ancient Ancestral South Indian component like Dravidian and tribal groups (albeit to a slightly lesser degree). This suggests that the Sinhalese originally spoke a Dravidian tongue and later adopted an Indo-European language.

A Brahmin man. Genome analysis indicates that, after Punjabi Jatts, Rors and Khatris, Brahmin individuals have the next lowest average frequency of the Ancient Ancestral South Indian component among modern populations in India (~50%).

A Punjabi Jatt man. The Punjabi Jatt are an Indo-European-speaking population of northwestern India and eastern Pakistan, who bear a slight predominance of West Eurasian ancestry (~60%). Most of this heritage consists of the Iran Neolithic and European Steppe genome components. The remainder of the Punjabi Jatt’s ancestry is largely composed of the Ancient Ancestral South Indian element (just under 40%). This is the lowest average frequency of this ancestral component among all modern groups in India.

Appendix

Genome analysis indicates that Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn region predominantly bear non-African ancestry (over 70% on average), which consists of a Levantine Natufian component and a Eurasiatic component (i.e. European Steppe and ancient East Asian elements). They also have minor Sub-Saharan African admixture (~25%) and trace Iberomaurusian admixture (under 5%).

The Universal Basic Genetic Model detects this same overall ancestral composition, albeit with less numerical precision. Below is a UBGM data table for Afro-Asiatic and Nilo-Saharan speakers in Northeast Africa, as well as UBGM data tables for each Horn population listed in the Global25 modern datasheet.

Universal Basic Genetic Model (rough ancestral composition):

Afro-Asiatic-speaking populations, Dinka and Mota on the Universal Basic Genetic Model.

Eritrean

Ethiopian Afar

Ethiopian Agaw

Ethiopian Amhara

Ethiopian Jew

Ethiopian Oromo

Ethiopian Tigray

Ethiopian Wolayta

Iraqw

Rendille

Kenya Somali

South Somali

For the exact frequencies of these various ancestral components, see the free form charts below, which cross-analyse the genomes of Cushitic, Nubian, Sudanese “Arab” (Arabized Nubian) and peninsular Arab individuals with those of over 7000 ancient samples from the Global25 ancient datasheet rather than just a handful of ancient samples. Also refer to Detecting the Eurasiatic genome component in Cushitic peoples and First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants.

Free-form genetic models (exact ancestral composition):

Genome analysis of an ancient Cushitic individual of the Pastoral Neolithic (Elmenteitan) using a free-form model. This person predominantly bears non-African ancestry (~74.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (25.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Afar individual using a free-form model. This person predominantly bears non-African ancestry (~81.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d element), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (Vietnam_LN.SG and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Agaw individual using a free-form model. This person predominantly bears non-African ancestry (~80.4%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Guam_Latte elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (19.6% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Amhara individual using a free-form model. This person predominantly bears non-African ancestry (~78.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d element), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (21.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Bataheen Nubian individual using a free-form model. This person predominantly bears non-African ancestry (~80.6%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian, Israel_IBA, Jordan_LBA and Syria_TellQarassa_Umayyad elements), a European Steppe component (Russia_North_Caucasus element), an Iran Neolithic component (Iran_C_TepeHissar element), a Dravidian component (India_RoopkundA element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (19.4% of the Kenya_Kakapel_LIA element).

Genome analysis of a Beni Amer Beja individual using a free-form model. This person predominantly bears non-African ancestry (~81.4%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an Iran Neolithic component (Iran_C_TepeHissar element), an ancient East Asian component (China_AmurRiver_N element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.6% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Gaalien Sudanese “Arab” individual using a free-form model. This person predominantly bears non-African ancestry (~71.6%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Skh002, Israel_Natufian and Turkey_Alakh_MLBA elements), a European Steppe component (Kyrgyzstan_TianShan_Saka.SG and Russia_North_Caucasus elements), an Iran Neolithic component (Turkmenistan_Gonur_BA_1 element), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (28.4% of the Kenya_Kakapel_LIA element).

Genome analysis of an Oromo individual using a free-form model. This person predominantly bears non-African ancestry (~82%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Israel_Natufian elements), a European Steppe component (Russia_North_Caucasus element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Saho individual using a free-form model. This person predominantly bears non-African ancestry (~83.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian, Egypt_ThirdIntermediatePeriod and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (16.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a South Somali individual using a free-form model. This person predominantly bears non-African ancestry (~81.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Haatimi Bravanese Somali individual using a free-form model. This person predominantly bears non-African ancestry (~81.6%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Jordan_PPNB elements), a European Steppe component (Russia_North_Caucasus element), a Caucasus Hunter-Gatherer component (Turkey_IA_o6 element), a Dravidian component (India_RoopkundA element), an ancient East Asian component (Vietnam_LN.SG and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he has minor Sub-Saharan African admixture (18.4% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Tigrayan individual using a free-form model. This person predominantly bears non-African ancestry (~81.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Kyrgyzstan_TianShan_Saka.SG elements), an Iran Neolithic component (Iran_C_TepeHissar element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Rashaida Arab individual. This person entirely bears non-African ancestry (100%). This non-African heritage consists of a Levantine component (Levant_Tell_Qarassa_Early_Antiquity, Levant_SYR_Tell_Masaikh and Levant_Natufian_EpiP elements) and an Iran Neolithic component (IRN_Ganj_Dareh_N element).

A Somali man and woman (Cushitic). Genome analysis indicates that the Cushitic-descended peoples of the Horn region bear a predominant non-African ancestry (over 70% on average). This non-African heritage consists of a Levantine Natufian component and a Eurasiatic component (European Steppe and ancient East Asian elements). Additionally, they have some minor Sub-Saharan African admixture (~25%). They also possess minute North African Iberomaurusian admixture (under 5%).

An Emirati Arab bedouin man (Semitic). Among Afro-Asiatic-speaking populations, EmiratiC bedouin Arabs have the most Ancient Ancestral South Indian admixture (10.9%). This is due to contact between their Semitic ancestors and natives of the Indian peninsula.

An Assyrian man (Semitic). Among Afro-Asiatic-speaking populations, Assyrians have the most Iran Neolithic admixture (17.3%). This is due to contact between their Semitic ancestors and peoples from South/Central Asia.

An Ashkenazi Jew man (Semitic). Among Afro-Asiatic-speaking populations, Ashkenazi Jews have the most Anatolian Neolithic admixture (32.8%). This is due to contact between their Semitic ancestors and native southern Europeans.

See also:

Genome analysis of modern Semitic individuals

Correcting a faulty genetic model #1

Detecting the Eurasiatic genome component in Cushitic peoples

Genetic affinities of the Middle Neolithic pastoralists of Skhirat, Morocco

First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants

Ashkenazi Jews and Sephardic Jews are of the same Levantine ancestral origin as Mizrahi Jews, Palestinians and other Semites

Y-DNA haplogroup E1b1a (formerly E3a) appears to have originated in the Middle East

Tutsi Bantus are of Nilotic origin

Tutsi Bantus are of Nilotic origin – Photographic adjunct

Baggara/Shuwa “Arabs”: last remaining descendants of the original Chadic speakers

Phenotype of the Canaanites, ancestors of modern Semitic peoples of the Levant and Mesopotamia

Genetic affinities of Nubians and Sudanese “Arabs” (Arabized Nubians)

Phenotypic affinities of ancient Cushitic peoples

Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa

A first look at Capsian ancient DNA and other updates

Myths about the Afro-Asiatic-speaking populations debunked

Genome analysis of the Tutsi Bantus

On Tutsi Bantus are of Nilotic origin, we saw how genetic and anthropological analyses have established that the Tutsi and Hima peoples of the Great Lakes region are basically Nilotes. As such, their ancestors appear to have switched from speaking Nilo-Saharan languages to their present adopted Bantu tongues. This biological affinity between the Tutsi-Hima and Nilotic groups like the Maasai is something which was also pointed out by Razib Khan on GNXP (see Tutsi are genetically very similar to the Maasai).

My earlier analysis of the genomes of Tutsi individuals was based on Global25 scaled coordinates since, at the time, that was the only kind of G25 data on the Tutsi-Hima that was available to me. Scaled coordinates are filtered versions of unscaled/raw/original coordinates. Because of this, results generated using scaled coordinates are less accurate than those generated using unscaled coordinates. Luckily, a reader has now pointed me to some Global25 unscaled coordinates belonging to Tutsi individuals.

Let us have a look these genomes.

Genome analysis

The samples in question comprise five Tutsi persons from the Democratic Republic of the Congo (where the local Tutsis are known as Banyamulenge). In order to determine these individuals’ overall ancestral makeup, I started off by performing a distal analysis.

Distal analysis

All of the examined Tutsis wound up deriving the core of their ancestry from the Late Iron Age Nilotes of Kakapel, Kenya. As explained on Guide to the Universal Basic Genetic Model, this Kenya_Kakapel_LIA genome element is the signature ancestral component of both the Bantu and Nilotic peoples of eastern Africa. It is today found in its most pristine form among the Luhya Bantus and Luo Nilotes. Additionally, the Tutsi individuals each bear low frequencies of the ancient East African Hunter-Gatherer component, a vestigial ancestral element associated with stone age foragers from Chencherere, Malawi.

The remainder of the sampled Tutsis’ ancestry consists of non-African components. Of these, primarily Levantine Natufian, European Steppe and Upper Paleolithic European elements can be discerned, components that are typical of Cushitic individuals. This suggests that the Tutsis, like their Nilotic Maasai kinsmen, acquired most of their non-African admixture through contacts with Cushitic people.

Genome analyses of Tutsis from the Democratic Republic of the Congo (distal). All of these individuals trace the core of their ancestry to the Late Iron Age Nilotes of Kakapel, Kenya. They also have minute ancient East African Hunter-Gatherer admixture (Malawi_Chenchere_LSA_5200BP element). Additionally, they bear significant non-African admixture, which mostly consists of Levantine Natufian, European Steppe and Upper Paleolithic European components.

Proximal analysis

Next, to find out which specific populations contributed to the Tutsi’s ancestral composition, I conducted a proximal genome analysis. Alongside the 7000+ official samples listed on the Global25 ancient datasheet, I included both of the unofficial G25 samples associated with the Middle Neolithic pastoralists of Skhirat, Morocco and the Early Neolithic pastoralists of Takarkori, Libya.

It turns out that the Tutsi samples trace the bulk of their recent ancestry to Nilote/Bantu samples — either to the Kenya_Historic_2 sample (which is most typical of the Dinka and other northern Nilotic pastoralists of the Nile Valley), or the Pastoral Iron Age sample (which is most typical of the Maasai and other southern Nilotic pastoralists of the Great Lakes), or the Kenya_Kakapel_LIA  sample (which is most typical of “purer” Nilotic/Bantu groups, such as the Luo Nilotes and Luhya Bantus). By contrast, only a minority of the examined Tutsis’ recent ancestry can be attributed to Cushitic peoples (i.e. less than 30% of Pastoral Neolithic and Kulubnarti/Sudan Early Christian elements).

Genome analyses of Tutsis from the Democratic Republic of Congo (proximal). All of these individuals trace the majority of their recent ancestry to Nilotic/Bantu samples — either to the Kenya_Historic_2 sample (most typical of the Dinka and other northern Nilotic pastoralists of the Nile Valley), or the Pastoral Iron Age sample (most typical of the Maasai and other southern Nilotic pastoralists of the Great Lakes), or the Kenya_Kakapel_LIA  sample (most typical of “purer” Nilotic/Bantu groups, such as the Luo Nilotes and Luhya Bantus). By contrast, only a minority of the examined Tutsis’ recent ancestry can be attributed to Cushitic peoples (under 30% of Pastoral Neolithic and Kulubnarti/Sudan Early Christian elements).

That said, one of the Banyamulenge samples stands out from the rest. Unlike the other Tutsis, who are clearly of Nilotic origin, this individual (identified as Peter below) appears to descend from assimilated Cushitic people. He traces just over half of his ancestry to the Cushites of the Pastoral Neolithic, with the remainder primarily consisting of Nilotic/Bantu-related admixture. This suggests that this particular individual’s family line, although now substantially intertwined with the Tutsi proper, may actually have South Cushitic (Iraqw) roots.¹

Genome analysis of a Tutsi individual from the Democratic Republic of Congo (proximal). This person traces just over 50% of his recent ancestry to the Cushites of the Pastoral Neolithic. By contrast, all of the other Tutsi samples above derive most of their recent ancestry from Nilotic populations. Hence, this individual likely descends from South Cushitic outsiders, who in the recent past were assimilated into the Tutsi culture group.

Genome analysis of an Iraqw individual from Tanzania. This person traces almost 70% of his ancestry to Cushitic-associated Pastoral Neolithic and Kulubnarti/Sudan Early Christian components (Kenya_PastoralN, Sudan_EarlyChristian_R and Tanzania_PN elements).

Conclusion

Distal (remote) analysis of Tutsi individuals in the Democratic Republic of the Congo (Banyamulenge) indicates that, like all Bantus and Nilotes in eastern Africa, they derive the core of their genome ancestry from the Late Iron Age peoples of Kakapel, Kenya. They also bear considerable non-African admixture, which was obtained via interaction with Cushitic peoples. Furthermore, proximal (recent) analysis reveals that most of this ancestral composition was specifically inherited from Nilotic populations of the Iron Age. However, one individual in the dataset traces instead just over 50% of his recent ancestry to the Cushites of the Pastoral Neolithic. Altogether, this confirms that the Tutsis as a population are of Nilotic origin, though they did receive some gene flow from neighboring communities. The Tutsi also historically assimilated outsiders into their culture group, including apparently a few individuals of recent South Cushitic origin.

Tutsis in the Democratic Republic of the Congo (Banyamulenge). Genome analysis indicates that Tutsi individuals derive most of their ancestry from Nilotic populations. This Nilotic affinity is also evident in the physiognomy of the average Tutsis, who bear a close resemblance to the Maasai and other Nilotes.

Tutsi leader Paul Kagame. Genome analysis has found that a minority of Tutsi individuals appear to be of foreign origin and were only recently assimilated into the Tutsi culture group. Although now admixed with the Tutsi proper, these individuals have a distinct physiognomy and also a bear higher than usual Cushitic ancestry.

An Iraqw woman (Cushitic). Historically, South Cushitic speakers occupied large parts of the Great Lakes region. They encountered and culturally influenced many neighboring Bantu and Nilotic populations, including the Tutsis’ ancestors.

Notes

¹It is also interesting to note that, in the present genome analysis, the proportion of what appears to be assimilated Cushitic individuals (1 person out of 5 or 20%) is roughly the same as the proportion of Tutsi individuals in Burundi that Trombetta et al. (2015) report carry the South Cushitic-affiliated haplogroup E1b1b-M293 (~22%; see Supplementary Table 7).

See also:

Tutsi Bantus are of Nilotic origin

Tutsi Bantus are of Nilotic origin – Photographic adjunct

Guide to the Universal Basic Genetic Model

Genetics of the Hausa, Chadic speakers of Niger-Congo origin

Baggara/Shuwa “Arabs”: last remaining descendants of the original Chadic speakers

A very Eurasiatic Toubou sample

First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants

A first look at Capsian ancient DNA and other updates

Hey there folks! After a brief hiatus, it is good to be back.

2024 was monumental for ancient DNA, anthropology in general, and this blog. Over the course of that year, we explored many topics related to the Afro-Asiatic-speaking populations and debunked a number of associated myths (some of them long-held!).

A Somali man (Cushitic).

A Saudi Arab man (Semitic).

A Beja man (Cushitic).

Discoveries and breakthroughs

Among other things, we learned that:

• The Dynastic Race Theory might be true.
• The Y-DNA haplogroup E1b1a (formerly E3a) appears to have originated in the Middle East.
• Cushitic-descended peoples predominantly bear non-African ancestry (over 70% on average). This non-African heritage generally consists of a Levantine Natufian component, a European Steppe component, and a newly-detected Upper Paleolithic European component. They also have minor Sub-Saharan African admixture (~25%) and trace North African Iberomaurusian admixture (under 5%). The non-African heritage which Cushitic individuals possess is/has often been underreported in analyses due to bad genetic modelling. Details on how to easily detect this missing non-African ancestry are provided on Detecting the Eurasiatic genome component in Cushitic peoples, Correcting a faulty genetic model #1 and Correcting a faulty genetic model #2.
• During the Bronze Age and earlier, the ancestral Semitic peoples of the Levant and Mesopotamia almost exclusively had dark skin, dark hair and dark eyes. That includes the ancient Canaanites of Israel/Palestine and the early Semitic societies of Jordan, Alalakh and Sidon.
• Ashkenazi Jews and Sephardic Jews are of the same Levantine ancestral origin as Mizrahi Jews, Palestinians and other Semites.
• The Mahra are among the “purest” Arabian peoples. As such, they are distinct from Afro-Arabs such as Al-Akhdam.
• The Northern Somalis from Puntland (Majerteen Darod) have the most non-African ancestry in the Horn.
• The ancient Egyptian Queen Tiye was of Eurasiatic origin.
• The Baggara Arabs (Shuwa Arabs) of the Sahel appear to be the last remaining descendants of the original Chadic speakers.
• Benadiri/Bravanese Somalis share most of their ancestry with Cushitic individuals of the northern Horn, confirming that they are of ethnic Somali origin. However, Benadiri/Bravanese Somalis also have secondary Persian, Indian and Bantu admixtures, which vary in frequency between individuals.
• Many Cushitic individuals carry European Steppe-associated mtDNA haplogroups.
• The Tuareg Berbers are genetically similar to other Berber populations of the Maghreb. This confirms that the Tuareg are ethnic Berbers.
• The genetic firms 23andme and AncestryDNA misrepresent the ancestral composition of Afro-Asiatic speakers.
• We can use the mAUTO formula to infer autosomal DNA from Y-DNA and mtDNA.
• Medieval DNA from Socotra confirms that the Socotri are among the “purest” descendants of the Levantine Natufians.
• DNA testing allegedly disproves Jewish origin of Igbo Jews.
• Although Abyssinians (Habesha) today speak Semitic languages, they are genetically, physically and culturally identical to Cushitic-speaking Agaws. This confirms that Abyssinians are Agaw people, who switched from speaking their original Cushitic Agaw language(s) and adopted Semitic languages.
• The Ancient Egyptian Pharaoh Ramesses III and Unknown Man E (Prince Pentawere) carried haplogroup E1b1b-V22.
• The Haplogroups E1b1b-V12 and E1b1b-V32 appear to be old Arabian lineages.
• Although the Hausa today speak an Afro-Asiatic language of the Chadic branch, they are genetically similar to Niger-Congo speakers. This suggests that the Hausa originally spoke a Niger-Congo tongue and later shifted to speaking their present Chadic idiom.
• The Tutsi Bantus are of Nilotic origin. Although the Tutsi of Rwanda and Burundi today speak a Bantu language, genome and haplogroup analyses indicate that they and Hima individuals of Uganda are closely related to the Maasai and other Nilotic peoples of the Great Lakes region. All of these populations appear to descend from the Nilotes of the Pastoral Iron Age, who bore some admixture from the earlier Cushites of the Pastoral Neolithic.
• The Cushitic peoples carry European-derived Neanderthal alleles.
• Craniometric analysis of northern Somalis and Egyptians has established that these Afro-Asiatic speakers share close affinities. By contrast, these individuals are morphologically distinct from the Sub-Saharan African samples (including the African American cohort).
• The Cushitic-speaking Beja of Eritrea, Sudan and Egypt share the same overall ancestral composition as other Cushitic individuals of the northern Horn. However, Beja individuals possess more elevated Levantine Natufian ancestry and lower European Steppe and Upper Paleolithic European genome elements compared to other Cushitic samples. The Beja also carry a minor Caucasus Hunter-Gatherer admixture signal.
• The Cushitic-speaking Saho of Eritrea share the same overall ancestral composition as other Cushitic individuals of the northern Horn. However, the Saho carry E1b1b-V22 at near fixed quantities (almost 100% frequency), a Y-DNA haplogroup that is closely associated with ancient and modern Egypt. Saho individuals also harbor at low frequencies a late period Egyptian genome signal. Altogether, this suggests that in antiquity, a small group of haplogroup E1b1b-V22 carriers from Egypt brought this paternal lineage to the Horn, where (probably through polygamy) it gradually attained a high frequency among the Saho.
• Middle Neolithic pastoralists of Skhirat, Morocco, who mostly bore Levantine Natufian ancestry, were immediately ancestral to the Cushites of the Pastoral Neolithic complex. Additionally, the Skhirat herders seem to have been largely responsible for the cattle pastoralism tradition of modern herder groups in the northern Maghreb as well as those of Cushitic, Egyptian, Sudanese “Arab,” Toubou and Baggara/Shuwa “Arab” pastoralists in Northeast Africa and the Chad Basin.
• Nubians and Sudanese “Arabs” (Arabized Nubians) have the same overall ancestral makeup, confirming their shared origins. They are also genetically similar to adjacent Cushitic-descended peoples, but have lower frequencies of the latter’s core Levantine Natufian, European Steppe and Upper Paleolithic European components. This is largely because Nubians and Sudanese “Arabs” received extra gene flow from both West Asian and Nilotic population sources.

We are now well into 2025. And already, we are greeted with an exciting new ancient DNA study from the Maghreb.

Capsian genetics

Lipson et al. (2025) have analysed some old genomes affiliated with the Capsian culture, a pre-Neolithic complex. These Capsian individuals seem to be of the same ancestral stock as the earlier, Epipaleolithic Iberomaurusians of Taforalt, Afalou Bou Rhummel and other sites in Northwest Africa.

Genome analysis

The earliest analysed Capsian specimens, two individuals from the Djebba site in Tunisia (dated c. 8000 BP), almost exclusively harbor Iberomaurusian-related ancestry. This ancestry is similar to that borne by an Epipaleolithic individual from Afalou Bou Rhummel, Algeria. However, a later Capsian specimen from the Hergla site in Tunisa (c. 5900 BP) shows evidence of outside contact; namely, with the Middle Neolithic pastoralists of Skhirat, Morocco and the Early Neolithic cultivators of Kahf Taht Al Ghar, also in Morocco. This is because the Hergla individual bears a Levantine Natufian genome element like that which typifies the Skhirat herders, as well as some Early European Farmer admixture (i.e. Anatolian Neolithic ancestry) akin to that found in the Kahf Taht Al Ghar agriculturalists.

Ancestral composition of Early Holocene Capsian individuals from the Djebba and Hergla sites, Epipaleolithic Iberomaurusian individuals from Afalou Bou Rhummel, Algeria (ABR), and Neolithic individuals from the Doukanet el Khoutifa site in Tunisia (DEK1 and DEK2). The two analysed Capsian individuals from Djebba almost exclusively bear Iberomaurusian-related ancestry (blue component), with minute admixture associated with the West Hunter-Gatherers of Mesolithic Europe (purple component). A later Capsian specimen from Hergla also carries Iberomaurusian ancestry at high frequencies. However, in addition to this ancestry, the Hergla individual possesses some Levantine Natufian admixture (grey component) and Early European Farmer/Anatolian Neolithic admixture (orange component). This suggests that later Capsians interacted with the Levantine Natufian ancestors of the Skhirat pastoralists and the Iberian ancestors of the Kahf Taht Al Ghar cultivators (Lipson et al. (2025)).

Y-DNA

Additionally, Lipson et al. typed their Capsian samples for uniparental markers. The scientists report that all of the examined specimens bore Y-DNA and mtDNA haplogroups that are common among modern Maghrebi individuals, consistent with the genome affinities they observed.

The one Djebba sample whose Y-DNA they assessed turned out to belong to E1b1b1a1, better known as the M78 subclade of haplogroup E1b1b. In the Maghreb, M81 (referred to as the “Berber marker” because of its ubiquity among Berber groups) is today the dominant E1b1b sublineage. However, the M78 subhaplogroup also occurs at significant frequencies, particularly among local Arabic speakers (Arabized Berbers). For his part, the Capsian individual from Hergla was assigned to the T1a1a lineage. Simões et al. (2023) note that both of the ancient Skhirat pastoralists that they analysed for Y-DNA markers also carry this paternal haplogroup. This further confirms that the Capsians of Hergla received their Levantine Natufian admixture via interactions with the Skhirat herdsmen.¹

A Moroccan “Arab” man (Berber origin). Most Arabic speakers in Morocco have been observed to belong to the Y-DNA haplogroup E1b1b (72.7%-75.5% of individuals). Of these Moroccan “Arab” samples, almost all carry the M78 (11.4%-42.9% of individuals) and M81 (32.6%-52.3% of individuals) subclades of E1b1b (cf. Semino et al. (2004)). This points to genetic continuity between Moroccan “Arabs” and their Capsian ancestors because the ancient Capsians primarily bore the E1b1b-M78 subhaplogroup.

mtDNA

In terms of mtDNA, both of the examined Capsian individuals from Djebba were found to bear derivatives of the U6 haplogroup. This maternal lineage is typical of Iberomaurusian specimens and is still widespread in the Maghreb. The later Capsian individual from Hergla belongs instead to the R0a clade. R0a is a mitochondrial haplogroup that is today frequent in the Arabian peninsula, Levant and Horn of Africa — again, consistent with gene flow from the Skhirat pastoralists.

Lipson et al. summarize their findings thusly:

Uniparental markers are consistent with our genome-wide results in
indicating a majority of Maghrebi ancestry among the newly reported
individuals, with more admixture from other sources later in the
transect. Of the five individuals assigned male at birth, four could be assigned to Y-chromosome haplogroup E1b1b1a1, which is characteristic of northern Africa, particularly in ancient individuals with Maghrebi ancestry. The exception was I22852 (Hergla), who carried T1a1a, associated with Levantine farmers. For mtDNA, the individuals from ABR and Djebba, as well as both individuals from the DEK1 subgroup and one from DEK2, carried subclades of U6, also known primarily from ancient northern Africa. Haplogroup L3f1b + 16292 (I22867, DEK2) belongs to a clade hypothesized to have originated in eastern Africa and spread to other parts of the continent, while R0a2 (I22852, Hergla) has a wide distribution, but has also been observed in the Neolithic Levant. Finally, individual I22866 (DEK2) carried mtDNA haplogroup U5b2b1, which is characteristic of pre-Neolithic Europe, and is probably derived from European hunter–gatherers, either directly (hunter–gatherers crossing the Strait of Sicily) or by means of WHG ancestry in European farmers.

Y-DNA and mtDNA haplogroups carried by the ancient Capsians of Djebba and Hergla. The Capsian individual at Djebba whose paternal markers were analysed belongs to E1b1b1a1, better known as the M78 subclade of haplogroup E1b1b. This lineage was common among the Epipaleolithic Iberomaurusians and is still found at elevated frequencies in the Maghreb. The later Capsian specimen from Hergla bears instead the T1a1a clade, a haplogroup ubiquitous among the Middle Neolithic pastoralists of Skhirat, Morocco. In terms of mtDNA, both of the analysed Djebba specimens carry the U6 lineage. This maternal haplogroup is typical of the Iberomaurusians and remains common today in the Maghreb. For his part, the Hergla individual belongs to the R0a clade, a mitochondrial lineage that is widespread in the Arabian peninsula and Levant. This again confirms that the later Capsians of Hergla received gene flow from the Skhirat herders, who bore Levantine Natufian ancestry (Lipson et al. (2025)).

Craniometric affinities

Furthermore, Lipson et al. (2025)’s observations concur with craniometric analysis by Sereno et al. (2008). The latter researchers report that their Capsian and Iberomaurusian skeletal samples are closely related. Both of these specimens are also distinct from the Paleolithic Aterian samples, the earliest skeletons which the scientists analysed (c. 40,000 years old).

Craniometric analysis of Iberomaurusians (WMI, EMI), Capsians (EMC, WMC), “Mechtoids” from Mali and Mauritania (Mali, Maur), early Holocene Kiffians from Gobero (Gob-e), mid-Holocene Tenerians from Gobero (Gob-m), and ancient Aterians (Ater). The Iberomaurusian, Capsian, Mechtoid and Kiffian samples cluster together, showing Upper Paleolithic European ties. The Tenerian cohort sits apart, likely because of its “Mediterranean” affinities. The Aterian specimens, which are the oldest at over 40,000 years of age, are also an outlier (Sereno et al. (2008)).

Notes:

¹The Skhirat pastoralists appear, in turn, to have acquired their Iberomaurusian admixture from the Capsians. Such contacts between the Capsians and the Skhirat shepherds would explain why the lithic industry of the Cushites of the Pastoral Neolithic, who descend from the Skhirat herdsmen, shows affinities with that of the Capsians. Correspondingly, the Pastoral Neolithic complex of the Great Lakes region (which consists of the Savanna Pastoral Neolithic and Elmenteitan cultures) was formerly known as the Kenya Capsian, in acknowledgement of the archaeological ties that exist between it and the Capsian industry of Northwest Africa (cf. Cole (1954)).

See also:

Guide to the Universal Basic Genetic Model

First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants

Genetic affinities of Nubians and Sudanese “Arabs” (Arabized Nubians)

Today, we shall examine the genetic affinities of Nubians and Arabized Nubians (better known as Sudanese “Arabs”).

But just who are Nubians? They are a population of the Nile Valley, living in an area adjacent to Egyptians.

Nubians currently speak a Nilo-Saharan language called Nobiin. They appear to have adopted this tongue some time around the Iron Age. This is the period marking the rise of the X-Group (Post-Meroitic) culture, which succeeded the Meroitic civilization.

Archaeogenetic data indicates that the X-Group peoples were of the same ancestral stock as the preceding Meroites, with a slightly greater Sub-Saharan African admixture (cf. Cherifi and Amrani (2020)). At present, the Meroitic language remains undeciphered. Linguistic evidence, however, suggests that it may have been an Afro-Asiatic language (Kirsty Rowan (2006)).

Nubian women (Cushitic origin).

“Red Nubians” vs. “Black Nubians”

We know from ancient inscriptions, mural artwork and skeletal analysis that there were at least two distinct population groups inhabiting ancient Nubia: the “Red Nubians” and the “Black Nubians.” These peoples were of different ancestral origins, much like how European Australians and Aboriginal Australians (though both Australians) are ultimately of different ancestral origins.

“Red Nubians” were the Egyptian-related inhabitants of ancient Nubia. They include populations like the Meroites and Post-Meroitic/X-Group makers. Their modern descendants include most Nubians, who live in the area between central Sudan and southern Egypt. Nubians are physically, genetically and culturally similar to Afro-Asiatic-speaking populations.

“Black Nubians” were the Nilotic-related inhabitants of ancient Nubia. Their modern descendants include groups like the Nuba (also spelled Noba), who live in the Nuba Mountains (Kordofan Mountains or Nuba Hills) further south. Nuba are physically, genetically and culturally similar to Niger-Congo/Nilo-Saharan-speaking populations.

Inscriptions

Epigraphs belonging to the Axumite King Ezana explicitly differentiate between “Red Nubians” and “Black Nubians.” The “Red Nubians” are Cushitic/Egyptian-related peoples and the “Black Nubians” are “Negroes” associated with Nilotes (Asiatic Society (1968)):

The Nubians at that time also (as in previous centuries) were divided into the Red People or the Kasu or Cushites (Hamites like the Egyptians) and the Black People or Sudanian Negroids[…] these two peoples evidently were contrasted as “Red” and “Black” from their skin-colour; and as late as the fourth century A.D., the great Ethiopian King ‘Ezana, who conquered Nubia, differentiated between these two classes of Nubian.

The medieval chronicle of the Ethiopian Emperor Susenyos I further confirms this identification, as the royal court historian refers to the Afro-Asiatic-speaking non-slave populations as qayh (red) and the “shanqilla” or “Negro” groups as tsalim (black) (cf. Pankhurst (1976)).

Wall art

The Grand Procession is a mural in Thebes, Egypt, located within the tomb of Rekhmire, a vizier under Pharaoh Thutmose III. The wall painting contains several registers or levels, which depict various foreign envoys submitting tribute to the ancient Egyptians. These figures include Nubian, Puntite and Cretan emissaries.

The Grand Procession’s bottom register shows several reddish-skinned, orthognathous Nubian delegates, who strongly resemble the mural’s similarly copper-toned Egyptian, Puntite and Cretan figures. Standing amongst these “red” Nubians is a jet black-skinned, prognathous man, who is evidently of Nilotic origin. As the Egyptologist Kenneth Kitchen puts it, “beside the so called “Hamitic’ type (like Parahu [a Puntite chief]) not very different from Egyptians in appearance, others represented were clearly of Negro stock” (cf. Balanda (2005)).

The Grand Procession is paralleled by the Book of Gates mural in the tomb of Pharaoh Seti I, which similarly contrasts Egyptian-like figures with Nilotic “Negroes” (Richardson (2003)).

Foreign tributaries on the top three registers of the Grand Procession mural at Thebes, Egypt. Top row=Puntites (left), Nilotes (far right); Middle row=Cretans; Bottom row=Egyptian-like “Red Nubians,” Nilotic-like “Black Nubian” (center-right).

A Nilote woman retaining the physiognomy of the “Black Nubians,” the Nilotic inhabitants of ancient Nubia.

Skeletal analysis

In his capacity as chief anatomist of the Archaeological Survey of Nubia, Grafton Elliot Smith had the opportunity to forensically examine thousands of ancient Egyptian and Nubian skeletal remains — exponentially more than any other Egyptologist. He reported the existence of three different types of crania, which were excavated in Nubia: an Egyptian “Caucasoid” type, a non-Egyptian “Caucasoid” type, and a “Negroid” type.

Three different types of crania excavated in Nubia by Grafton Elliot Smith and other Egyptologists accompanying the Archaeological Survey of Nubia. Left=Skull of an individual of unknown origin; Center=Skull of an ancient Egyptian individual; Right=Skull of a “Negro” individual. This is consistent with archaeogenetic evidence (discussed below) indicating that Egyptian-related peoples inhabited areas of Nubia (Elliot Smith and Jones (1910)).

Brace (1993) likewise observed that his modern Nubian sample was craniometrically most similar to neighboring Afro-Asiatic speakers and distinct from both modern Niger-Congo speakers and the Mesolithic sample from Wadi Halfa, Sudan. Because of their “Negroid” affinities, the Mesolithic skeletons from Wadi Halfa and Khartoum are thought to belong instead to the ancestors of current-day Nilotes.

Analogously, Kemp (2006) found that his ancient and modern Egyptian samples were craniometrically closest to other Afro-Asiatic speakers and ancient & medieval Nubians inhabiting Northeast Africa. They were also more distantly related to populations in the Near East, but again shared no significant affinities with the ancient and modern “Negroid” populations in Africa.

Craniometric analysis of ancient/medieval Nubians and various other global samples. The Nubian specimens cluster with Afro-Asiatic speakers of Northeast Africa (ancient/modern Egyptians and modern Cushitic peoples from the Horn). All of these samples are distinct from the modern and archaic “Negroid” samples (Kemp (2006)).

Genetics

Y-DNA

Hassan et al. (2008) analysed various Afro-Asiatic and Nilo-Saharan-speaking populations of Sudan for Y-DNA. The scientists report that haplogroup J was the most common lineage among both their Nubian (17 out of 39 examined individuals or ~44%) and Sudanese “Arab” samples (16 out of 24 Arakien individuals or ~67%, 12 out of 28 Messeria individuals or ~43%, and 20 out of 50 Gaalien individuals or 40%). Furthermore, they observed that most of their tested Nubian and Sudanese “Arab” individuals specifically bore the J1 subclade, which is common among peninsular Arabs and Jews. This suggests past interactions between the Nubian peoples and Semitic speakers of the Middle East.¹

The remaining Nubian samples in Hassan et al.’s dataset primarily carried the haplogroup E1b1b (9 out of 39 individuals or ~23%). Of these E1b1b carriers, most belonged to the V12* subclade. E1b1b is a signature paternal lineage of Afro-Asiatic speakers, especially those in Africa. Hassan et al. (2009) observed that 3 out of 4 Meroitic samples (75%) were positive for the YAP insertion, which is ancestral to the E1b1b clade. This supports a connection between present-day Nubians and the ancient Meroites. Nubians also show ties with Egyptians since the V12 subclade of E1b1b is today the most common Y-DNA clade among males in southern Egypt (~74.5%; see Supplementary Table 7 in Trombetta et al. (2015)). E1b1b in general is also the modal haplogroup among males in northern Egypt, as it is among the Berber and Maghrebi “Arab” (Arabized Berber) populations in Northwest Africa.

The rest of the examined Nubian individuals belonged to the R1b haplogroup (4 out of 39 individuals or ~10%), F haplogroup (4 out of 39 individuals or ~10%), and I haplogroup (2 out of 39 individuals or ~5%). These lineages are, respectively, mainly linked with the spread of Indo-European speakers, peoples bearing Anatolian Neolithic ancestry, and peoples bearing Western Hunter-Gatherer ancestry. Some of the Nubian samples also carried the archaic African haplogroup A (~8%), which is frequent amongst northern Nilotic populations.

For their part, the rest of the Sudanese “Arab” samples also primarily bore the E1b1b subclade (9 out of 50 Gaalien individuals or 18% and 4 out of 24 Arakien individuals or ~17%). Most fell under either the V12 subclade or its V32 derivative. However, in the case of the Messeria samples, haplogroup R1b was instead their second most common paternal lineage (7 out of 28 examined individuals or 25%).

A Nubian boy (Cushitic origin).

mtDNA

In terms of mtDNA, Hassan (2009) indicates that his Nubian and Sudanese “Arab” samples predominantly belonged to the Africa-centered macrohaplogroup L. 83% of the tested Nubians carried various L subclades, with only 17% bearing derivatives of the non-African M and N macroclades. Similarly, 94% of Messeria, 91% of Gaalien, and 82% of Arakien Sudanese “Arab” samples belonged to L subhaplogroups.

Among the Arakien in particular, this haplogroup pattern is primarily due to marked admixture with local Nilotic females. This is clear from their high frequency of L2d1 (52.9%), a Sub-Saharan African mtDNA lineage which, in Hassan (2009)’s dataset, is also common among the Nuer Nilotes (37.5%).

That said, other mitochondrial studies have observed high frequencies of non-African M and N derivatives among their Nubian samples (48%; cf. Table 3-3 in Non (2010)). This suggests that, in the case of Nubians, the elevated percentages of macrohaplogroup L which Hassan (2009) observed may be a product of limited sampling.

mtDNA affinities of Afro-Asiatic, Niger-Congo and Nilo-Saharan-speaking populations in eastern Africa. Many Nubian individuals from Sudan carry derivatives of the non-African M and N mtDNA haplogroups like adjacent Afro-Asiatic speakers: Tigray=63.6%; Somali=60%; Afar=56.2%; Gurage=55%; Oromo=53.3%; Sudanese Nubian=48%; Amhara=46%. This supports the suggestion that Nubians originally spoke an Afro-Asiatic language and only later adopted a Nilo-Saharan tongue (Non (2010)).

For a nuanced discussion on the haplogroup L3, which Hassan (2009) detected amongst his Nubian and Sudanese “Arab” samples, see Genetic affinities of the Cushitic-speaking Beja.

A Sudanese “Arab” woman (Nubian origin).

Genome analysis

To gain an understanding of the ancestral composition of Nubians and Sudanese “Arabs” (Arabized Nubians), I performed a genome analysis on several official and unofficial Global25 samples (see Appendix below). For Nubians proper, this consisted of Mahas, Halfawieen, Danagla and Bataheen individuals from Sudan. For Sudanese “Arabs,” this consisted of Shaigia, Messeria, Kababish and Gaalien individuals. Except for the Kababish samples (whose coordinates are listed on the official Global25 modern datasheet), all of these Nubian and Sudanese “Arab” samples were originally published in Fortes-Lima et al. (2022) and later converted to Global25 coordinates (see here for those unscaled or raw G25 coordinates).

The Nubian and Sudanese “Arab” persons turned out to be genetically similar to local Afro-Asiatic speakers. As with the Cushitic Beja and most other Cushitic, Ethiosemitic and North Omotic-speaking individuals of the Horn region, the Nubian peoples predominantly bear non-African ancestry (over 70% on average). This non-African heritage generally consists of a Levantine Natufian component, a European Steppe component, and an Upper Paleolithic European component. They also have minor Sub-Saharan African admixture (~25%), and trace North African Iberomaurusian admixture (under 5%).

One characteristic difference between the Nubian/Sudanese “Arab” samples and neighboring Afro-Asiatic speakers is that the former bear moderate Caucasus Hunter-Gatherer and Iran Neolithic admixtures, as well as trace Dravidian admixture. The Caucasus Hunter-Gatherer element is clearly due to contact with Y-DNA haplogroup J carriers since this clade originated in the Caucasus and was introduced to the Semitic gene pool relatively recently, during the Bronze Age (see Notes below).

Another noticeable difference is that the Sub-Saharan African admixture of the Nubian and Sudanese “Arab” samples, although comparable in average frequency to that borne by Afro-Asiatic speakers from the Horn, exclusively consists of the ancient Nilo-Saharan component. This agrees with the high frequency of Nilotic-associated mtDNA L lineages among Nubians and especially Sudanese “Arabs.”

To determine which specific population sources contributed the ancestral components above, I conducted a followup proximal genome analysis (see here and here). As predicted, almost all of the Nubian and Sudanese “Arab” individuals wound up tracing much of their recent ancestry to the medieval Nubians of Kulubnarti, Sudan. These Christian period inhabitants of Nubia are genetically closely related to the Cushites of the Pastoral Neolithic, thus confirming the Cushitic affinities of most Nubians and Sudanese “Arabs.”

Moreover, genetic distance analysis of the Nubian and Sudanese “Arab” samples indicates that they share closest affinity with each other, as well as with Cushitic-descended individuals of the Horn and Nile Valley. This, again, is due to the fact that Nubians and Sudanese “Arabs” have the same overall ancestral makeup, bearing the same core ancestral components at similar frequencies as Cushitic peoples (see Appendix below).

A Sudanese “Arab” man (Nubian origin).

Conclusion

Nubians and Sudanese “Arabs” have the same overall ancestral makeup, confirming their shared origins. They are also genetically similar to adjacent Cushitic-descended peoples, but have lower frequencies of the latter’s core Levantine Natufian, European Steppe and Upper Paleolithic European components. This is largely because Nubians and Sudanese “Arabs” received extra gene flow from both West Asian and Nilotic population sources.

Appendix

Genome analysis of a Bataheen Nubian individual using a free-form model. This person predominantly bears non-African ancestry (~80.6%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian, Israel_IBA, Jordan_LBA and Syria_TellQarassa_Umayyad elements), a European Steppe component (Russia_North_Caucasus element), an Iran Neolithic component (Iran_C_TepeHissar element), a Dravidian component (India_RoopkundA element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (19.4% of the Kenya_Kakapel_LIA element).

Genome analysis of a Danagla Nubian individual using a free-form model. This person predominantly bears non-African ancestry (72%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), a Caucasus Hunter-Gatherer element (Turkey_IA_o6 element), an Iran Neolithic component (Turkmenistan_Gonur_BA_1 element), an ancient East Asian component (China_AmurRiver_N element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (28% of the Kenya_Kakapel_LIA element).

Genome analysis of a Gaalien Sudanese “Arab” individual using a free-form model. This person predominantly bears non-African ancestry (~71.6%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Skh002, Israel_Natufian and Turkey_Alakh_MLBA elements), a European Steppe component (Kyrgyzstan_TianShan_Saka.SG and Russia_North_Caucasus elements), an Iran Neolithic component (Turkmenistan_Gonur_BA_1 element), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (28.4% of the Kenya_Kakapel_LIA element).

Genome analysis of a Halfawieen Nubian individual using a free-form model. This person predominantly bears non-African ancestry (~75.4%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Skh002 and Jordan_PPNB elements), a European Steppe component (Kyrgyzstan_TianShan_Saka.SG and Russia_North_Caucasus elements), a Caucasus Hunter-Gatherer element (Turkey_IA_o6 element), an Iran Neolithic component (Iran_C_TepeHissar element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (24.6% of the Kenya_Kakapel_LIA element).

Genome analysis of a Kababish Sudanese “Arab” individual using a free-form model. This person predominantly bears non-African ancestry (~79.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Lebanon_IA3_o2.SG, Syria_TellQarassa_Umayyad, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), an Iran Neolithic component (Turkmenistan_Gonur_BA_1 element), a Dravidian component (India_RoopkundA element), an ancient East Asian component (Guam_LateUnai_noUDG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (20.2% of the Kenya_Kakapel_LIA element).

Genome analysis of a Mahas Nubian individual using a free-form model. This person predominantly bears non-African ancestry (~74.4%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Kyrgyzstan_TianShan_Saka.SG and Russia_North_Caucasus elements), an Iran Neolithic component (Turkmenistan_Gonur_BA_1 element), a Caucasus Hunter-Gatherer element (Turkey_IA_o6 and Turkey_MBA elements), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (25.6% of the Kenya_Kakapel_LIA element).

Genome analysis of a Messiria Sudanese “Arab” individual using a free-form model. This person bears significant non-African ancestry (~47.8%). This non-African heritage consists of a Levantine component (Skh002 and Israel_Natufian_d elements), a European Steppe component (Russia_North_Caucasus, Scotland_Orkney_MBA.SG and Kyrgyzstan_TianShan_Saka.SG elements), an Anatolian Neolithic component (Ktg001 element), a Caucasus Hunter-Gatherer element (Turkey_IA_o6 element), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has substantial Sub-Saharan African admixture (52.2% of the Kenya_Kakapel_LIA element).

Genome analysis of a Shaigia Sudanese “Arab” individual using a free-form model. This person predominantly bears non-African ancestry (~71.6%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Kyrgyzstan_TianShan_Saka.SG and Russia_North_Caucasus elements), an Iran Neolithic component (Turkmenistan_Gonur_BA_1 element), a Caucasus Hunter-Gatherer element (Turkey_IA_o6 element), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (28.4% of the Kenya_Kakapel_LIA element).

Genome analysis of a Beni Amer Beja individual using a free-form model. This person predominantly bears non-African ancestry (~81.4%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an Iran Neolithic component (Iran_C_TepeHissar element), an ancient East Asian component (China_AmurRiver_N element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.6% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a South Somali individual using a free-form model. This person predominantly bears non-African ancestry (~81.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Haatimi Bravanese Somali individual using a free-form model. This person predominantly bears non-African ancestry (~81.6%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Jordan_PPNB elements), a European Steppe component (Russia_North_Caucasus element), a Caucasus Hunter-Gatherer component (Turkey_IA_o6 element), a Dravidian component (India_RoopkundA element), an ancient East Asian component (Vietnam_LN.SG and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he has minor Sub-Saharan African admixture (18.4% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Oromo individual using a free-form model. This person predominantly bears non-African ancestry (~82%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Israel_Natufian elements), a European Steppe component (Russia_North_Caucasus element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Amhara individual using a free-form model. This person predominantly bears non-African ancestry (~78.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d element), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (21.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Tigrayan individual using a free-form model. This person predominantly bears non-African ancestry (~81.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Kyrgyzstan_TianShan_Saka.SG elements), an Iran Neolithic component (Iran_C_TepeHissar element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genetic distance analysis of a Kababish Sudanese “Arab” individual. This person shares closest affinity with Nubians and Cushitic-descended individuals of Northeast Africa. This is due to the fact that these individuals share the same overall ancestral makeup, bearing the same core genome components at similar frequencies.

Genetic distance analysis of a Mahas Nubian individual. This person shares closest affinity with Sudanese “Arabs” and Cushitic-descended individuals of Northeast Africa. This is due to the fact that these individuals share the same overall ancestral makeup, bearing the same core genome components at similar frequencies.

Genetic distance analysis of a South Somali individual. This person shares closest affinity with other Cushitic peoples of the Horn as well as Sudanese “Arabs” and Nubians of Northeast Africa. This is due to the fact that these individuals share the same overall ancestral makeup, bearing the same core genome components at similar frequencies.

Genetic distance analysis of an Amhara individual. This person shares closest affinity with other Cushitic peoples of the Horn as well as Sudanese “Arabs” and Nubians of Northeast Africa. This is due to the fact that these individuals share the same overall ancestral makeup, bearing the same core genome components at similar frequencies.

Notes

¹Ancient DNA analysis has found that modern Semitic peoples of the Levant genetically cluster with Bronze Age Levantines, who in turn descend from the Pre-Pottery Neolithic makers of the Levant, the immediate descendants of the Natufians. Moreover, the Semites’ Pre-Pottery Neolithic and Mesolithic Natufian ancestors did not carry the haplogroup J and instead primarily bore the E1b1b lineage. This indicates that the ancestral Semites acquired the J clade relatively recently, during the Bronze Age, through interaction with peoples from the Iranian plateau/Caucasus. On this genetic contact, Haber et al. (2017) write: “We compiled frequencies of Y-chromosome haplogroups in this geographical area and their changes over time in a dataset of ancient and modern Levantine populations (Figure S12), and note, similarly to Lazaridis et al., that haplogroup J was absent in all Natufian and Neolithic Levant male individuals examined thus far, but emerged during the Bronze Age in Lebanon and Jordan along with ancestry related to Iran_ChL.”

See also:

Haplogroups E1b1b-V2 and E1b1b-V32 appear to be old Arabian lineages

Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa

Genetic affinities of the Cushitic-speaking Beja

Genetic affinities of the Cushitic-speaking Saho

Genetic analysis of the Cushitic-speaking Agaws (Agaw proper) and Semitic-speaking Agaws (Abyssinians/Habesha)

Genetic affinities of North Omotic-speaking populations

Detecting the Eurasiatic genome component in Cushitic peoples

European Steppe-associated mtDNA haplogroups among the Cushitic peoples

A forgotten ancient DNA study on the Meroites

Genetic affinities of the Middle Neolithic pastoralists of Skhirat, Morocco

Lactase persistence (lactose absorption) rates among Egyptians, Ashkenazi Jews, Iraqi Jews and Moroccan Jews

On Genetic affinities of the Middle Neolithic pastoralists of Skhirat, Morocco and First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants, we saw how many Egyptian, Cushitic, Sudanese “Arab,” Baggara/Shuwa “Arab,” Toubou and northern Maghrebi pastoralists bear Levantine Natufian-related ancestry, much of which they inherited from Middle Neolithic herders of Skhirat, Morocco. The ancient Skhirat culture bearers also appear to be largely responsible for these groups’ cattle pastoralism tradition.

However, most Egyptians are traditionally fellahin (farmers) and carry little, if any, Skhirat-derived ancestry. What are the lactase persistence rates of these individuals, considering their agricultural background?

A recent study examined this issue in a global context.

Low LP rates among Egyptians and Jews

Bayless et al. (2017) compared the lactose persistence (LP) rates of the general Egyptian population with those of Ashkenazi Jews, Iraqi Jews, Moroccan Jews and various European, Sub-Saharan African, Asian, Amerindian and Oceanian populations.

The researchers indicate that most of the examined Egyptian individuals (73%) were lactose malabsorbers (i.e. lactase non-persistent). For the Jewish individuals, even lower rates of lactase persistence were reported: Iraqi Jews=93% lactose malabsorbers (7% LP), Ashkenazi Jews=83% lactose malabsorbers (17% LP), and Moroccan Jews=82% lactose malabsorbers (18% LP).

This was in sharp contrast to the European groups, most of whom had high rates of lactase persistence (especially northern Europeans; as few as 10% of Swedes, 12% of Danes and 14% of Irish were lactose malabsorbers). Oppositely, virtually all East Asian, Bantu & Yoruba, Amerindian and Aboriginal Australian samples were lactose malabsorbers.

Global prevalence of lactose malabsorption/lactase non-persistence. Most of the Egyptian, Jewish, Sub-Saharan African, East Asian, Amerindian and Oceanian samples consist of lactose malabsorbers/lactase non-persistent individuals, due to these populations’ respective agrarian or hunter-gatherer backgrounds. Conversely, most Europeans are lactose absorbers/lactase persistent (Bayless et al. (2017)).

Discussion

The low lactase persistence rate of the general Egyptian population is not surprising given the cultivator background of most natives of Egypt. This agrarian tradition also seems to be of longstanding.

Archaeogenetic analysis has established that ancient Egyptians of the Old Kingdom almost entirely bore Levantine Natufian ancestry (for details, see The Dynastic Race Theory might be true). The Mesolithic Natufians were hunter-gatherers who practised a rudimentary form of agriculture. As such, they were not milk drinkers and did not carry any lactase persistence/lactose tolerance mutations.¹ Ancient Egyptians from the later Old Kingdom-Middle Kingdom and Middle Kingdom periods received extra gene flow, mainly from outsiders bearing Caucasus Hunter-Gatherer ancestry. Modern Egyptians, both Coptic and Muslim, directly descend from the ancient Egyptians of the Old Kingdom-Middle Kingdom. This explains their low rates of lactase persistence.

Like most northern Semites, Ashkenazi Jews, Iraqi Jews and Moroccan Jews carry a mixture of Levantine Natufian ancestry — the original ancestral component of the Semitic peoples — and Caucasus Hunter-Gatherer, Iran Neolithic and Anatolian Neolithic elements. Anatolian Neolithic notwithstanding (an ancestral element which today occurs at low frequencies in many Semitic populations), all of these components are exclusively affiliated with forager and cultivator lifestyles. Ergo, the low lactase persistence rates of the Jewish groups are again to be expected.

Conversely, among the European samples, the high rates of lactase persistence have been attributed to recent positive selection for the T-13910 allele, a Europe-specific LP mutation (see Gerbault (2013)). Among the East Asian, Bantu & Yoruba, Amerindian and Aboriginal Australian samples, the low-to-zero rates of lactase persistence are clearly due to these populations’ traditionally farmer and hunter-gatherer subsistence modes.

An Egyptian man (Egyptian). Most Egyptian individuals are lactase non-persisent (73%), due to their traditionally farmer (fellahin) background.

An Ashkenazi Jew man (Semitic). Ashkenazi Jews and other Jewish individuals are mostly lactase non-persisent (83%), due to their traditionally agrarian background.

A Danish man (Germanic). Most Danes (88%) and other Europeans are lactase persistent due to positive selection for the T-13910 allele, a Europe-specific LP mutation.

Notes

¹Lactase is an enzyme that helps break down lactose (milk sugar). Humans naturally produce lactase until they are past the breast-feeding age. Thereafter, most people become lactase non-persistent unless they carry specific LP mutations, which allow their body to continue producing the enzyme.

See also:

Genetic affinities of the Middle Neolithic pastoralists of Skhirat, Morocco

First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants

The Dynastic Race Theory might be true

Ashkenazi Jews and Sephardic Jews are of the same Levantine ancestral origin as Mizrahi Jews, Palestinians and other Semites

Genetic affinities of the Middle Neolithic pastoralists of Skhirat, Morocco

On First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants, we saw how multiple population movements from outside Africa introduced pastoralism to the continent (both cattle pastoralism and camel pastoralism). One of the most important of these ancient migrations involved cattle herders from the Middle East, who settled at the Skhirat site in northern Morocco during the Middle Neolithic.

Archaeogenetic analysis indicates that the Skhirat peoples bore the Y-DNA haplogroup T, the mtDNA haplogroup M1, and Levantine Natufian genome ancestry. They appear to have been largely responsible for the cattle pastoralism tradition of modern herder groups in the northern Maghreb as well as those of Cushitic, Egyptian, Sudanese “Arab,” Toubou and Baggara/Shuwa “Arab” pastoralists in Northeast Africa and the Chad Basin.

An Ababda Beja man (Cushitic). Camel pastoralism seems to have been introduced quite recently to Northeast Africa, apparently in association with the spread of the Arabian E1b1b-V12 and E1b1b-V32 lineages. Archaeogenetics, excavations and ancient rock art suggest that prior to that, Cushitic herders raised cattle, having inherited their cattle pastoralism tradition from Middle Neolithic Skhirat ancestors.

Y-DNA and mtDNA

In 2023, scientists from Uppsala University analysed the DNA of ancient individuals excavated in the Maghreb, including one Epipaleolithic specimen, one Early Neolithic specimen from the Ifri n’Amr or Moussa site, four Early Neolithic specimens from the Kahf Taht Al Ghar site, and three Middle Neolithic specimens from the Skhirat site (cf. Simões (2023a)). Their findings are of particular interest vis-a-vis archaeogenetic studies on Sudan and Northeast Africa generally.

Two of the three examined Skhirat individuals were found to carry the paternal haplogroup T, and one bore the mtDNA haplogroup M1a1b. Sirak et al. (2021) have reported instances of both the LT parent clade and some T subclades (viz. T-L208 and T-Y31477) among Christian-era Nubian samples buried at the Kulubnarti site in Sudan (cf. Supplementary Figure 6).

The T clade occurs today at highest frequencies among Cushitic-speaking Somalis in the Horn of Africa (up to 80%; cf. Plaster (2011)). Additionally, the mtDNA haplogroup M1 is a signature mitochondrial lineage of the contemporary Afro-Asiatic-speaking populations in the Horn and Nile Valley (Stevanovitch et al. (2004); Non (2010), Table 3-3; Holden (2005)).

Y-DNA and mtDNA haplogroups carried by Middle Neolithic individuals excavated at Skhirat, a site in northern Morocco. The ancient pastoralists were found to bear the T paternal clade and the M1 maternal lineage (Simões (2023a)). These non-African haplogroups are signature uniparental markers of the modern Afro-Asiatic-speaking populations in the Horn and Nile Valley.

A Somali man (Cushitic). Haplogroup T, the signature Y-DNA lineage of the ancient Skhirat pastoralists, has been observed at frequencies of up to 80% among Somali individuals.

Genome analysis

In her more detailed 2023 dissertation, the lead researcher Luciana G. Simões asserts that the Middle Neolithic individuals from Skhirat harbor ancient Levantine ancestry, akin to that borne by early Cushitic individuals of the Pastoral Neolithic culture in eastern Africa:

We also generated new genomic data from the Maghrebi Middle Neolithic and identified the introduction of new ancestry, from the Levant. This ancestry, which has not been observed on the European side of the Mediterranean, is identified in association with a new ceramic tradition in northern Morocco and a pastoralist lifestyle, as a consequence of the expansion of cattle pastoralism in the current Sahara territory [168]. Levantine gene flow has been previously identified in pastoralist Neolithic East African individuals living around 4,000 years ago [169].

Genome analysis of Cushitic individuals, both ancient and modern, indicates that they predominantly bear non-African ancestry (over 70% on average). This non-African heritage consists of a Levantine Natufian component, a European Steppe component, and a newly-identified Upper Paleolithic European component. Additionally, they have minor Sub-Saharan African admixture (~25%), and trace North African Iberomaurusian admixture (under 5%).

The Skhirat pastoralists appear to have contributed a significant portion of the Levantine Natufian ancestry, which the Cushitic peoples carry (see Skh002 element in charts below).

Genome analysis of an ancient Cushitic individual of the Pastoral Neolithic (Elmenteitan) using a free-form model. This person predominantly bears non-African ancestry (~74.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (25.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Agaw individual using a free-form model. This person predominantly bears non-African ancestry (~80.4%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Guam_Latte elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (19.6% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a South Somali individual using a free-form model. This person predominantly bears non-African ancestry (~81.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Tigrayan individual using a free-form model. This person predominantly bears non-African ancestry (~81.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Kyrgyzstan_TianShan_Saka.SG elements), an Iran Neolithic component (Iran_C_TepeHissar element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Discussion

Taken together, the above appears to corroborate the traditional interpretation — supported by genetic analysis of modern African cattle breeds (cf. Decker et al. (2014)) — that pastoralism was first introduced to Africa by Afro-Asiatic-speaking herders, who ultimately arrived from the Middle East.

It also gives credence to Luis et al. (2004)‘s assertion that the T clade (formerly known as K2) was the original Y-DNA haplogroup in many areas of Africa. This paternal lineage would then have been replaced during subsequent expansions of other haplogroup carriers, particularly those bearing the E1b1b clade (formerly known as E3b). Luis et al. write:

K2-M70 is believed to have originated in Asia after the emergence of the K-M9 polymorphism (45–30 ky) (Underhill et al. 2001a). As deduced from the collective data (Underhill et al. 2000; Cruciani et al. 2002; Semino et al. 2002; present study), K2-M70 individuals, at some later point, proceeded south to Africa. These chromosomes are seen in relatively high frequencies in Egypt, Oman, Tanzania, Ethiopia, and Morocco and are especially prominent in the Fulbe (18% [Scozzari et al. 1997, 1999]), the highest concentration of this haplogroup found so far. The current patchy distribution of K2-M70 in Africa may be a remnant of a more widespread occupation. Subsequent demic events introducing chromosomes carrying the E3b-M35, E3a-M2, G-M201, and J-12f2 haplogroups may have overwhelmed the K2-M70 representatives in some areas. Like the R1*-M173 males, the M70 individuals could represent the relics of an early back migration to Africa from Asia, since these chromosomes are not associated with the G-M201, J-12f2, and R1-M173 derivatives, lineages that represent more-recent Eurasian genetic contributions (Semino et al. 2000; Underhill et al. 2001b). The K2-M70 expansion estimates in Egypt (17.5–13.7 ky; see table 3) are consistent with an early African diaspora. From the present-day African distribution of K2-M70, it is difficult to determine which of the two Africa/Asia migratory passages, if any, prevailed in its southward journey. However, the BATWING expansion estimates of both the Egyptian and Turkish K2-M70 lineages (13.7 ky and 9.0 ky, respectively) are much older than that of Oman (1.6 ky), which suggests that the Levantine corridor may have been used more extensively in the African dissemination of this lineage as well.

A recent lineage replacement does seem to have occurred in the Horn of Africa. Here, the V32 subclade of E1b1b is today ubiquitous despite the fact that it has only been observed in one pre-Iron Age individual (a Cushitic specimen of the Pastoral Neolithic). DNA analysis of ancient individuals in Sudan also hints that E1b1b-V32 may have been first brought to the Horn from the Nile Valley during the Meroitic period (see Hassan (2009)).

For details on this later expansion of E1b1b-V32 over a haplogroup T substrate, refer to Haplogroups E1b1b-V2 and E1b1b-V32 appear to be old Arabian lineages.

An Afar woman (Cushitic) and a Yemeni Arab woman (Semitic). Archaeogenetic analysis of Middle Neolithic herders from Skhirat, Morocco indicates that they bore Levantine Natufian genome ancestry. These ancient individuals appear to have been largely responsible for the cattle pastoralism tradition of Afro-Asiatic speakers in the Horn, Nile Valley, Chad Basin and northern Maghreb.

See also:

First genomes from the ancient Sahara detect two different groups of pastoralist inhabitants

Haplogroups E1b1b-V2 and E1b1b-V32 appear to be old Arabian lineages

Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa

Lactase persistence (lactose absorption) rates among Egyptians, Ashkenazi Jews, Iraqi Jews and Moroccan Jews

Genetic affinities of Nubians and Sudanese “Arabs” (Arabized Nubians)

Benadiri/Bravanese Somalis: Cushitic people with Persian, Indian and Bantu admixtures

A first look at Capsian ancient DNA and other updates

Genetic affinities of the Cushitic-speaking Beja

Recently, I stumbled upon some Global25 coordinates belonging to Beja individuals. I thought it would be apropos to have a look today at these samples’ genomes and see how closely they correspond to those of other Cushitic-descended peoples in Northeast Africa. We shall also examine the Beja’s uniparental markers (Y-DNA and mtDNA haplogroups), so as to get a handle on which population movements helped shape their gene pool.

Ababda Beja men (Cushitic).

Who are the Beja?

Here is some background info on the Beja, taken from Anthropometric analysis of the Cushitic-speaking Beja and Iraqw:

At present, the Beja are the most northerly situated Cushitic-speaking population. They are traditionally divided into four main subgroups: the Beni Amer (who live in Eritrea and Sudan), the Bisharin (who reside in Sudan), the Hadendoa (who also live in Sudan), and the Ababda (who dwell in Egypt).

Of these Beja divisions, the Bisharin have retained the native Cushitic language of the Beja people, known as Bedawiyet. Bedawiyet is the only surviving member of the North Cushitic branch of the Cushitic phylum. All of the other Beja subgroups have adopted other Afro-Asiatic languages. Most Beni Amer individuals now speak the Tigre language, an Ethiosemitic tongue spoken by the Tigre (a people who linguistic and genetic analyses suggest were themselves also originally Bedawiyet speakers). The Hadendoa and Ababda have, for their part, both switched to Arabic.

Despite these language shifts, genome analysis indicates that the Beja still share most of their ancestry with Cushitic/Ethiosemitic speakers of the northern Horn (cf. Dobon et al. (2015)).

A Bisharin Beja boy (Cushitic).

An Ababda Beja girl (Cushitic).

Y-DNA

Hassan et al. (2008) analysed various Afro-Asiatic and Nilo-Saharan-speaking populations of Sudan for Y-DNA. The scientists report that haplogroup E1b1b was the most common lineage among their Beja samples (22 out of 42 examined individuals or ~52%). This paternal clade is ubiquitous amongst Afro-Asiatic speakers, particularly in Africa. Furthermore, Hassan et al. observed that most of their tested Beja individuals specifically bore V32, a subclade of E1b1b that is typical of Cushitic speakers in Northeast Africa (for details, see Haplogroups E1b1b-V2 and E1b1b-V32 appear to be old Arabian lineages).

The remaining Beja samples in Hassan et al.’s dataset primarily carried the haplogroup J (~38%). All but one of these individuals bore the J1 subclade, which is prevalent among peninsular Arabs and Jews. This hints at past interactions between the Beja and Semitic speakers of the Middle East.¹ The rest of the examined Beja individuals belonged to the R1b haplogroup (2 out of 42 individuals or ~5%), a lineage mainly linked with the spread of Indo-European speakers; some also carried the archaic African haplogroup A (~5%), which is frequent amongst northern Nilotic populations.

A Beja man (Cushitic).

mtDNA

In terms of mtDNA, Hassan (2009) indicates that many of his Sudanese Beja samples bore the haplogroup L3 (17 out of 48 individuals or ~36%). Maca-Meyer et al. (2001) note that this “haplogroup L3 is more related to Eurasian haplogroups than to the most divergent African clusters L1 and L2,” owing to the fact that the L3 maternal lineage is immediately ancestral to the non-African M and N macroclades. Cabrerera et al. (2018) likewise propose that “carriers of mitochondrial DNA macrohaplogroup L3 basal lineages migrated back to Africa from Asia around 70,000 years ago.” As such, the original bearers of L3 may have introduced Upper Paleolithic European ancestry to Africa² (this newly-identified non-African ancestral element is a core component of Cushitic heritage — see Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa).

Many of the remaining Beja individuals belonged to various derivatives of the non-African M and N macroclades (~29%). These mitochondrial lineages consisted of the U6a1 clade (10.4%), followed by the preHV1 (8.3%), M1 (4.2%), T1 (2.1%), J1b (2.1%) and U3 (2.1%) haplogroups.

Additionally, a significant minority of the Beja samples bore (non-L3) derivatives of the Africa-centered macrohaplogroup L (~31%). These clades comprised L2 (16.7%), L1c (8.3%) and L0 (6.3%).

Overall, the Beja’s high frequencies of the L3 clade relative to non-African M and N mitochondria represents a characteristic difference between themselves and other Cushitic-descended peoples in the Horn. Non (2010) reveals that her Cushitic and Ethiosemitic-speaking samples bear high frequencies of M and N derivatives: Tigrai=63.6%, Somali=60%, Afar=56.2%, Gurage=55%, Oromo=53.3%, Nubian=48%, Amhara=46% (Table 3-3). This “urban pattern” appears to be relatively common among Afro-Asiatic-speaking populations in Africa and the Arabian peninsula because Kujanova ́ et al. (2009) reports a similar proportion of M and N lineages among Egyptians from El-Hayez oasis (68.6%), as does Bekada et al. (2015) among Algerians from Algiers (63%; cf. S5 Table), Coudray et al. (2009) among Figuig Berbers from northern Morocco (55.3%), Capocasa et al. (2023) among Douz Arabs (61%) and Rbaya Arabs (58%) of southern Tunisia (cf. Supplementary Table S2), and Kivisild et al. (2004) among Yemenis (56.4%).

A Beja woman (Cushitic).

Genome analysis

To gain an understanding of the Beja’s ancestral composition, I performed a genome analysis on two unofficial Global25 Beja samples. These consisted of Beni Amer and Hadendoa (Hadendowa) individuals from Sudan.

The Beja persons turned out to be genetically similar to other local Afro-Asiatic speakers. As with most other Cushitic, Ethiosemitic and North Omotic-speaking individuals of the Horn region, the Beja predominantly bear non-African ancestry (over 70% on average). This non-African heritage consists of a Levantine Natufian component, a European Steppe component, and the aforementioned Upper Paleolithic European component. They also have minor Sub-Saharan African admixture (~25%), and trace North African Iberomaurusian admixture (under 5%).

The foregoing supports the suggestion (discussed below under Notes) that much, if not all, of the L3 mtDNA subhaplogroups borne by the Beja were derived from a similar peninsular Arab source as the L3 which the Douz and Rbaya Arabs of Tunisia carry. This Arabian link is also supported by the fact that the Beni Amer Beja possess elevated Natufian ancestry like the Rbaya (~50%), and lower European Steppe and Upper Paleolithic European elements compared to other Cushitic samples from the Horn (correspondingly, the latter also have considerable Steppe-associated mtDNA lineages — see European Steppe-associated mtDNA haplogroups among the Cushitic peoples).

Another notable difference between the examined Beja individuals and other local Afro-Asiatic speakers is that both Beja samples show some evidence of Iran Neolithic admixture. However, unlike the Cushitic-speaking Saho of Eritrea — who bear a minor Caucasus Hunter-Gatherer admixture signal (see here); CHG is closely related to the Iran Neolithic component — this extraneous gene flow does not seem to have been mediated via late period dynastic Egyptian intermediaries. It instead appears to have been contributed by a more direct population source, one closer to the Iranian plateau. In the case of the Beni Amer sample, this is clear from the detection of Iran_C_TepeHissar, an Iran Neolithic element. In the case of the Hadendoa sample, the Iran Neolithic component is represented by the Turkmenistan_Gonur_BA_1 element.

Genome analysis of a Beni Amer Beja individual using a free-form model. This person predominantly bears non-African ancestry (~81.4%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an Iran Neolithic component (Iran_C_TepeHissar element), an ancient East Asian component (China_AmurRiver_N element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.6% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Hadendoa Beja individual using a free-form model. This person predominantly bears non-African ancestry (~73.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Skh002 and Israel_Natufian elements), a European Steppe component (Russia_North_Caucasus and Kyrgyzstan_TianShan_Saka.SG elements), an Iran Neolithic component (Turkmenistan _Gonur_BA_1 element), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (26.2% of the Kenya_Kakapel_LIA element).

Genome analysis of a Saho individual using a free-form model. This person predominantly bears non-African ancestry (~83.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian, Egypt_ThirdIntermediatePeriod and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (16.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a South Somali individual using a free-form model. This person predominantly bears non-African ancestry (~81.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Amhara individual using a free-form model. This person predominantly bears non-African ancestry (~78.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d element), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (21.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Tigrayan individual using a free-form model. This person predominantly bears non-African ancestry (~81.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d, Israel_Natufian and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Kyrgyzstan_TianShan_Saka.SG elements), an Iran Neolithic component (Iran_C_TepeHissar element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an ancient Cushitic individual of the Pastoral Neolithic (Elmenteitan) using a free-form model. This person predominantly bears non-African ancestry (~74.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (25.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Notes

¹Ancient DNA analysis has found that modern Semitic peoples of the Levant genetically cluster with Bronze Age Levantines, who in turn descend from the Pre-Pottery Neolithic makers of the Levant, the immediate descendants of the Natufians. Moreover, the Semites’ Pre-Pottery Neolithic and Mesolithic Natufian ancestors did not carry the haplogroup J and instead primarily bore the E1b1b lineage. This indicates that the ancestral Semites acquired the J clade relatively recently, during the Bronze Age, through interaction with peoples from the Iranian plateau/Caucasus. On this genetic contact, Haber et al. (2017) write: “We compiled frequencies of Y-chromosome haplogroups in this geographical area and their changes over time in a dataset of ancient and modern Levantine populations (Figure S12), and note, similarly to Lazaridis et al., that haplogroup J was absent in all Natufian and Neolithic Levant male individuals examined thus far, but emerged during the Bronze Age in Lebanon and Jordan along with ancestry related to Iran_ChL.”
²Hassan (2009) observed an even higher frequency of L3 mitochondrial subclades among his Nubian and Sudanese “Arab” (Arabized Nubian) samples. Farrell et al. (2013) also identified certain L3-rich areas in the Arabian Peninsula, where the local populations do not show any apparent Sub-Saharan African genomic influence. The Rbaya bedouins of Tunisia, one of the few populations in Africa that genuinely appear to be of recent peninsular Arab descent, seem to have originated from these areas since they have been observed to carry L3 at high frequencies (41%) while simultaneously bearing little Sub-Saharan African genome admixture (cf. Capocasa et al. (2023)). Consequently, Farrell et al. suggest that the L3 sublineages borne by some Afro-Asiatic and Nubian speakers in Africa could originally have been imported from the Middle East:  “We performed formal tests for a history of admixture and found no evidence of African admixture in the Saudi after the split. Taken together, these analyses suggest that the L3 haplogroup found in the Saudi were present before the bottleneck 50,000 YBP. Given the TMRCA estimates for the L3 haplogroup of approximately 70,000 YBP and the timing of the Out-of-Africa split, these analyses suggest that L3 haplogroup arose in the Middle East with a subsequent back migration and expansion into Africa over the Horn-of-Africa during the lower sea levels found during the glacial period bottleneck.” (*N.B. For the latest evidence on the suggested non-African origin of the mtDNA macrohaplogroup L3, see Cabrera et al. (2022).)

See also:

Haplogroups E1b1b-V2 and E1b1b-V32 appear to be old Arabian lineages

Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa

Anthropometric analysis of the Cushitic-speaking Beja and Iraqw

Genetic affinities of the Cushitic-speaking Saho

Genetic analysis of the Cushitic-speaking Agaws (Agaw proper) and Semitic-speaking Agaws (Abyssinians/Habesha)

Northern Somalis from Puntland (Majerteen Darod) have the most non-African ancestry in the Horn

Genetic affinities of North Omotic-speaking populations

Genetic affinities of Nubians and Sudanese “Arabs” (Arabized Nubians)

Benadiri/Bravanese Somalis: Cushitic people with Persian, Indian and Bantu admixtures

Phenotype of the Late Bronze Age Semites of Jordan

On Phenotype of the Canaanites, ancestors of modern Semitic peoples of the Levant and Mesopotamia, Genetic and phenotypic affinities of the Bronze Age Semites of Sidon, Lebanon and Genotype and phenotype of the Middle-to-Late Bronze Age Semites of Alalakh, we saw in detail how empirical genetic data indicates that, as recently as the Bronze Age, the Semitic peoples of the Levant and Mesopotamia almost all had relatively dark skin, dark eyes and dark hair. None were pale in complexion and very few had light eyes or light hair.

But what about the Late Bronze Age Semites of Jordan? Did they also have a swarthy physiognomy?

Let us find out.

Phenotypic traits

Lazaridis et al. (2022) conducted a comprehensive analysis of phenotypic traits borne by ancient individuals excavated in Europe and Asia, including the Late Bronze Age Semitic peoples of Jordan.

With regard to skin color, the scientists report that a little over half of the Jordanians were deeply pigmented (53%), having either a dark skin tone (5/15 specimens or 33%) or a dark-to-black skin tone (3/15 specimens or 20%). The other Jordanian individuals had an intermediate skin tone (7/15 specimens or 47%). None of the Jordanian individuals reportedly had a pale skin tone (0/15 specimens or 0%).

Similarly, in terms of hair color, Lazaridis et al. note that all of the Jordanians had darker manes (15/15 or 100%). Of these individuals, most had black hair (11/15 specimens or 73%) and a few had brown hair (4/15 or 27%). None of the Jordanian individuals had either red hair or blond hair.

Finally, with respect to eye color, all of the Jordanian individuals had brown eyes (15/15 or 100%). None were reported to possess blue eyes.

Phenotypic traits of ancient Asian and European individuals, including the Late Bronze Age Semitic peoples of Jordan. These Jordanian specimens were found to be all dark-haired and brown-eyed, and to have mostly intermediate-to-black skin tones. No Jordanian individual was pale-skinned, blue-eyed, or red-haired or blond-haired (Lazaridis et al. (2022)).

Discussion

The phenotypic findings above — which indicate that, on average, the Late Bronze Age Semites of Jordan were intermediate-to-black in skin color and exclusively had dark hair and brown eyes — are to be expected. This is because the Natufians of the Mesolithic (the ultimate ancestors of the Semitic peoples) did not possess any genetic variants which confer light skin, light eyes or light hair. These Jordanian results are also very similar to those reported for the ancient Canaanites and other Bronze Age Semitic peoples of the Levant and Mesopotamia (see links below in See also).

Light pigmentation-associated mutations first appear in the ancestral Semitic gene pool after the Natufian period, during the ensuing Neolithic era. Through contact with outsiders from the Mediterranean, who introduced Anatolian Neolithic ancestry to the Levant, Arabian peninsula and Mesopotamia, the Pre-Pottery Neolithic makers (immediate descendants of the Natufians) acquired the derived SLC24A5 allele. This is the main causal variant responsible for lighter skin coloration among populations of Europe, South Asia, the Middle East, North Africa and the Horn of Africa.

In the Late Bronze Age, the Semitic natives of the Levant, Arabia and Mesopotamia would have further interactions with foreigners from the Caucasus/Iranian plateau and Mediterranean. These contacts would serve to introduce additional light skin-associated variants into the Semitic gene pool (notably, the SLC45A2 allele). As a result, based on objective skin reflectance data, the Semitic peoples of the Middle East today have complexions which, on average, are intermediate between those of the lighter-skinned Iranic peoples (Kurds) and Anatolian Turks and the darker-skinned populations in most of Africa (see Appendix below).

Ancient wall painting from a monumental tomb at Beni Hasan, Egypt. The mural depicts early Levantine individuals. Note the figures’ brown skin tone, consistent with actual phenotype analysis of the Canaanites and other early Semitic peoples of the Bronze Age.

Modern adaptation of the Beni Hasan mural, falsely portraying the Bronze Age Semitic figures as being lighter-skinned than how they are actually shown on the original wall painting.

Jordanian Arab bedouin men, retaining the dark-skinned, dark-haired and dark-eyed phenotype of their Bronze Age Semitic ancestors.

Appendix

Gradation of skin colors for modern human populations in Africa, the Middle East and South Asia, based on observed skin reflectance values at the 685 nm reading. Shading from darkest to lighest gray represents greatest to least melanization. White areas are places where no skin reflectance data has been collected. On average, the examined Semitic populations in the Middle East have skin pigmentation levels between those of Iranic Kurds & Anatolian Turks and most examined African populations. The foregoing is consistent with ancient DNA data indicating that outsiders from the Caucasus/Iranian plateau vicinity interbred with the ancestors of the Semites in the Levant/Arabia/Mesopotamia. This admixture event(s) introduced the Caucasus Hunter-Gatherer component to these areas, an ancestral element which contains alleles that confer lighter skin pigmentation. As a result, the Semitic natives of northern Arabia, the Levant and Mesopotamia got fairer relative to their Canaanite forebears and other darker-complexioned ancestors of the Bronze Age ( (Jablonski and Chaplin (2000)).

Predicted skin color variation in modern human populations, based on multiple environmental factors (Chaplin (2004)). Note the difference between this hypothetical global skin color map (which is based on projected data) and the real global skin color map above (which is based on observed skin reflectance data). Both maps are from the same author.

Skin color variation in modern human populations, based on observed and extrapolated von Luschan scale data. According to Jablonski (2006), “this map gained broad circulation during the latter half of the twentieth century, even though Biasutti lacked information on skin color for some regions and was forced to extrapolate from known areas to fill in the gaps.” That said, “a more accurate map, based on a compilation of standardized skin reflectometry readings of indigenous peoples” can be seen above.

See also:

Phenotype of the Canaanites, ancestors of modern Semitic peoples of the Levant and Mesopotamia

Genetic and phenotypic affinities of the Bronze Age Semites of Sidon, Lebanon

Genotype and phenotype of the Middle-to-Late Bronze Age Semites of Alalakh

Phenotypic affinities of ancient Cushitic peoples

Genome analysis of modern Semitic individuals

Guide to the Universal Basic Genetic Model

Cushitic peoples carry European-derived Neanderthal alleles

On Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa, we saw how the ancient Zlatý kůň forager woman bore some Neanderthal-associated alleles (estimated at 3% of her genome). Since it turns out that Zlatý kůň-like ancestry is a core element of Cushitic heritage, modern Cushitic speakers who carry this Upper Paleolithic European genome component should also have inherited some Neanderthal variants.

Let us briefly investigate and confirm if this is in fact the case.

Origin and spread of Neanderthal variants

Schaefer et al. (2021) compared Neanderthal genomes to those of various modern populations, identifying haplotype block lengths that are indicative of Neanderthal genetic introgression. The scientists observed that their Somali, Mozabite Berber and Sahrawi samples were distinct from their other African samples and instead had haplotype block lengths similar to their non-African samples.

From this, Schaefer et al. conclude that “these Neanderthal haplotype blocks may have originated in ancient European migrants to eastern Africa.”

With the recent sequencing of the Zlatý kůň specimen, genome analysis now supports this claim.

Mean Neanderthal haplotype block lengths, which identify Neanderthal admixture in various modern populations. The Somali sample has haplotype block lengths similar to those of the North African (Mozabite Berber and Sahrawi) and non-African samples. Schaefer et al. (2021) suggest that this is because “these Neanderthal haplotype blocks may have originated in ancient European migrants to eastern Africa.” This is consistent with the Upper Paleolithic European ancestry and European Steppe ancestry, which have been detected in genome analyses of Cushitic individuals (Schaefer et al. (2021)).

Discussion

When properly genetically modelled, Cushitic, Ethiosemitic and North Omotic-speaking individuals of the Horn region predominantly bear non-African ancestry (over 70% on average). This non-African heritage consists of Levantine Natufian, European Steppe and Upper Paleolithic European components. Additionally, they have minor Sub-Saharan African admixture (~25%), and trace North African Iberomaurusian admixture (under 5%) (see Appendix below).

Because Levantine Natufian and ancient Sub-Saharan African specimens have been found to lack Neanderthal alleles, this implies that, among Cushitic-descended peoples, Neanderthal gene flow was indeed likely derived from folks who bore Upper Paleolithic European ancestry and/or European Steppe ancestry.

A Somali man (Cushitic). Cushitic peoples have been found to bear Neanderthal-associated alleles, like most non-African individuals. Haplotype analysis suggests that they inherited these genetic variants specifically from ancestors who bore Upper Paleolithic European ancestry and/or European Steppe ancestry. 

A Sahrawi girl (Berber origin). Peoples of Berber origin, including Sahrawi and Mozabite Berbers, have also been found to bear Neanderthal-associated alleles.

Appendix

Genome analysis of a South Somali individual using a free-form model. This person predominantly bears non-African ancestry (~81.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus and Hungary_DanubeTisza_EAvar elements), an ancient East Asian component (Vietnam_LN.SG element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (18.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of an Amhara individual using a free-form model. This person predominantly bears non-African ancestry (~78.2%). This non-African heritage consists of a Levantine component (Israel_Natufian_d element), a European Steppe component (Russia_North_Caucasus element), an ancient East Asian component (China_AmurRiver_N and Belize_9400BP elements), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (21.8% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

Genome analysis of a Wolayta individual using a free-form model. This person predominantly bears non-African ancestry (~74.8%). This non-African heritage consists of a Levantine component (Israel_Natufian_d and Skh002 elements), a European Steppe component (Russia_North_Caucasus element), and an Upper Paleolithic European component (Czechia_Bohemia_UP_HG element). Additionally, he/she has minor Sub-Saharan African admixture (25.2% of the Kenya_Kakapel_LIA and Malawi_Chencherere_LSA_5200BP elements).

See also:

Zlatý kůň: Upper Paleolithic European ancestry detected in Northeast Africa

European Steppe-associated mtDNA haplogroups among the Cushitic peoples

Detecting the Eurasiatic genome component in Cushitic peoples

Genome analysis of modern Semitic individuals

The contemporary Semitic-speaking populations of the Middle East are genetically and physically diverse. Most researchers have attributed this heterogeneity to differing admixture strains, which were superimposed on a shared Levantine Natufian ancestral base. A few have instead argued that this diversity exists because these groups actually have separate ethnic origins.

I wanted to find out the truth underlying these claims, so I performed a genome analysis.

An Ashkenazi Jew man (Semitic).

An Assyrian man (Semitic).

An Emirati Arab man (Semitic).

An Iraqi Arab man (Semitic).

A Jordanian Arab bedouin man (Semitic).

A Lebanese Arab man (Semitic).

A Mahra man (Semitic).

A Mizrahi Jew woman (Semitic).

An Omani Arab bedouin man (Semitic).

A Palestinian Arab man (Semitic).

An Rbaya Arab bedouin girl (Semitic).

A Saudi Arab man (Semitic).

A Sephardic Jew woman (Semitic).

A Socotri girl (Semitic).

A Syrian Arab man (Semitic).

A Yemeni Arab man (Semitic).

Samples

On the Vahaduo Admixture JS program, for the Target samples, I used most of the Semitic-speaking individuals listed in Eurogenes’ official Global25 modern datasheet (unscaled). For the Source samples, I utilized the Global25 ancient datasheet’s least-admixed representatives for each of the main West Eurasian and Sub-Saharan African ancestral components. I also included three additional ancient reference samples in order to capture any existing Iberomaurusian, East Eurasian and South Eurasian ancestries.

Least-admixed West Eurasian reference samples:

IRN_Ganj_Dareh_N (Iran Neolithic component)

Levant_Natufian_EpiP (Levantine Natufian component)

AZE_Caucasus_lowlands_LN (Caucasus Hunter-Gatherer component)

TUR_Marmara_Barcin_N (Anatolian Neolithic component)

WHG (Western Hunter-Gatherer component)

Yamnaya_RUS_Samara (European Steppe component)

Least-admixed Sub-Saharan African reference samples:

CMR_Shum_Laka (ancient Pygmy component)

COG_Kindoki_230BP (ancient Niger-Congo component)

KEN_Kakapel_300BP (ancient Nilo-Saharan component)

MWI_Chencherere (ancient East African Hunter-Gatherer component)

ZAF_2000BP (ancient Khoisan component)

Least-admixed Iberomaurusian reference sample:

MAR_Taforalt (Iberomaurusian component)

Least-admixed East Eurasian reference sample:

CHN_Huatuyan_500BP (ancient East Asian component)

Least-admixed South Eurasian reference sample:

LAO_Hoabinhian (Ancient Ancestral South Indian component)

Analysis

The resulting analytical table is as follows:

Semitic populations

Ashkenazi Jew (Belarus)

Ashkenazi Jew (Lithuania)

Ashkenazi Jew (Poland)

Ashkenazi Jew (Ukraine)

Assyrian

BedouinA

BedouinB

EmiratiA

EmiratiC

Iranian Jew

Iraqi

Jordanian

Kurdish Jew

Lebanese Christian

Lebanese Druze

Lebanese Muslim

Palestinian

Saudi

SaudiB

Sephardic Jew

Syrian

Syrian Jew

Yemenite Al Jawf

Yemenite Amran

Yemenite Dhamar

Yemenite Jew

Yemenite Mahra

Yemenite Marib

Discussion

From the data tables above, we may note that:

• The Iran Neolithic component (Iran_Ganj_Dareh_N) is found at highest frequency in a Syrian individual (27.8%), and at lowest frequency in various Yemeni, Saudi, Bedouin, Ashkenazi Jew and Sephardic Jew individuals (0%). The Assyrian samples have the highest average percentage of this Iran Neolithic ancestral element (17.3%), whereas the Yemenite Al Jawf samples bear this component at the lowest average frequency (1.4%). Overall, this Iran Neolithic component occurs at an average frequency of 7% among our Afro-Asiatic-speaking samples.
• The Levantine Natufian component (Levant_Natufian_EpiP) is found at highest frequency in a Yemenite Mahra individual (80.2%), and at lowest frequency in four Syrian, Syrian Jew, Ashkenazi Jew and Assyrian individuals (0%). The Yemenite Mahra samples also have the highest average percentage of this Levantine Natufian ancestral element (73.4%), while the Sephardic Jew samples carry this component at the lowest average frequency (5.2%). Overall, this Epipaleolithic Levantine Natufian component occurs at an average frequency of 31.1% among our Afro-Asiatic-speaking samples.

  

  Mahra men (Semitic). Among Semitic peoples, the Levantine Natufian genome component attains its highest average frequency among Yemenite Mahra individuals (73.4%).

• The Caucasus Hunter-Gatherer component (AZE_Caucasus_lowlands_LN) is found at highest frequency in a Syrian individual (78.6%), and at lowest frequency in various Ashkenazi Jew individuals and another Syrian individual (0%). The Assyrian samples have the highest average percentage of this Caucasus Hunter-Gatherer ancestral element (56.4%), whereas the Yemenite Mahra samples bear this component at the lowest average frequency (19.3%). Overall, this Caucasus Hunter-Gatherer component occurs at an average frequency of 33.6% among our Afro-Asiatic-speaking samples.

  

  Assyrian children (Semitic). Among Semitic peoples, the Caucasus Hunter-Gatherer genome component attains its highest average frequency among Assyrian individuals (56.4%).

• The Anatolian Neolithic component (TUR_Marmara_Barcin_N) is found at highest frequency in an Ashkenazi Jew individual (50.4%), and at lowest frequency in various Yemeni, Saudi, Emirati, Bedouin, Iraqi and Assyrian individuals (0%). The Belarus Ashkenazi Jew samples have the highest average percentage of this Anatolian Neolithic ancestral element (32.8%), while most Yemenite, Saudi, EmiratiA and BedouinB samples carry this component at the lowest average frequency (0%). Overall, this Anatolian Neolithic component occurs at an average frequency of 11.6% among our Afro-Asiatic-speaking samples.

  

  An Ashkenazi Jew man (Semitic). Among Semitic peoples, the Anatolian Neolithic genome component attains its highest average frequency among Ashkenazi Jew individuals (32.8%).

• The Western Hunter-Gatherer component (WHG) is found at highest frequency in an Ashkenazi Jew individual (6.4%), and at lowest frequency among most other Semitic individuals (0%). The Ukraine Ashkenazi Jew samples have the highest average percentage of this Western Hunter-Gatherer ancestral element (1.6%), whereas most of the other Semitic samples bear this component at the lowest average frequency (0%). Overall, this Western Hunter-Gatherer component occurs at an average frequency of 0.3% among our Afro-Asiatic-speaking samples.
• The European Steppe component (Yamnaya_RUS_Samara) is found at highest frequency in an Ashkenazi Jew individual (27.2%), and at lowest frequency among most Yemeni, Saudi and Iraqi Jew individuals (0%). The Belarus Ashkenazi Jew samples have the highest average percentage of this European Steppe ancestral element (21.2%), while the Yemenite Mahra and Yemenite Marib samples carry this component at the lowest average frequency (0%). Overall, this Steppe component occurs at an average frequency of 8% among our Afro-Asiatic-speaking samples.
• The ancient Pygmy component (CMR_Shum_Laka) is found at highest frequency in a Syrian individual (2.8%), and at lowest frequency among almost all other Semitic individuals (0%). The BedouinA samples have the highest average percentage of this Pygmy ancestral element (0.4%), whereas almost all of the other Semitic samples bear this component at the lowest average frequency (0%). Overall, this ancient Pygmy component occurs at an average frequency of 0% among our Afro-Asiatic-speaking samples.
• The ancient Niger-Congo component (COG_Kindoki_230BP) is found at highest frequency in a Syrian individual (8.8%), and at lowest frequency among most other Semitic individuals (0%). The Syrian samples have the highest average percentage of this Niger-Congo ancestral element (1.3%), while most of the other Semitic samples carry this component at the lowest average frequency (0%). Overall, this ancient Niger-Congo component occurs at an average frequency of 0.2% among our Afro-Asiatic-speaking samples.

  

  Syrian Arabs (Semitic).

• The ancient Nilo-Saharan component (KEN_Kakapel_300BP) is found at highest frequency in a SaudiB individual (13.4%), and at lowest frequency in various mainly Yemeni, Ashkenazi Jew, Sephardic Jew, Assyrian and Syrian individuals (0%). The Jordanian samples have the highest average percentage of this Nilo-Saharan ancestral element (6.1%), whereas the Yemenite Mahra samples bear this component at the lowest average frequency (0%). Overall, this ancient Nilo-Saharan component occurs at an average frequency of 3% among our Afro-Asiatic-speaking samples.

  

  Jordanian Arab men (Semitic). Among Semitic peoples, the ancient Nilo-Saharan genome component attains its highest average frequency among Jordanian individuals (6.1%).

• The ancient East African Hunter-Gatherer component (MWI_Chencherere) is found at highest frequency in an Ashkenazi Jew individual (1.6%), and at lowest frequency among almost all other Semitic individuals (0%). The Ukraine Ashkenazi Jew samples have the highest average percentage of this East African Hunter-Gatherer ancestral element (0.1%), while all of the other Semitic samples carry this component at the lowest average frequency (0%). Overall, this ancient East African Hunter-Gatherer component occurs at an average frequency of 0% among our Afro-Asiatic-speaking samples.
• The ancient Khoisan component (ZAF_2000BP) is found at highest frequency in an Ashkenazi Jew individual (1.4%), and at lowest frequency among almost all other Semitic individuals (0%). The Lithuania Ashkenazi Jew samples have the highest average percentage of this Khoisan ancestral element (0.2%), whereas almost all of the other Semitic samples bear this component at the lowest average frequency (0%). Overall, this ancient Khoisan component occurs at an average frequency of 0% among our Afro-Asiatic-speaking samples.
• The Iberomaurusian component (MAR_Taforalt) is found at highest frequency in a Sephardic Jew individual (10.2%), and at lowest frequency among various Yemeni, Bedouin, Saudi, Emirati, Jordanian, Palestinian, Lebanese and Assyrian individuals (0%). The Sephardic Jew samples have the highest average percentage of this Iberomaurusian ancestral element (6.9%), while the Yemenite, Saudi, EmiratiA and BedouinB samples carry this component at the lowest average frequency (0%). Overall, this ancient Iberomaurusian component occurs at an average frequency of 1.9% among our Afro-Asiatic-speaking samples.

  

  A Sephardic Jew man (Semitic). Among Semitic peoples, the Iberomaurusian genome component attains its highest average frequency among Sephardic Jew individuals (6.9%).

• The ancient East Asian component (CHN_Huatuyan_300BP) is found at highest frequency in a Palestinian individual (both 10.2%), and at lowest frequency in various mainly Yemeni, Emirati, Bedouin, Saudi and Lebanese individuals (0%). The Palestinian samples also have the highest average percentage of this East Asian ancestral element (5.2%), whereas the Saudi and EmiratiA samples bear this component at the lowest average frequency (0%). Overall, this ancient East Asian component occurs at an average frequency of 2.2% among our Afro-Asiatic-speaking samples.

  

  A Palestinian Arab man (Semitic). Among Semitic peoples, the ancient East Asian component attains its highest average frequency among Palestinian individuals (5.2%).

• The Ancient Ancestral South Indian component (LAO_Hoabinhian) is found at highest frequency in an EmiratiC individual (29.2%), and at lowest frequency among most other Semitic individuals (0%). The EmiratiC samples also have the highest average percentage of this Ancient Ancestral South Indian element (10.9%), while the Syrian Jew, Lebanese Druze, Lebanese Christian, BedouinB and Assyrian samples carry this component at the lowest average frequency (0%). Overall, this Ancient Ancestral South Indian component occurs at an average frequency of 1% among our Afro-Asiatic-speaking samples.

  

  Emirati Arab men (Semitic). Among Semitic peoples, the Ancient Ancestral South Indian genome component attains its highest average frequency among EmiratiC individuals (10.9%).

Conclusion

In summation, the genomic evidence indicates that the present-day Semitic-speaking populations of the Middle East may be divided into seven basic groups based on their ancestral composition:

1. The first group consists of populations having a predominant Levantine Natufian ancestry (>70% on average), with secondary Caucasus Hunter-Gatherer-related elements (CHG & Iran Neolithic components). These include the Mahra and other Yemenis, BedouinB, Saudi and EmiratiA individuals.
2. The second group comprises populations with an almost equal apportionment of Levantine Natufian and Caucasus Hunter-Gatherer ancestries. These primarily consist of BedouinA individuals.
3. The third group consists of individuals with an excess of Caucasus Hunter-Gatherer ancestry, but still bearing significant Levantine Natufian ancestry as well as moderate Anatolian Neolithic ancestry. This includes Palestinians, Jordanians, Iraqis, most Syrians, most EmiratiC individuals, most Lebanese (Muslims and Christians), and most Lebanese Druze.
4. The fourth group comprises individuals with predominant Caucasus Hunter-Gatherer ancestry (>70% on average), with ancillary Levantine Natufian and Anatolian Neolithic ancestries. This includes some Syrians and some Assyrians.
5. The fifth group consists of individuals who almost entirely bear Caucasus Hunter-Gatherer-related ancestry (mostly CHG & some Iran Neolithic), with very little Levantine Natufian and Anatolian Neolithic ancestry. This comprises most Assyrian individuals.
6. The sixth group comprises individuals with elevated Caucasus Hunter-Gatherer ancestry, moderate Anatolian Neolithic ancestry, lesser European Steppe ancestry, and minimal Levantine Natufian ancestry. This includes Sephardic Jew, Syrian Jew and many Ashkenazi Jew individuals.
7. The seventh and last group consists of many Ashkenazi Jew individuals with elevated Anatolian Neolithic ancestry, moderate Caucasus Hunter-Gatherer ancestry, notable European Steppe ancestry, and lesser Levantine Natufian ancestry.

One data point which stands out is the very heterogeneous nature of the modern Syrian population. Moreover, the fact that all but two of the Semitic samples lacked the MWI_Chencherere component of the ancient East African Hunter-Gatherers is quite telling. Since this population element is found at an average frequency of 8.6% among Afro-Asiatic-speaking populations of the Horn (see here), this reveals that the Cushitic component that certain researchers have detected at low frequency in the genomes of some contemporary Middle Eastern Semitic speakers (<10%) was specifically derived from Sudanese “Arabs” and/or Nubians (i.e. Cushitic-related peoples from the Nile Valley, where East African Hunter-Gatherer admixture is practically non-existent and Nilo-Saharan admixture is instead more pronounced; cf. Gopalan et al. (2019)). Finally, the presence of the Iberomaurusian/Taforalt component at highest frequencies among the Ashkenazi Jew samples suggests a connection with North Africa. Such a link is also supported by these individuals’ excess Anatolian Neolithic ancestry, which has long been a common ancestral element in the Maghreb.

See also:

Genotype and phenotype of the Middle-to-Late Bronze Age Semites of Alalakh

How and when the Ashkenazi Jew and Sephardic Jew gene pool was formed

Brief note on the Levantine origin of Ashkenazi and Sephardic Jews

Ashkenazi Jews and Sephardic Jews are of the same Levantine ancestral origin as Mizrahi Jews, Palestinians and other Semites

The difference between “pure” Semites (Mahra) and Afro-Arabs (Akhdam)

Y-DNA haplogroup E1b1a (formerly E3a) appears to have originated in the Middle East

Guide to the Universal Basic Genetic Model

23andme and AncestryDNA misrepresent the ancestral composition of Afro-Asiatic speakers

Medieval DNA from Socotra confirms that the Socotri are among the “purest” descendants of the Levantine Natufians

Haplogroups E1b1b-V2 and E1b1b-V32 appear to be old Arabian lineages

Phenotype of the Canaanites, ancestors of modern Semitic peoples of the Levant and Mesopotamia

Genetic and phenotypic affinities of the Bronze Age Semites of Sidon, Lebanon

Genetic analysis of the Cushitic-speaking Agaws (Agaw proper) and Semitic-speaking Agaws (Abyssinians/Habesha)

Myths about the Afro-Asiatic-speaking populations debunked

Tutsi Bantus are of Nilotic origin

I have read some propaganda online claiming that the Tutsi, a Bantu-speaking people living in Central-East Africa, are of Cushitic origin. This could not be further from the truth. The Tutsi are actually of Nilotic origin, as should be obvious from their close physical resemblance to the Maasai and other Cushitic-admixed Nilotes inhabiting the Great Lakes region.

Tutsi Bantu men and woman.

Craniometrics and physiognomy

Craniometrically, the anthropologist G. P. Rightmire (1975) observed that his Iron Age Rwanda male and female samples cluster with his Bantu and Khoisan samples. All of these specimens are also morphologically distinct from the ancient Cushitic male and female samples, which instead cluster with the ancient Egyptian male and female samples (Baharini and Makalia in particular):

Furthermore, because the Tutsis’ Bantu ancestors in the past assimilated some Southern Cushites, the average Tutsi individual today tends to have narrower facial features than other Niger-Congo/Nilo-Saharan speakers. However, the nasal width of the Tutsi (estimated by Jean Hiernaux (1974) at 39 mm, similar to the Maasai Nilotes) remains significantly broader than that of Afro-Asiatic speakers (estimated at 36 mm for the Warsangali Darod Somali of northern Somalia).

Likewise, David Blackwell (1984) photogrammatically analysed the nasal morphology of the Tutsi Bantus as compared to that of the Hehe Bantus and found that these Niger-Congo-speaking populations are similar to each other on most indices, including nasal prominence, nasal bridge distance, columella length, nasal height, nasal angle, nasal wing/septum relationship, and nasal region vertical relationship. Moreover, although the Tutsis generally have a lower nasal index than their Hehe brethren, the Tutsi also have a considerably larger nasal breadth index and inter-occular nasal width index. This “again suggests the Tutsi’s noses are wider relative to face width than the Hehe’s”.

Additionally, Hiernaux (1974) reports that all Tutsis have kinky hair texture like the Hutu Bantus, Maasai Nilotes and other Niger-Congo, Nilo-Saharan and Khoisan populations.

Altogether, this tells us that the Tutsi’s Iron Age ancestors were not Cushitic people, but instead morphologically “Negroid” and thus similar in physiognomy to other Bantus, Nilotes and Khoisan.

Blood groups

Serological (blood group) analyses indicate that, overall, the Tutsis are similar to other Bantus/Nilotes. Lawrence Oschinsky (1960) states:

The Batutsi are tall, narrow nosed, narrow faced Negroids showing evidence of previous Caucasoid admixture, the Bahutu are intermediate in stature and have broader noses and faces, and the Batwa represent a local variant of Congo pygmy racial type. In the ABO, Rh and MN blood group distributions the Batutsi and the Bahutu are similar.

J. D. Fage (2013) likewise notes that:

The origin of the Tutsi/Hima/Chwezi ruling class in the lacustrine Bantu kingdoms is an intriguing question. Serologically they are Blacks, and this seems to rule out the possibility of a Cushitic origin.

Genetics

Genetically, the Tutsis again show close ties with the Maasai Nilotes and Hutu Bantus.

Peer-reviewed studies report that 75% of Tutsi individuals in Rwanda and Burundi carry the E1b1a haplogroup, a clade formerly known as E3a (Trombetta et al. (2015); Luis et al. (2004); Luis et al. (2007)). This is the most common paternal lineage borne today by the Hutu Bantus and other Niger-Congo-speaking individuals, as well as most Maasai. Only 1% of Tutsis in Rwanda and 22% of Tutsis in Burundi have been found to harbor M296, a subclade of the Afro-Asiatic-affiliated haplogroup E1b1b (formerly E3b), which is associated with the ancient spread of Southern Cushites:

In terms of mtDNA, Göbel et al. (2019) report that almost all of their Rwanda sample belongs to derivatives of the macrohaplogroup L. Much of this mitochondrial variation was also shared with the Kenya cohort. The scientists did, however, detect a minor Cushitic influence in their Rwandan dataset via the presence of the West Eurasian maternal lineages M1a1 (2.6%), N1a1a (1.3%) and K1a (0.6%) as well as the U6a clade.

For their overall ancestry, all studies that have examined the autosomal DNA of Tutsi individuals indicate that their STR markers are most similar to those of the Hutu and other Niger-Congo-speaking populations (cf. Coudray et al. (2006); Simms et al. (2008); Shepard and Herrera (2006b); Shepard and Herrera (2006a)):

Miao et al. (2024) compared the genomes of Rwandan individuals with those of various global populations. They observed that their Rwandan sample clustered with their Nilote Luo sample from Kenya, consistent with the Nilotic roots of the Tutsi. According to the researchers:

Rwanda is known as the heart of Africa, reflecting the history of the world. Colonization and genocide have led to Rwanda’s existing genetic structure. Herein, we used massively parallel sequencing to analyze 296 loci in 185 Rwandans and constructed a database for Rwandan forensic data for the first time. We found the following results: First, forensic parameters demonstrated that all loci were highly informative and could be used for forensic identification and paternity tests in Rwandans. Second, we found that the differences in genetic background between Rwandans and other African populations were similar but slight, as indicated by the massively parallel sequencing panel. Rwandans belonged to the African population and were inseparable from populations from neighboring countries. Also, Rwandans were closer to the European and American populations because of colonization, war, and other reasons. There was no scientific basis for racial classification established by colonization. Further research still needs to be carried out on more loci and larger Rwandan samples.

Autosomal DNA analysis of Rwandans and other global populations. The Rwandan individuals (RW) cluster with the Nilote Luo sample from Kenya (LWK), consistent with the Nilotic origins of the Tutsi (Miao et al. (2024)).

Autosomal SNP analysis by 23andme, AncestryDNA and other commercial genome testing labs similarly report that Tutsi individuals share most of their ancestry (close to 70%) with Niger-Congo/Nilo-Saharan/Khoisan individuals. Of this predominantly Sub-Saharan African ancestry, much of it is associated with Southern East African Bantus (44.8% for the Rwandan Tutsi individual below, tested with 23andme). They also have some admixture from Nilotic peoples from the Sudan area (16.2% below). Additionally, Tutsis bear admixture derived from Cushitic peoples of Ethiopia and Eritrea (34.2% below), whom their Nilotic/Bantu ancestors absorbed:

Finally, genome analysis of Tutsi individuals using the Vahaduo Admixture JS program indicates that, like the Maasai and other Nilotic peoples inhabiting the Great Lakes region, they trace most of their proximal or recent ancestry to the Cushitic-admixed Nilotes of the Pastoral Iron Age. This confirms that the Tutsi are indeed of Nilotic origin since all peoples of genuine Cushitic origin (such as the Somali, Oromo, Afar, Agaw/Abyssinians) trace instead their proximal ancestry to either the Cushites of the Pastoral Neolithic or the Cushitic-related inhabitants of medieval Kulubnarti. Genetic distance analysis also indicates that the Tutsi and Hima share closest genetic affinity with the Nilotic Ogiek people of Kenya and other mostly Nilotic/Bantu populations, further confirming the Nilotic origins of the Tutsi-Hima.

An Ogiek woman (Nilotic). The Ogiek, a Nilo-Saharan-speaking population of Kenya, mostly bear the ancient Nilo-Saharan component. Genetic distance analysis also indicates that, among all modern groups, the Ogiek are the people with whom the Tutsi and Hima Bantus share the closest affinity. This further confirms the Nilotic ancestral origin of the Tutsi-Hima.

For a complete biological analysis of the Tutsi Bantus, I recommend the excellent essay The Elongated African fallacy on the Land of Punt website. It covers all of the above and much more in detail, including the Tutsi’s HLA antigen affinities and other classical genetic markers.

Appendix

Tutsi men and women:

Among affluent Tutsi individuals and models, skin bleaching/lightening and the wearing of hair weaves or chemical hair straightening is relatively common. These cosmetic alterations serve to obscure the Tutsis’ predominant Nilotic/Bantu ancestry (over 70%), while highlighting their minority Cushitic admixture (under 30%). This practice can be seen below in the person of Ange Kagame, a Tutsi individual, and daughter of the Rwandan president Paul Kagame:

Tutsi socialite Ange Kagame (Bantu), in her natural state. Note Ange’s very dark skin tone, owing to the Tutsi’s Nilotic ancestral origins. Also notice her naturally kinky hair texture, which all Tutsi individuals have.

Ange Kagame, after skin lightening treatments and wearing straight hair extensions. She can now pass for some of the mixed peoples in the northern Great Lakes/southern Horn, a contact zone where Bantus/Nilotes have intermixed with Cushitic peoples.

Tutsi Bantu women au naturel.

Overall, the Tutsi closely resemble the Hutu, another Bantu-speaking population inhabiting Rwanda and Burundi.

Tutsi and Hutu men, women and children:

Maasai Nilotic men and women — note the close physical resemblance to the Tutsi Bantus, stemming from their shared Nilo-Saharan origins:

For additional images of the Tutsi Bantus, see Tutsi Bantus are of Nilotic origin – Photographic adjunct.

See also:

Tutsi Bantus are of Nilotic origin – Photographic adjunct

A very Eurasiatic Toubou sample

Baggara/Shuwa “Arabs”: last remaining descendants of the original Chadic speakers

Anthropometric analysis of the Cushitic-speaking Beja and Iraqw

Genetics of the Hausa, Chadic speakers of Niger-Congo origin

Genome analysis of the Tutsi Bantus