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11 Saturday Mar 2023
The ancestral origin of the Afro-Asiatic-speaking groups inhabiting the Horn of Africa has long been a subject of debate among scholars. In the 2000s, with the advent of genomics, a number of scientists proposed that these communities were formed through contact between ancient West Eurasian and Sub-Saharan African individuals. They principally based this on admixture analyses, which compared the genomes of persons from various reference populations with those of Afro-Asiatic speakers from the Horn region. The earliest of these genomic studies typically used modern Europeans and modern West Africans as their proxy groups. However, as more global populations were analysed, it became clear to researchers that virtually every living individual is mixed to some degree, thereby making contemporary groups inadequate reference populations for genomic analysis. This realization prompted scientists to increasingly turn to ancient specimens, which they assumed — usually correctly — would be less admixed than modern individuals and therefore more reliable proxies.
Llorente et al. published one such paleogenetics study in 2015, analysing for the first time an indigenous hunter-gatherer from the Horn (see Ancient DNA from Ethiopia). At the time of publication, this forager individual, called Mota, was believed to be “purely” African and thus free of non-African admixture. Subsequent examination found that there, in fact, had been a software-related error; the specimen apparently did harbor some Eurasian ancestry, albeit a trivial amount (cf. Erratum). Consequently, researchers on the genetic affinities of the Horn’s Afro-Asiatic speakers continued to utilize Mota as their favored ancient Sub-Saharan African proxy, with newly-analysed early Levantine populations (viz. Mesolithic Natufians, Pre-Pottery Neolithic makers) serving as their preferred stand-ins for West Eurasian ancestry.
In 2020, Wang et al. released an archaeogentics paper that included a number of never before analysed ancient specimens from eastern Africa. These newly-published samples helped unveil several layers of ancestry in the Mota forager, which had previously been hidden. Mota’s ancestral makeup was revealed to actually consist of various hunter-gatherer elements (Hadza, Mbuti and Khoisan-related components), as well as minor Niger-Congo/Nilo-Saharan-related admixture and considerably higher Eurasian admixture than previously had been realized (~25%). Additionally, genome analysis found that the earliest Afro-Asiatic-speakers yet to be analysed in eastern Africa (i.e. the Cushitic settlers of the Pastoral Neolithic) carried a predominant West Eurasian ancestry related to ancient North African/Levantine groups, with ancillary Sub-Saharan African admixture. The analysis also showed that a 300 BP individual from the Kakapel site in Kenya bore the most such Sub-Saharan African ancestry, thus supplanting Mota as an ideal proxy (cf. Supplementary Material).
Thanks to these new samples, we can now estimate with much greater accuracy than ever before the actual ancestral composition of the Afro-Asiatic-speaking populations from the Horn of Africa.
Admixture Analysis
To assist us in this endeavor, we will make use of the Vahaduo Admixture JS program, which interprets data from the official Global25 datasheets. Global25, or G25, is a powerful genetic system originally developed by the Eurogenes project. Based on Principal Coordinate Analysis (PCA) coordinate data, it allows users to compare the genome of any modern or ancient individual against that of well over 6000 ancient samples culled from around the world, as well as several thousand modern samples.
According to Genoplot:
Global 25 (G25) ancestry modeling is currently the most effective way to perform free-form discovery of ancestry. The set of coordinates used by Global 25 are a product of Eurogenes. They were created as a means to facilitate ancestral admixture modeling[…]
This type of ancestry modeling allows for very precise and granular determination of ancestry. Unlike admixture calculators which come locked-in to pre-defined ancestral components, G25 modeling allows for selection of any number of potential ancestral sources.
Furthermore, through its Single function, Vahaduo has the capacity to cross-analyse thousands of genomes in a matter of minutes. This gives it a distinct advantage over other genetic modeling programs, which typically can only process a handful of genomes at a time. As Bulbul and Kidd (2021) note, the accuracy of an autosomal SNP analysis is dependent upon which reference populations have been selected and the sample size of each of those reference populations. If a key reference population has been omitted from study or if a selected reference sample is too small in terms of the number of individuals in contains, the results produced will be incorrect and unrepresentative. It is therefore imperative for accurate ancestry estimation to have as large as possible a global coverage of reference populations:
One can only estimate an individual’s ancestry among the existing reference populations. Initially, a panel of SNPs is good only for the set of populations used to select the SNPs. As more populations are tested, the SNP panel may have broader applicability. Basically, the inference of ancestry can only be as good as the global coverage of the reference populations and the reference populations need to provide reasonable estimates of the allele frequencies of all the SNPs in the specific panel of SNPs.
Hence, Global25 ancestry modeling is the best tool for our genome inquiry.
Step 1: Identify the “purest” Sub-Saharan African reference population available
As a first step in our genetic investigation, we will try and determine which ancient Sub-Saharan African population the contemporary Nilotic peoples share the most affinity with. We are specifically interested in such individuals because genome analysis by Prendergast et al. (2018) and Wang et al. (2020) shows that the Cushites of the Pastoral Neolithic absorbed an African population carrying ancestry similar to that borne by modern Nilotes.
We begin by running a Vahaduo Admixture JS Distance analysis on the ancient African samples listed on the Global25_PCA datasheet. In order to not invalidate our findings, these Source populations must exclude any ancient specimens possessing substantial Eurasian ancestry (viz. the Pastoral Neolithic, Kulubnarti, Pastoral Iron Age, Kenya Iron Age, ancient Egyptian, ancient Moroccan, Guanche and Zanzibar Euro specimens). For our Target populations, we will employ all of the modern Nilote samples listed on the Global25_PCA_modern datasheet. The resulting Vahaduo Distance analysis, shown here, indicates that every single Nilotic sample (except for one heavily mixed Datog individual) shares greatest genetic affinity with the Kakapel 900BP cohort from Kenya.
In Wang et al.’s Figure S1-B below, we can further see that this Kakapel 900BP sample primarily bears Nilo-Saharan/Niger-Congo-related ancestry (purple component), similar to that which defines the contemporary Dinka Nilote sample from Sudan. Kakapel 900BP thus seems to be the earliest appearance of a modern Dinka-like specimen in the archaeogenetic record. However, like the Dinka of today, Kakapel 900BP also bears a non-trivial amount of West Eurasian admixture (red component). This makes the sample an unsuitable proxy for inferring “pure” ancient Nilotic ancestry.
To find a less admixed Sub-Saharan African reference sample, we turn instead to the Kakapel 300BP sample from the same site. As with Kakapel 900BP, we note that the Kakapel 300BP cohort appears among the top eight ancient African specimens with whom the modern Nilote samples show the closest genetic ties on Vahaduo’s Distance parameter. We know that this affinity is due to shared ancient Nilotic ancestry rather than shared non-African admixture since, in Wang et al.’s Figure S1-B above, Kakapel 300BP carries only a small proportion of the West Eurasian red component, unlike both Kakapel 900BP and the Mota forager from Ethiopia (labeled Ethiopia_4500BP).
As an additional precaution, in order to ensure that Kakapel 300BP is indeed the most optimal source of “pure” Sub-Saharan African ancestry available, we will conduct a Vahaduo Multi analysis (process described below). We shall use the Kakapel 300BP sample as a Source population to infer ancient Nilo-Saharan ancestry in all of the contemporary Nilotic samples listed on the Global25_PCA_modern datasheet. Our resulting genetic model, displayed here, is successful; it produces plausible admixture levels for all of the examined Nilotic individuals and at acceptable Distance fits of <9%. By contrast, tests using other ancient Sub-Saharan African specimens, which have the highest levels of the Nilo-Saharan/Niger-Congo purple component above, are all failures. These experiments either grossly exaggerate the non-African admixture levels in the modern Nilotic samples and/or do not capture any Natufian admixture, which these specimens are known to possess (since they assimilated some Cushitic peoples, who bear such West Eurasian ancestry).
Finally, we will consult Wang et al.’s height data on the Kakapel 300BP sample (cf. Supplementary Material). Since the Sidamo and Sab, respectively, are both the most African-admixed and shortest of the Afro-Asiatic-speaking populations in Ethiopia and Somalia (see Ancient DNA from Ethiopia), this will enable us to determine whether the Kakapel 300BP specimen represents the main ancient Sub-Saharan African peoples whom the early Afro-Asiatic speakers in Northeast Africa absorbed. Kakapel 300BP, a young adult woman, is listed by Wang et al. at a diminutive stature of 156 cm or 5’1.5″, confirming that she likely does represent that ancient Sub-Saharan African contact population.
Step 2: Identify which Eurasian ancestries the modern Afro-Asiatic speakers of the Horn carry
Next, we will try and ascertain which specific Eurasian ancestries the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn region actually bear. To accomplish this, we will avail ourselves of the Vahaduo Admixture JS program’s Single tab and compare these modern samples (which are listed on the Global25_PCA_modern datasheet) with all 6000+ ancient Eurasian samples listed on the Global25_PCA datasheet. To this official G25 datasheet we shall add the coordinates for the newly-analysed EGY_1879BCE sample, which belong to the Middle Kingdom ancient Egyptian aristocrat Nakht-Ankh:
| EGY_1879BCE:Nakht-Ankh,0.0012,0.129,-0.044,-0.0965,-0.0031,-0.0534,-0.017,-0.0078,0.0551,-0.0049,0.0138,-0.0172,0.0306,-0.0015,0.0069,-0.0072,-0.0111,0.0053,-0.004,0.0042,-0.0012,0.0046,-0.0078,0.0026,-0.0013 |
Before proceeding, we will also make sure to exclude any ancient samples that have non-trivial Sub-Saharan African admixture. The end result, shown below, has identified four principal non-African ancestries, three of which are West Eurasian (ancient Egyptian component, European-related Steppe component, and Levantine Natufian component) and one which is East Eurasian (East Asian component).
In order to verify our preliminary findings, we will again run Vahaduo’s Single analysis. However, now we shall include our “pure” early Nilotic sample (the Kakapel 300BP specimen) as a Source population alongside all of the ancient Eurasian populations from the Global25_PCA datasheet, while keeping our modern Afro-Asiatic-speaking individuals from the Horn as our Target populations.
As an additional cautionary measure, we shall again conduct a Vahaduo Single analysis. Now we will add all of the other ancient Sub-Saharan African samples that have the least Eurasian admixture. These samples represent the “purest” Niger-Congo (COG_Kindoki_230BP:KIN002), East African Hunter-Gatherer (MWI_Chencherere:I4421_new_all), Pygmy (CMR_Shum_Laka:I10874_new_all), and Khoisan (ZAF_2000BP:bab001) specimens. Appending these samples will help us ascertain whether our Target Horn populations realistically carry any extra Sub-Saharan African admixture and from which sources.
As can be seen above, all of the aforementioned Eurasian ancestries appear once more, confirming their authenticity. Among the West Eurasian elements, the ancient Egyptian component is best represented by the EGY_1879BC sample, which is also the oldest Egyptian sample whose Global25 coordinates are available; the European-related Steppe component is best represented by the RUS_Baikal_BA specimens from Bronze Age Russia; and the Levantine Natufian component is best represented by the Levant_Natufian_EpiP specimens, which date to the Mesolithic. The East Eurasian element is best represented by the CHN_Huatuyan_500BP sample from late medieval China. We can also, for the first time, observe robust estimates of the actual ancestral proportions that characterize the Afro-Asiatic speakers analysed. On average, modern Cushitic, Ethiosemitic and North Omotic-speaking individuals appear to carry over 70% Eurasian ancestry (most of which is West Eurasian, with a significant East Eurasian element), with around 25% Sub-Saharan African admixture (primarily derived from early Nilo-Saharan speakers like Kakapel_300BP, and secondarily derived from ancient East African Hunter-Gatherers like MWI_Chencherere) and under 5% Epipaleolithic North African ancestry (i.e. Iberomaurusian/Taforalt component).
Step 3: Quantify these ancestral proportions
To further break down the apportionment of these ancestral elements, we will now make use of the Vahaduo Admixture JS program’s Multi function. In order to efficiently interpret our findings, we shall utilize the five ancient Eurasian and two ancient Sub-Saharan African “best representative” samples, which we have just identified above. As expected, the end result (shown below) appears very similar to Vahaduo’s Single analysis:
With regards to the table above, we may note that:
Tables of ancestral proportions for each Afro-Asiatic-speaking population (derived from the Vahaduo Admixture JS program’s Multi function):
Summary Vahaduo Multi table of the averages of each genome component for all the examined Horn populations:
| Ancient East Asian (CHN_Huatuyan_500BP) | Ancient Pygmy (CMR_Shum_Laka) | Ancient Niger-Congo (COG_Kindoki_300BP) | Ancient Egyptian (EGY_1879BCE) | Iran Neolithic (Ganj_Dareh_N) | Ancient Nilo-Saharan (KEN_Kakapel_300BP) | Natufian (Levant_Natufian_EpiP) | Iberomaurusian (MAR_Taforalt) | East African Hunter-Gatherer (MWI_Chencherere) | European Steppe (RUS_Baikal_BA) | Anatolian Neolithic (TUR_Marmara_Barcin_N) | Ancient Khoisan (ZAF_2000BP) | Average non-African Ancestry | Average Sub-Saharan African Ancestry | Average Ancient North African Ancestry | |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Afar (Ethiopia) | 23.9% | 0.8% | 0% | 18.2% | 0% | 14.4% | 24.3% | 0.9% | 5.5% | 12.1% | 0% | 0% | 78.4% | 20.7% | 0.9% |
| Agaw (Ethiopia) | 21.9% | 0.2% | 0% | 19% | 0% | 13.9% | 23.1% | 1.0% | 8.1% | 12.8% | 0% | 0% | 76.8% | 22.2% | 1.0% |
| Amhara (Ethiopia) | 22.2% | 0.2% | 0% | 18.1% | 0% | 14.7% | 24.9% | 1.0% | 7.7% | 11.3% | 0% | 0% | 76.4% | 22.6% | 1.0% |
| Eritrean | 20.4% | 1.1% | 0% | 16.7% | 0% | 16.7% | 30.0% | 0.2% | 4.5% | 10.3% | 0% | 0% | 77.5% | 22.3% | 0.2% |
| Ethiopian Jew | 22.6% | 0.3% | 0% | 17.6% | 0% | 15.1% | 26.4% | 0.3% | 7.5% | 10.3% | 0% | 0% | 76.8% | 22.9% | 0.3% |
| Iraqw (Tanzania) | 22.9% | 0% | 0% | 18.2% | 0% | 21.7% | 5.2% | 2.4% | 18.3% | 11.3% | 0% | 0% | 57.6% | 40.0% | 2.4% |
| Oromo (Ethiopia) | 22.9% | 0.1% | 0% | 17.2% | 0% | 17.6% | 19.4% | 0.8% | 11.6% | 10.4% | 0% | 0% | 69.9% | 29.3% | 0.8% |
| Rendille (Kenya) | 25.6% | 0% | 0% | 18.1% | 0% | 25.7% | 10.7% | 1.8% | 9.6% | 8.5% | 0% | 0% | 62.9% | 35.3% | 1.8% |
| Somali (Kenya) | 23.6% | 0.2 | 0% | 18.5% | 0% | 20.4% | 14.1% | 2.1% | 9.1% | 12.1% | 0% | 0% | 68.2% | 29.7% | 2.1% |
| Somali (South Somalia) | 23.6% | 0% | 0% | 19.3% | 0% | 20.0% | 13.3% | 3.0% | 7.7% | 13.1% | 0% | 0% | 69.3% | 27.7% | 3.0% |
| Tigray (Ethiopia) | 21.7% | 0.1% | 0% | 17.8% | 0% | 15.5% | 26.3% | 0.8% | 6.4% | 11.5% | 0% | 0% | 77.2% | 22.0% | 0.8% |
| Wolayta (Ethiopia) | 21.7% | 0.9% | 0% | 16.8% | 0% | 17.5% | 15.6% | 1.8% | 14.7% | 10.9% | 0% | 0% | 65.1% | 33.1% | 1.8% |
Step 4: Corroborate these findings with other scientific evidence
As a fourth step in our analysis, we shall further establish the veracity of our findings by corroborating them with other scientific evidence gathered from different disciplines. We will focus on the Steppe element because the ancient Egyptian component is fully discussed on Punt: an ancient civilization rediscovered.
In regards to the European-related Steppe ancestry, which we have just identified above, uniparental markers (both Y-DNA and mtDNA) support the existence of such an influence in Northeast Africa. Gad et al. (2020) report that the 18th Dynasty ancient Egyptian Pharaoh Amenhotep III, his son Pharaoh Akhenaten and his grandson Pharaoh Tutankhamun, who governed from the Amarna site in Upper Egypt, belong to the Y-DNA haplogroup R1b (cf. Gad et al. (2020a); Gad et al. (2020b)). iGENEA further specifies that these Amarna royals fall under the clade’s M269 branch, the most common paternal lineage carried today by European males. Yatsishina et al. (2021) likewise divulge that one of the ancient Egyptian mummies they analysed at the Kurchatov Institute bears the R1b-M269 haplogroup. Maternally, Khairat et al. (2013) state that a mummified ancient Egyptian individual they studied belongs to the I2 mtDNA haplogroup. This mitochondrial lineage has been detected among various early Steppe cultures of Europe. It is nowadays quite rare (typically under 5%), attaining a global frequency peak of 23% among Cushitic-speaking remnant groups in the Great Lakes region (Castrì et al. (2008)). Moreover, the basal I* haplogroup has only been observed among three persons worldwide; two of these individuals are from Somalia and the other is from Iran (Olivieri (2013)). White et al. (2023) also detected the rare H4a1 mtDNA clade in the mummy of Takabuti, an ancient Egyptian noblewoman from Thebes, Upper Egypt. The scientists note that this maternal lineage has been “described in antiquity only in Central Europe,” and propose that the haplogroup is “suggestive of the introduction of new gene pools during the Late Period of ancient Egyptian history.”
Y-DNA haplogroups of the ancient Egyptian Pharaoh Amenhotep III, his son Pharaoh Akhenaten, and his grandson Pharaoh Tutankhamun, who ruled from the Amarna site in Upper Egypt. These 18th Dynasty kings belong to the R1b clade, which today is the most common paternal lineage borne by Europeans. This finding supports an ancient presence in the Nile Valley of peoples bearing European-related Steppe ancestry (Gad et al. (2020a)).
Autosomal DNA analysis of the Amarna kings has also unveiled a clear Steppe-related ancestral affinity. Hawass et al. (2010) typed these ancient Egyptian monarchs for short tandem repeats (STRs or microsatellites). The genetic testing company DNA Tribes then compared these specimens’ microsatellite markers against those belonging to various modern populations contained within its internal database. It reported that these specimens showed greatest genetic affinity with the autosomal STR profiles of contemporary Sub-Saharan African individuals. However, this conclusion is rendered doubtful by the relatively small size of DNA Tribes’ internal reference population database. Using the PopAffiliator autosomal STR prediction software (which is now defunct), Gourdine et al. (2018) similarly asserted that these Amarna mummies had a greater probability of assignment to PopAffiliator’s “Sub-Saharan Africa” cluster (41.7%-93.9%) than to its “Eurasia” (3%-41.5%) or “Asia” (0.3%-16.7%) clusters. However, a glance at PopAffiliator’s STR Worldwide Database, where it lists the reference populations against which it compared the Amarna specimens’ microsatellites, reveals that these comparison samples were few in number, consisting of just 80 populations in total. PopAffiliator, moreover, included a sample from Somalia among its four reference populations from Sub-Saharan Africa. This had the effect of biasing and invalidating its microsatellite assignments because autosomal STR analysis of ethnic Somalis has found that these Cushitic-speaking individuals actually possess considerable Eurasian affinities (Steele et al. (2014), the largest global autosomal STR analysis, groups its Somalia sample within its IC6 Middle East cluster). Y-DNA (Trombetta et al. (2015), Supplementary Table 7)), mtDNA (Stevanovitch et al. (2004)), and autosomal DNA (Hodgson et al. (2014), Supplementary Text S1; Dobon et al. (2015); Almarri et al. (2021), Table S4) analyses likewise indicate that Somalis and other Afro-Asiatic speakers from the Horn share close ties with both ancient and modern Egyptians. Altogether, this explains why PopAffiliator clustered the Amarna mummies as it did: the specimens were showing affinity with the Somalia reference sample, specifically.
Furthermore, a cross-analysis of the Amarna royals’ autosomal STRs with the microsatellites listed on the more extensive Allele Frequency Database (ALFRED) — which includes well over 400 different worldwide populations, including many understudied South Asian groups — demonstrates instead a close affiliation with populations in South Asia and Europe (see our study Autosomal STR Analysis of the Ancient Egyptian Amarna Royal Family, Pharaoh Ramesses III, and Unknown Man E (Prince Pentawere). We know that this affinity is specifically tied to ancient Steppe-bearing peoples because the alleles with a primary South Asian affiliation also often include among their top results ALFRED’s eastern European samples, and eastern Europe is where the Steppe component is believed to have ultimately originated (e.g. 22.50% of Croatians carry the Pharaoh Akhenaten’s FGA=23 allele, which peaks at 40% in a Reddy/Vanne sample from South Asia). The reverse is also true. That is, the alleles with a primary European affiliation likewise frequently include South Asian groups among their top results (e.g. 48% of the Drokpa in South Asia carry the courtier Thuya’s D7S820=10 allele, which peaks at 52.60% in a Croatian sample). This finding agrees with our Vahaduo Admixture JS analysis, as well as with the Steppe-associated uniparental markers that these ancient Egyptians carry.
What’s more, analysis of limb proportions is suggestive of contact with populations inhabiting the temperate zone. Holliday (2013) examined various old and recent global samples, including an ancient Egyptian cohort (dating from the Predynastic to Middle Kingdom), a Kerma sample (dating from the Middle Kingdom i.e., the same time period as the Global25 sample EGY_1879BCE, which belongs to Nakht-Ankh), and a Christian-era Nubian sample (dating from the 4th to 7th centuries CE). He reports that his Middle Ages Nubian cohort had a more cold-adapted body plan, similar to his medieval European samples. On the other hand, Holliday’s ancient Egyptian cohort and Kerma sample had more linear, tropically-adapted limbs, similar to the “leptosome tendency” which Coon (1939) affirms is typical of Bedouin Hadhramis in southern Arabia. This is consistent with our analysis below, which indicates that the Christian period inhabitants of Kulubnarti in Nubia bore some Steppe-related admixture. It therefore appears that during Nakht-Ankh’s lifetime, Steppe ancestry, although already present since the Neolithic period, had not yet spread throughout the Nile Valley.
The cephalic indices (CI) of the ancient Egyptian Amarna monarchs also attest to a European Steppe affiliation. Kemp and Zink (2012) report that, with the exception of the courtier Yuya, who is markedly dolichocephalic (long-headed, with a CI of 70.3), all of these royal mummies are either mesocephalic (medium-headed) or brachycephalic (broad-headed). That is, they possess cephalic index values of 75 or greater. Akhenaten and Tutankhamun are, in fact, brachycephalic, with high cephalic indices of 81.0 and 83.9, respectively. This is atypical for both ancient and modern Egyptians, who for the most part tend toward dolichocephaly (cephalic indices under 75). However, brachycephaly is common in eastern Europe, where frequencies of the Steppe ancestral component are today maximized (cf. Godina (2011)).
Additionally, philological evidence backs a Steppe connection. For example, Bahadur (1917) notes that “Eusebius states that Ethiopians [Meroites] emigrating from the River Indus settled in the vicinity of Egypt [Meroe].” Nilus similarly relayed to Apollonius Tynaeus that “the Indi are the wisest of all mankind. The Ethiopians [Meroites] are a colony from them: and they inherit the wisdom of their forefathers.” The Indus river mostly traverses Pakistan, an area that historically was settled by Indo-European speakers, who carried Steppe ancestry.
Craniometric analysis likewise points to a close association between Afro-Asiatic speakers in Northeast Africa, Indo-European-speaking and Dravidian-speaking populations of South Asia, and Europeans. This affinity extends back in time to subsume early groups in these areas, including the ancient Egyptians and post-Neolithic Nubians. Brace (1993), for instance, asserts that “insofar as India has metric ties with any other populations, it combines with Nubia [Bronze Age/X-Group and Medieval/Christian Era samples] and then the Somalis to join Europe and the Egyptians [Predynastic and Late Dynastic samples] as a last link before that set of branches ties in with the rest of the world.” Multiple other studies have also observed a similar affiliation (e.g. Morton (1854), Stoessiger (1927), Coon (1939), Sergent (1997), Brace et al. (2006)). This aligns well with a diffusion of ancient Steppe-bearing peoples into Northeast Africa and South Asia, either indirectly from a common waypoint near Central Asia or directly from Europe.
Craniometric analysis of ancient and contemporary global populations. The modern Afro-Asiatic-speaking populations in Northeast Africa, the ancient Egyptians, the Bronze Age Nubians (X-Group) and Medieval Nubians (Christian period), modern Indo-European-speaking and Dravidian-speaking groups of India, and modern and ancient Europeans cluster together. This is consistent with a spread of Steppe-bearing peoples from Europe to these areas, either directly or indirectly via Central Asia (Brace (1993)).
Furthermore, hair morphology supports an early European-associated presence in Northeast Africa. Lazaridis et al. (2022) conducted a comprehensive analysis of phenotypic traits borne by ancient individuals exhumed in Europe and Asia. The scientists indicate that blond hair was most common among their ancient European specimens and that red hair was exclusively found among these samples. This is key since Brothwell and Spearman (1963), employing reflectance spectrophotometry, observed a number of authentically blond ancient Egyptian individuals; their hair samples were not affected by either hair dye or cuticular damage. Strikingly, Griggs (1988) reports that excavations he led at a Roman-Christian era cemetery in Seila, located in the Fayum region of Egypt, yielded the remains of many individuals with blond or red hair. He affirms that “of the 37 adults whose hair was still preserved[…] there were 4 redheads, 16 blondes, 12 with light or medium brown hair, and only 5 with dark brown or black hair. Of those whose hair was preserved 54% were blondes or redheads, and the percentage grows to 87% when light-brown hair color is added.” Similarly, Janssen (1978) writes that “330 graves were excavated in cemetery 221 (Meroitic) and a proportion of blond individuals of Caucasoid type found.” Hrdy (1978), also analysing Meroitic remains, notes that many ancient individuals buried at Semna South in Sudanese Nubia had blond or red hair. He suggests that this “probably points to a significantly lighter-haired population than is now present in the Nubian region.”
Correspondingly, several of the ancient Egyptian mummies belonging to the Amarna royal lineage, including the aristocrats Yuya, Thuya and Tiye as well as the Pharaoh Ramesses III, also have blond or red hair. These are the same monarchs who, as we have just seen, carry some European-affiliated autosomal STR markers. In the case of Yuya, Hawass and Saleem (2016) argue that his blondism was caused by either the fading of henna dye on white hair or the interaction of embalming materials and applied henna. However, the Egyptologist Janet Davey of the Victorian Institute of Forensic Medicine later demonstrated through laboratory experimentation that natron used in the mummification process has no effect on hair color, regardless of whether or not a specimen’s hair had been dyed with henna during embalming. This finding was confirmed by a followup microscopic analysis conducted by Gale Spring of RMIT (cf. Smith (2016)). Hamed and Maher (2021), using various analytical processes (viz. microscopic examination, FTIR spectroscopy, gas chromatography and raman spectroscopy), also observed that Yuya’s natural hair color is indeed a reddish-blond. Elemental analysis of his hair shafts showed a high concentration of sulphur, an element that occurs in greater quantities in blond or red hair than in black hair. Tiye and the other royal mummies likewise reportedly have naturally red or blond hair. What’s more, a tomb painting of the Pharaoh Amenhotep III depicts him with a light reddish mane. This altogether underlines the fact that blond and red hair were prevalent among the ancient Egyptian Amarna nobles.
In summary, the existence of an old European Steppe ancestral component in Northeast Africa, analogous to that which we have detected through our Vahaduo Admixture JS genome analysis, is supported by studies on the Amarna royal family. These ancient Egyptian monarchs ruled from the Amarna site in Upper Egypt during the 18th Dynasty. Their mummies have been found to bear Steppe-related autosomal STR markers, Steppe-related uniparental haplogroups (notably, the R1b Y-DNA clade and the I mtDNA clade), primarily mesocephalic or brachycephalic skulls, and blond or red hair, much like populations in eastern Europe where the Steppe component originally dispersed from. Moreover, craniometric examination of modern Afro-Asiatic speakers and Nubians in Northeast Africa reveals close ties with groups in South Asia and Europe, consistent with population movements from the European Steppe into Central Asia and from there into both the Nile Valley/Horn and the Indian subcontinent. Such early migrations are also backed by ancient texts, which assert that peoples from the Indus river vicinity settled near Egypt, in the Meroë area situated in present-day Sudan.
Step 5: Confirm whether the ancient Pastoral Neolithic and Kulubnarti specimens share this ancestral composition
Fifthly, we shall explore if the ancient Cushitic settlers of the Pastoral Neolithic and the Christian-era Kulubnarti specimens from Nubia carry these same ancestral components and at similar frequencies. This will help us determine whether the Eurasian elements, which we have just identified, are legitimate ancestries or mere statistical constructs. It will also give us some insight as to which of these components are the original ancestries of the Afro-Asiatic speakers and which components are instead intrusive (i.e., elements acquired later through interbreeding).
We start off by launching another Vahaduo Single run, this time using the Pastoral Neolithic samples as our Target populations. The “pure” ancient Sub-Saharan African specimens (Kakapel_300BP, MWI_Chencherere, COG_Kindoki), CMR_Shum_Laka, ZAF_2000BP) and the 6000+ ancient Eurasian specimens listed on the Global25_PCA datasheet will once more serve as our Source populations. The end result is displayed below. It again primarily shows the same assortment of West Eurasian (viz. ancient Egyptian, Steppe and Natufian components) and East Eurasian (East Asian component) ancestries, which occur at similar total frequencies as before (over 70%). The remaining minority of the Pastoral Neolithic individuals’ ancestral composition consists of Sub-Saharan African admixture (~25%) and North African Iberomaurusian admixture (under 5%), just like the modern Afro-Asiatic speakers from the Horn region.
When we repeat this process for the Kulubnarti samples, we encounter the same ancestral components occurring again at practically identical percentages. However, one key difference between the Kulubnarti individuals on the one hand, and the Pastoral Neolithic and modern Horn samples on the other, is that the Kulubnarti specimens have little East African Hunter-Gatherer admixture. Almost all of their ancient Sub-Saharan African admixture instead consists of the Nilo-Saharan Kakapel300BP element.
Lastly, when we run a Vahaduo Multi analysis on both our Pastoral Neolithic and Kulubnarti ancient samples, here too we come away with similar ancestral components and at comparable percentages:
With regards to the Pastoral Neolithic table above, we may note that:
Tables of ancestral proportions for each Pastoral Neolithic group:
Step 6: Confirm whether modern Egyptians share this ancestral composition
Sixthly, we will examine whether contemporary Egyptian individuals have the same ancestral makeup as just outlined. As our Target population on Vahaduo Admixture JS, we shall use the Egyptian samples listed on Eurogenes’ official Global25_PCA_modern datasheet. For our Source populations, we will again utilize the Eurasian samples on the Global25_PCA datasheet, alongside the five “pure” ancient Sub-Saharan African representatives. With our Target and Source populations now loaded, we will tap into Vahaduo’s Single function, letting the program sift through Global25’s massive 6000+ samples to pinpoint for us the most optimal sources of ancient Eurasian ancestry.
From the above, it is apparent that modern Muslim Egyptians (i.e. non-Coptic Egyptians) do generally share the same ancestral composition as the Afro-Asiatic-speaking populations in the Horn of Africa. West Eurasian elements (ancient Egyptian, European-related Steppe, and Levantine Natufian components) and an East Eurasian element (East Asian component) are most prominent here too, and minor Sub-Saharan African admixture and a minute North African Iberomaurusian admixture can again be detected. That said, many of the analysed Muslim Egyptian individuals also show an affinity with the contemporary Levant/Arabia, as exemplified by the Levant_Tell_Qarassa_Early_Antiquity component. This West Eurasian ancestral element — which mainly consists of Natufian ancestry, with some later acquired Anatolian Neolithic and Caucasus Hunter-Gatherer/Iran Neolithic admixtures — is most typical of modern Arabic speakers, as well as many Yemeni Jews and some Mahra individuals (see Vahaduo Single analysis here).
Like before, to re-confirm the main ancestries that we have just identified and organize them into table format for easier interpretation, we will end with a Vahaduo Multi analysis. We shall include the Levant_Tell_Qarassa_Early_Antiquity sample to capture the modern Arabian admixture that is present in Egyptians. Next, we shall re-run our Multi analysis and include G25’s TUR_Marmara_Barcin_N and IRN_Ganj_Dareh_N samples among our Source populations so as to account for, respectively, the Anatolian Neolithic and Iran Neolithic elements that are inherent in the Levant_Tell_Qarassa_Early_Antiquity component:
As can be seen in the tables above, the Muslim Egyptian individuals still bear the same ancestries as other Afro-Asiatic speakers in the Horn of Africa and at comparable frequencies, confirming the observed affinities. On average, modern Egyptians carry around 84% non-African ancestry, the majority of which consists of West Eurasian elements (viz. primarily ancient Egyptian, Steppe and Natufian components, as well as some extra Anatolian Neolithic and Iran Neolithic admixtures) and a minority of which comprises an East Eurasian element (East Asian component). Furthermore, they also harbor around 13% Sub-Saharan African admixture (primarily consisting of the Nilo-Saharan element, and secondarily of the Niger-Congo element) and a very minor North African Iberomaurusian admixture (3%).
Coptic Egyptians are not yet included on the official Global25 datasheets, but they seem to have a somewhat different ancestral composition. This is suggested by the Coptic samples on IllustrativeDNA, a service which uses G25 technology to process its own coordinates (not official Global25 coordinates). When these coordinates are run through Vahaduo Admixture JS’s Single function, the Coptic individuals appear to derive over 99% of their ancestry from the EGY_1879BCE ancient Egyptian sample. However, since IllustrativeDNA’s samples are often low coverage, producing bloated fits on Vahaduo’s Distance parameter of >15%, they frequently do not allow for precise identification of all the ancestral components an individual might bear nor can they accurately quantify those elements (e.g. when used as Target populations against Global25’s ancient Eurasian Source populations, IllustrativeDNA’s Afro-Asiatic-speaking samples from the Horn have Vahaduo Distance fits which are around three times higher than Eurogenes’ official Global25 Horn samples; compare this with this). We must therefore perform an additional Vahaduo Distance analysis on our Coptic samples to make sure that they are reliable. When this is done, our Copts and other Egyptian samples from IllustrativeDNA wind up showing almost identical Distance fits as Eurogenes’ official Global25 Egyptian samples (see here and here).
This confirms that Coptic Egyptians indeed descend directly from Egyptians dating from at least the earlier Dynastic period — something which was already strongly implied by their traditional language, Coptic, a later iteration of the ancient Egyptian tongue. As such, Coptic Egyptians seem to have been largely unaffected by the aforementioned gene flow into the Nile Valley, which we have detected in both the ancestral Cushites of the Pastoral Neolithic and the later Dynastic period Egyptians. Having said that, besides the Y-DNA haplogroup E1b1b common among Afro-Asiatic speakers, some modern Coptic individuals also bear the R1b and J paternal clades (E1b1=74% and J1=1% among Copts in Upper Egypt according to Crubézy et al. 2010; E1b1b=21%, J=45% and R1b=15% among Copts in Sudan according to Hassan et al. (2008)). The R1b and J lineages were first brought to the Egypt area by newcomers bearing, respectively, European-related Steppe ancestry (seemingly introduced as early as the Neolithic) and Caucasus Hunter-Gatherer/Iran Neolithic ancestry (introduced during the later Dynastic period). Hence, there is genetic evidence of contact between Coptic Egyptians and foreigners. Our Vahaduo Single analysis below, however, demonstrates that this gene flow was ultimately also of negligible importance because, unlike Muslim Egyptians, Copts (both in Sudan and Egypt) for the most part do not bear these ancestral components.
We will finish by conducting a Vahaduo Multi analysis on all of the Coptic and Egyptian samples discussed above. To these we shall add Hollfelder et al. (2017)’s Coptic sample, as well as that paper’s Nubian cohort and other Sudanese Afro-Asiatic-speaking groups. The resulting data table, shown below, indicates that Hollfelder et al.’s samples, including their Coptic representative, are unreliable since they have the tell-tale inordinately high Distance fits (on average >15%; one Sudanese “Arab” Shaigia sample has an absurd Distance fit of 47%), few identified ancestral elements (usually 2 to 3 maximum), and skewed component frequencies typical of low coverage samples. However, it is interesting to note that the Egyptian samples from Cairo and Mansoura share an almost identical genetic profile as Coptic Egyptians, albeit with a small Sub-Saharan African admixture. Both of these samples are from IllustrativeDNA and have acceptable Distance fits of <9%. Since Cairo and Mansoura are cities located in northern Egypt, where the EGY_1879BCE specimen was excavated from at Deir Rifeh, we therefore now have, for the first time, genetic evidence substantiating the existence of Upper Egyptian/southern Egyptian and Lower Egyptian/northern Egyptian types.
Anthropologists have long observed such a cleavage in the Nile Valley’s skeletal record. For example, Batrawi (1946) remarks that the ancient Egyptians appeared to have been divided into two distinct but related physiognomies, and that “the study of the available measurements of the living, however, apparently suggests that the modern population all over Egypt conforms more closely to the southern type” (also see G. Billy (1975)). We can now state that the primary distinguishing factor between these two osteological types seems to have been gene flow from the European Steppe and later also from the Arabian peninsula rather than Sub-Saharan African admixture. This is because the contemporary Egyptian samples from Global25 and the northern Egyptian samples from IllustrativeDNA have almost the same low level of Sub-Saharan African admixture (13% and 11%, respectively). Ergo, after the early Dynastic period, foreign influences from Europe and Asia significantly impacted northern Sudan and southern Egypt as compared to northern Egypt.
Step 7: Confirm whether modern Sudanese “Arabs” also share this ancestral composition
For the seventh step in our analysis, we shall inquire whether Sudanese “Arabs” share the same ancestral composition as the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa. We will follow the exact same procedure as just described above for Egyptians, using Kababish as our Sudanese “Arab” cohort. To these we shall add two other samples from Sudan, of Rashaayda Arabs and Baggara “Arabs.” In addition, we will examine Daza (Gorane or southern Toubou) individuals and other Baggara “Arabs,” both from Chad. All of these samples were originally published in Fortes Lima et al. (2022) and later converted to Global25 coordinates for use in the Vahaduo Admixture JS software program (see here for the unscaled or raw G25 coordinates; though not official Global25 samples, they are high coverage/decent quality).
Our resulting Vahaduo Single analysis is as follows:
Judging by the Single analysis above, it is apparent that the Kababish “Arabs” generally bear the same ancestral makeup as the Horn’s Afro-Asiatic speakers.
To better organize our thoughts and strengthen our interpretation, we will finish by conducting a Vahaduo Multi analysis:
From the data table above, it is clear that the Kababish “Arabs” of Sudan do have the same overall ancestral composition as the Afro-Asiatic speakers from the Horn of Africa. The Kababish individuals bear a predominant Eurasian ancestry (averaging almost 70%), comprising majority West Eurasian elements (ancient Egyptian, European Steppe, and Levantine Natufian components) and a minority East Eurasian element (East Asian component). Furthermore, these individuals carry some Sub-Saharan African admixture (close to 30%) and a trace Iberomaurusian admixture (under 1%). One characteristic difference, however, is that the Kababish’s Sub-Saharan African admixture almost entirely consists of the “pure” ancient Nilo-Saharan component (KEN_Kakapel_300BP). Their comparatively trivial frequencies of the East African Hunter-Gatherer component (MWI_Chencherere), averaging just 1%, are instead more congruent with our modern Egyptian and ancient Kulubnarti results above, as well as our results for contemporary Libyans below. All of these samples have negligible frequencies of this forager element, which emphasizes that this component is indeed autochthonous to eastern Africa rather than the Nile Valley.
For their part, the Baggara “Arab” samples from Sudan and Chad are very similar to the Sudanese Kababish “Arab” cohort. These populations, in fact, appear to be of the same origin. However, it is evident from the figures above that the Baggara have intermixed more with their Nilo-Saharan-speaking neighbors since they harbor greater average percentages of the KEN_Kakapel_300BP component (40.2% for the Baggara in Chad and 47.3% for the Baggara in Sudan).
The samples belonging to the Daza (Gorane), or southern Toubou, are almost identical to those of the Baggara “Arabs.” Just one key difference separates these two groups, and that is the Daza’s appreciable frequencies of the North African Iberomaurusian component (averaging 14.5%). In this respect, the Daza individuals appear more similar to the modern Libyans (discussed below), who have a comparable average percentage of this Taforalt element. It may be that the Daza and Teda or northern Toubou — as hypothesized successors of the ancient Garamantes/Garamantians, whose old territory they currently occupy — were originally Afro-Asiatic speakers of Libyan stock. Hence, as Kirwan (1934) observes, the etymological connection between the ethnonyms Goran and Garamantes. However, since the Daza and other Toubou intermingled significantly with their Nilo-Saharan neighbours, it is conceivable that they eventually adopted the latter’s language. This ultimately would have served to obscure their Berber origins.
The Rashaayda Arabs (Rashaida) of Sudan and Eritrea are relative newcomers to Northeast Africa. Their arrival from the Hejaz region of Saudi Arabia during the 19th century is well-documented. It is, therefore, completely expected that they should mostly bear Natufian-related ancestry. Nevertheless, just to confirm that these individuals are indeed of peninsular Arab origin, we shall run a final Vahaduo Multi analysis with all of our other Sudanese and Chadian “Arab” samples, but this time include the Levant_Tell_Qarassa_Early_Antiquity cohort to capture any such recent ancestry. We will also include the TUR_Marmara_Barcin_N and IRN_Ganj_Dareh_N specimens to see if our Rashaayda individuals and other samples carry any extra Anatolian Neolithic and Iran Neolithic admixture, respectively. The resulting analytical table below shows that, unlike the Kababish “Arab”, Baggara “Arab” and Daza Toubou samples, the Rashaayda Arabs clearly are of recent peninsular Arab origin. They are the only Arabic-speaking population of Sudan and Chad in our dataset that mostly belongs to the associated Levant_Tell_Qarassa_Early_Antiquity component (68.6% on average). Among the Kababish and Baggara, Arabian admixture is instead restricted to a handful of individuals (notably, the Chadian Baggara individual ABA032, who has an elevated 41.4% frequency of the Levant_Tell_Qarassa_Early_Antiquity element). These outliers are likely descendants of peninsular Arab Muslims, who introduced the Islamic faith and the Arabic language to the Sudan area during the medieval period.
Tables of ancestral proportions for each Arabic-speaking Sudanese and Chadian group as well as the Daza Toubou:
Step 8: Confirm whether modern Maghrebis also share this ancestral composition
As an eighth step in our analysis, we shall explore whether contemporary Afro-Asiatic speakers of the Maghreb region in northwestern Africa also share the ancestral composition described above. We will begin, as previously, by conducting a Vahaduo Distance test on all of Global25’s ancient African samples that possess the least Eurasian admixture. This should help us determine which population mainly contributed the Sub-Saharan African admixture that modern Maghrebis carry.
Of the top eight results listed above, the Congo Kindoki specimens are the most suitable Niger-Congo proxies for our Maghrebi samples. This is because, along with the Congo NgongoMbata 220BP cohort, they are the “purest” available ancient specimens bearing Niger-Congo-related ancestry (see Step #9 below on how we know that), and Bekada et al. (2015) have found that contemporary Maghrebis harbor such Yoruba-like admixture. However, since the COG_Kindoki_230BP:KIN004 sample belongs to the Indo-European-associated Y-DNA haplogroup R1b1 (cf. Wang et al. (2020), Table S10), we shall avoid using it. We will instead utilize COG_Kindoki_230BP:KIN002 as our reference sample, for it bears the E1b1a paternal clade common among modern Niger-Congo speakers.
Moving forward, we shall now perform a Vahaduo Single analysis on our Maghrebi samples. The non-African specimens listed on the Global25_PCA datasheet will, alongside Congo_Kindoki and the other “pure” ancient Sub-Saharan African samples, again serve as our Source populations. Such a test leverages Vahaduo Admixture JS’s processing capabilities, allowing the program to find for us the exact ancestries our Target populations carry.
From the above, we can see that Maghrebis do generally share a similar ancestral makeup as other Afro-Asiatic speakers in the Horn and Nile Valley. Here too we may note the now-familiar array of majority West Eurasian elements (ancient Egyptian, European-related Steppe, and Levantine Natufian components) and a minority East Eurasian element (East Asian component), with a low Sub-Saharan African admixture (primarily derived here from the ancient Niger-Congo sample Congo Kindoki 230BP, with ancillary gene flow from the ancient Nilo-Saharan sample Kakapel 300BP). However, a major difference between these populations is that the Iberomaurusian/Taforalt component forms a large portion of the ancestry of virtually all modern Maghrebi individuals, whereas this element is found at very low frequencies toward the east (typically under 5%). Anatolian Neolithic-related ancestry (represented by TUR_Marmara_Barcin_N) is also an important admixture element, particularly among coastal populations in the north, with Iran Neolithic-related admixture (represented by IRN_Ganj_Dareh_N) also present. Furthermore, one population, the Arabic-speaking Rbaya of Tunisia, seems to be descended from recent settlers from the Arabian peninsula rather than Arabized Berbers. These Rbaya individuals carry a predominant Natufian ancestry like many peninsular Arabs, and (notwithstanding Sub-Saharan African admixture) low percentages of said quintessential Maghrebi ancestral elements.
To better marshal the findings above and more easily observe the identified trends via table format, we shall conclude by running a Vahaduo Multi analysis on our Maghrebi Target populations:
Fregel et al. (2018) studied Late Neolithic individuals excavated at the Kelif el-Boroud site in Morocco, and report that these ancient specimens bore ancestry comprised of roughly equal Natufian and Anatolian Neolithic genome elements. Given this discovery, we must conduct a Vahaduo Multi test using Global25’s MAR_LN ancient sample as an additional Source population. This will help us determine whether the Anatolian Neolithic ancestry, which we have just observed above in our Maghrebi samples, was primarily 1) inherited from these Late Neolithic Moroccans, or 2) acquired later through absorption of peoples arriving from southern Europe or western Asia. From the data table below, it is apparent that scenario #2 is correct; most Maghrebi groups did not derive their Anatolian Neolithic admixture from the Late Neolithic specimens from Kelif el-Boroud. Contemporary Maghrebi individuals carry the MAR_LN component at a low average of 4.4%, with little change in average frequencies of the Anatolian Neolithic-related TUR_Marmara_Barcin_N component (23.4% vs. 25.8%). This is also evidenced by the fact that haplogroup T, the only paternal clade that Fregel et al. observed among their Late Neolithic Moroccan specimens, is rare among just about all modern Maghrebi groups. The latter populations instead largely belong to the E1b1b lineage, as do most Epipaleolithic Iberomaurusians, Mesolithic Natufians, Pre-Pottery Neolithic makers, Early Neolithic specimens exhumed from the Ifri n’Amr or Moussa site in Morocco, ancient Cushites of the Pastoral Neolithic, and ancient Egyptian individuals.
When Libyans are added to our Maghrebi dataset, they appear most similar to Muslim Egyptians inhabiting the Nile Valley; they also share appreciable ties with the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn. The Libyan individuals have the same basic ancestral makeup as these Muslim Afro-Asiatic speakers to their immediate east, carrying majority West Eurasian ancestries (viz. ancient Egyptian, European-related Steppe, and Natufian components) and a minority East Eurasian ancestry (East Asian component), as well as a bit of Sub-Saharan African admixture and a small North African Iberomaurusian/Taforalt admixture. However, Libyans (13.5%) have a higher average frequency of the Iberomaurusian component than Egyptians (2.9% among Muslims, 0% among Copts) and Horn populations (1.2%), though significantly lower than Maghrebis (28.5%). Like Muslim Egyptians and to a lesser extent Maghrebi groups, Libyans also sustained recent gene flow from the Arabian peninsula. This is evidenced by the presence of the Levant_Tell_Qarassa_Early_Antiquity component in our Libyan dataset, an ancestral element which again is typical of modern Arabic speakers.
Step 9: Repeat analytical steps above with a control population to ensure accuracy and replicability
So as to ensure the accuracy and replicability of our admixture analysis, we will broadly repeat the steps above with a control population. For this purpose, we will utilize peninsular Arab samples from the Global25_PCA_modern datasheet as our Target group, including Yemeni Jews and Mahra individuals.
To start, we shall aim to identify the “purest” Sub-Saharan African reference population available for our Arabian Target population. We will achieve this by first conducting a Distance analysis on the Vahaduo Admixture JS program, using as our Source populations the ancient African samples listed on the Global25_PCA datasheet (again excluding the aforementioned ancient African samples with substantial Eurasian ancestry). We then take note of the top eight African samples with whom our Target population shares the closest genetic ties. We are only interested in the top eight results because these are the groups that are most likely to have contributed genes to our Target population.
Next, we scour the existing genetic literature to find out which of these eight African proxy groups has the least documented non-African admixture. This necessary step will help us avoid depressing Eurasian ancestry/inflating Sub-Saharan African admixture in our Arabian cohort. According to Wang et al. (2020)’s admixture analysis, the COG_Kindoki_230BP:KIN002 sample from the Democratic Republic of the Congo has the least Eurasian admixture (red component):
We have thus found our “pure” Sub-Saharan African proxy sample for our examined Arabian individuals. This discovery informs us that the ancient contact population which contributed most of the Sub-Saharan African admixture in the Arabian peninsula (as represented by Congo Kindoki 230BP) was different from that which did the same in the Horn of Africa and Nile Valley (as represented by Kakapel 300BP) — a fact which is especially clear when we perform a two-way Vahaduo Multi analysis with peninsular Arab individuals, Afro-Asiatic speakers from the Horn, and Maghrebis, using the COG_Kindoki_230BP sample as our Sub-Saharan African Source population and the ancient Egyptian EGY_Late_Period sample as our ancient Eurasian Source population. The peninsular Arab samples wind up showing the highest average Niger-Congo ancestry in comparison to the other examined Afro-Asiatic-speaking groups, confirming their preference for this component as their main source of ancient Sub-Saharan African admixture:
Now, we will carry out a Vahaduo Single analysis, using as our Source populations all of the ancient non-African samples listed on the Global25_PCA datasheet, except those with non-trivial Sub-Saharan African admixture. This step identifies the Levant_Tell_Qarassa_Early_Antiquity sample as the predominant Eurasian ancestry borne by virtually all of the examined Arabian individuals (see here).
We will end by running a Vahaduo Multi analysis, using Levant_Tell_Qarassa_Early_Antiquity as our “pure” ancient non-African reference sample and Congo Kindoki 230BP as our “pure” ancient African reference sample. The analysis produces acceptable Distance fits of <9%, with a sensible estimated average African admixture of ~17%. We may also note that the Mahra samples from Yemen have the least Sub-Saharan African admixture. This is consistent with previous research, which has established that Mahra individuals are on average the “purest” living Semites, having retained the most ancient Natufian ancestry and the lowest extraneous genetic influences (cf. Vyas (2017)).
Conversely, when a Vahaduo Multi analysis is conducted using KEN_IA_Deloraine — the ancient African sample with whom our Arabian individuals showed the greatest genetic affinity in the Vahaduo Distance analysis above (except Emiratis, most of whom preferred instead the COG_NgongoMbata_220BP ancient African sample) — the Arabian individuals appear to have a more elevated Sub-Saharan African admixture of ~19% on average (see here). If we consult again Wang et al.’s genome analysis, it is clear why that is: the early Bantu sample from the Deloraine farm in Kenya harbors substantial Eurasian admixture (red component) specifically related to Arabians, which, compared to other ancient African samples, is bringing it genetically closer to the modern Arabian individuals. This again highlights the importance of using ancient African proxy groups that have as little Eurasian admixture as possible.
Step 10: Re-confirm our findings using other ancient Egyptian samples
As a penultimate step, we will repeat our Vahaduo Admixture JS analysis using other ancient Egyptian samples in lieu of the EGY_1879BCE cohort. Although all of our modern Afro-Asiatic-speaking samples from the Horn demonstrated a clear preference for EGY_1879BCE in the Vahaduo Single analysis in Step #3 above, EGY_1879BCE is not listed on Eurogenes’ official Global25_PCA datasheet. We must therefore re-confirm our findings, this time using Global25’s official ancient Egyptian samples.
To start, we shall carry out a Vahaduo Distance analysis on all of the Eurasian samples listed on the Global25_PCA datasheet. Doing so will help us identify which of these ancient specimens our modern Cushitic, Ethiosemitic and North Omotic-speaking individuals share the nearest affinity with. The Distance analysis indicates that our Afro-Asiatic speakers show a preference for the Levant_Beirut_IAIII_Egyptian:SFI-44 cohort followed by EGY_Late_Period:JK2134, which are ancient Egyptian samples dating from the Iron Age and later Dynastic epoch, respectively (see here).
We will now perform a Vahaduo Multi analysis, utilizing the Iron Age Levant_Beirut_IAIII_Egyptian:SFI-44 sample in place of the earlier Dynastic period EGY_1879BCE sample:
As can be seen in the table above, our Afro-Asiatic-speaking samples again wind up with almost the same average non-African ancestry (just over 70%), with approximately 27% Sub-Saharan African admixture and around 3% North African Iberomaurusian/Taforalt admixture. The average Distance fit is also similar. However, the apportionment of the Eurasian ancestries differs appreciably from before. The frequency of the ancient Egyptian component increases about 10 percentage points, going from an average of 18% to 28.1%. Additionally, the average frequency of the Levantine Natufian component rises over 10 percentage points, spiking from 20.9% to 31.3%. These boosts ultimately come at the expense of the European-related Steppe component and the East Asian component, which, respectively, drop from an average of 11.3% to 0% (~11 percentage points) and 22.6% to 12.6% (10 percentage points). Hence, the Steppe and East Asian components appear to be embedded within the Iron Age Levant_Beirut_IAIII_Egyptian cohort, representing constituent elements of that sample.
These changes inform us that:
1. The northern Egyptian population to which the earlier Dynastic period specimen EGY_1879BCE (i.e., the Middle Kingdom nobleman Nakht-Ankh) belonged had not yet interbred with the peoples who brought the European Steppe and East Asian-related ancestries to the Nile Valley, nor with the folks responsible for the Sub-Saharan African admixture element. This is also suggested by Vahaduo Multi analysis of ancient Egyptian individuals, which reveals that EGY_1879BCE only bore embedded Natufian (~81%) and Anatolian Neolithic ancestry (~19%):
2. By the time of the Iron Age Egyptian specimen Levant_Beirut_IAIII_Egyptian:SFI-44, these newcomers had largely been absorbed. From the cumulative data available, we may presume that this assimilation process was concentrated in northern Sudan and southern Egypt since the older Cushites of the Pastoral Neolithic — who seem to have originated from northern Sudan; see Wang et al. (2022) — already bore Steppe and East Asian components, as well as a minor Sub-Saharan African admixture element and a tiny Iberomaurusian admixture.
We will finish by conducting the same Vahaduo Multi analysis on the later Dynastic-era EGY_Late_Period:JK2134 sample, which the Afro-Asiatic speakers of the Horn also appear to favor. Predictably, the end result is virtually identical to the Levant_Beirut_IAIII_Egyptian:SFI-44 Multi analysis, with only incremental differences in Distance fit and ancestral component frequencies (see here). This apprises us that the Iron Age Egyptian profile actually dates earlier, to at least the later Dynastic period. It also lets us know that our interpretations vis-a-vis the EGY_1879BCE sample are correct.
When we perform the same confirmatory Vahaduo Multi tests on the Coptic Egyptian samples and northern Egyptian (Cairo and Mansoura) samples from IllustrativeDNA and the Muslim Egyptian samples from Eurogenes, alternately using the later Dynastic period EGY_Late_Period:JK2134 sample and Iron Age Levant_Beirut_IAIII_Egyptian:SFI-44 sample as our ancient Egyptian Source population in place of the earlier Dynastic period EGY_1879BCE cohort, the Coptic and northern Egyptian samples have ballooned Distance fits approaching 15%. By contrast, the general Muslim Egyptian samples maintain acceptable Distance fits of <9%. Hence, Coptic Egyptians and northern Egyptians indeed appear to derive most of their ancestry specifically from the earlier Dynastic period Egyptians, as represented by the EGY_1879BCE sample:
Step 11: Consolidate our findings for all local Afro-Asiatic speakers
Finally, to conclude our admixture study, we shall distil our findings by grouping all local Afro-Asiatic speakers (Horn African and North African alike) into one Vahaduo Multi table. This will allow us to more easily observe and describe broad trends in the data, and to spot any patterns or relationships that we may have overlooked.
From the data table above, we can discern three general ancestry patterns:
Besides the foregoing, we may also note that the Elmolo, a vestigial Cushitic-speaking group inhabiting Kenya, carry similar ancestral elements as the Horn’s Afro-Asiatic-speaking populations. They are distinguished by a substantially higher Sub-Saharan African admixture (~52% on average), which primarily consists of the “pure” ancient Nilo-Saharan element (Kakapel 300BP) followed by the “pure” ancient East African Hunter-Gatherer element (MWI_Chencherere). This is consistent with a heavy absorption of local Nilotic and forager individuals. Furthermore, the South Omotic-speaking Ari seem to derive the bulk of their ancestry from the Sub-Saharan African components (98%). This proportion is probably an overestimation since the Ari sample (taken from IllustrativeDNA) has a grossly unrealistic Distance fit of ~40%.
We will end by performing another Vahaduo Multi analysis on our Afro-Asiatic-speaking samples. This time, we shall employ the Levant_Tell_Qarassa_Early_Antiquity cohort as a Source population so as to quantify the amount of modern Arabian admixture present in our Target populations:
The data table above affirms that the Afro-Asiatic-speaking populations of the Horn, as well as Coptic Egyptians and northern Egyptians, do not bear any recent admixture from the Arabian peninsula. All of the examined Cushitic, Ethiosemitic and North Omotic-speaking individuals and Coptic and northern Egyptian individuals have a 0% frequency of the Levant_Tell_Qarassa_Early_Antiquity component, an ancestral element typical of contemporary peninsular Arabs. This suggests that most of the gene flow from Arabia into Northeast Africa predates the Islamic era. On the other hand, Muslim Egyptians and Libyans do harbor this Arabian component at low frequencies of around 17% and 19%, respectively. In the Maghreb, average frequencies of this Arabian element are a bit lower, except among the Arabic-speaking Rbaya of Tunisia. Rbaya individuals on average carry the Levant_Tell_Qarassa_Early_Antiquity component at an elevated percentage of 44%. This strongly supports their traditions of descent from Arab forefathers. Intriguingly, most persons from the Arabic-speaking Douz community in Tunisia also have high frequencies of this Arabian ancestral component, in keeping with their own traditions of descent from Arab settlers.
Conclusion
The admixture analysis above tells us a number of things about the biogenesis of the modern Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa. We may summarize these key findings as follows:
11 Saturday Mar 2023
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Abdel Monem A. H. Sayed, Adulis, AECR, Afro-Asiatic, Ahmed El-Batrawi, Ahmed Ibrahim Awale, Alula, Amelia Edwards, ancient DNA, Ancient Egypt, ancient inscriptions, Antiquities Service of Egypt, Arrian of Nicomedia, Ati, Auguste Mariette, Axumite Kingdom, Édouard Naville, Barbaria, Bia-Punt, Book of Gates, Boswellia frereana, Carleton Coon, Caterina Cozzolino, Caucasoid, Cushitic, D'mt, Dalbergia melanoxylon, David O’Connor, Deir el-Bahri, Dimitri Meeks, Diodorus Siculus, Egyptian pantheon, El-Osbolé, Emmett Sweeney, Epiphanius of Constantia, Eric Robson, Ernest-Théodore Hamy, Ernesto Schiaparelli, F. Nigel Hepper, Flinders Petrie, frankincense, G. Billy, G. W. B. Huntingford, Gaston Maspero, George A. Hoskins, George Andrew Reisner, Georges Révoil, Giuseppe Sergi, Grafton Elliot Smith, Grand Procession, Hamitic, Hathor, Hatshepsut, Heinrich Karl Brugsch, Horus, Hyphaene thebaica, James Bruce, Johannes Maria Hildebrandt, John Desmond Clark, Kathryn Bard, Kenneth Kitchen, Laas Geel, Lady of Punt, Land of Punt, Lionel Casson, Medri Bahri, Meroitic culture, Mersa Gawasis, myrrh, Nathaniel Dominy, Neville Chittick, Northeast Africa, Palermo Stone, Papio hamadryas, Perahu, Periplus of the Erythraean Sea, Precambrian, PROTA, Red Sea, Richard Pankhurst, Robert Bennett Bean, Robert Hoyland, Rodolfo Fattovich, Saww, Solomonid Dynasty, Stanley Balanda, stela, Ta netjer, Theodor von Heuglin, Toby Wilkinson, Wadi Gawasis, Zoscales
The Land of Punt was a major civilization in the ancient world. Located to the south and east of the Nile Valley, it was Dynastic Egypt’s main trading partner and the source of much of the frankincense, myrrh and other coveted products that were used by the Pharaohs in their traditional rituals and ceremonies. According to the Egyptian First Dynasty rulers or Horus-Kings, Punt (Ta netjer or “God’s Land”) was also their ancestral homeland.
Given its historical importance, Egyptologists have long debated just where exactly this mysterious territory was situated. The riddle appeared to have been finally solved in 2010, when preliminary isotopic analysis narrowed down the prospective locations to present-day Eritrea. However, follow-up isotopic work as well as DNA studies conducted since then, analysis of clay from pots that were brought to Egypt from Punt, and little-known botanical evidence and epigraphs firmly locate the ancient land in a broader region stretching from northern Somalia, Djibouti and the Eritrea/Ethiopia corridor to northeastern Sudan. The recent discovery of the first actual Puntite artifacts and their similarity to those of ancient Egypt has, in particular, confirmed the close ties between both areas.
Route to Punt
One of the key factors in pinpointing the location of the Land of Punt is its geographical proximity to ancient Egypt. In this regard, scholars have often indicated that the territory was situated to the south and east. But what exactly do they base this on?
In the 1850s, the Antiquities Service of Egypt discovered hieroglyphic texts in the vicinity of Thebes (Luxor) in Upper Egypt. These inscriptions identify Punt as a source of aromatics found to the east of Egypt. This, in turn, would prompt the Egyptologist Heinrich Karl Brugsch to postulate that the ancient land was located in the Arabian peninsula. A few years later, his colleague the archaeologist Auguste Mariette came upon geographical lists at the Karnak Temple, which had been left by the Pharaoh Thutmose III of the Eighteenth Dynasty (r. 1458–1425 BCE). These hieroglyphics include Punt among the territories that lay to the south of Egypt. The Egyptologist Abdel Monem A. H. Sayed explains:
The first lists of this kind, and the most comprehensive, are those of Thutmoses III, where the regional and site names are arranged in a manner coinciding with their geographical locations.
The list of toponyms of the African side of the Red Sea begins with the heading Kush, the Egyptian name for Upper Nubia. Under this heading are recorded 22 toponyms. Then comes the regional name Wawat or Lower Nubia, with 24 toponyms listed under it. After that, the list begins again from the south, recording regional and site names closer to the Red Sea shore. The regional name Punt is mentioned as a heading for 30 toponyms. After Punt comes Mejay as a heading of 17 toponyms. Lastly comes the regional name Khaskhet extending along the Red Sea shore of Egypt, with 22 toponyms listed (Schiaparelli 1916, 115-9).
This clear hieroglyphic account allows the following important deductions[…] The relation between Punt and the other regional names in the list (Kush, Wawat, Mejay and some of the toponyms under the heading Khaskhet), of which the African locations are agreed among Egyptologists, shows clearly that in the time of Thutmoses III Punt was the most southerly region and adjacent to the Red Sea coast. This is of great value for locating Punt during the New Kingdom in general, and the time of Queen Hatshepsut in particular, with which I deal later.
A 26th Dynasty stela was also recovered from the ancient site of Dafnah (Daphnae) near the Egyptian Delta, which contains an inscription stating that “when rain falls on the mountain of Punt, the Nile floods.” This is a clear allusion to the northern Ethiopian highlands, around Lake Tana where the Blue Nile rises (cf. Sayed (1989)). The Papyrus of Hunefer, a 19th Dynasty document by an Egyptian royal scribe, strengthens this association, for it too includes the Blue Nile within the confines of Punt. Van Auken (2011) notes:
The Greek historian Herodotus wrote, “Egypt was the gift of the Nile.” Three tributaries created this amazing river. The first is the White Nile, a long gentle river flowing from Lake Victoria — a high mountain lake bordered by Kenya, Tanzania, and Uganda. This tributary joins with the shorter but more voluminous and nutrient-richer Blue Nile, springing from Lake Tana in Ethiopia. And in ancient times, a third tributary joined these two, the Yellow Nile flowing from the eastern highlands of Chad. The Yellow Nile is now dry but was once a part of this trinity of rivers creating the ancient Nile.
Each year during the rainy season, the ancient Nile River would overflow its banks and inundate Egypt; as it retreated, it left behind nutrient-rich black silt that made Egypt one of the most fertile lands in recorded history. Today, two dams now control the Nile — there is no flooding and no rich silt fertilizer.
Around this river of life-giving water grew one of the greatest cultures on the planet. The ancient Egyptians called the river Iteru, meaning “Great River”; the modern name comes from the Greek Neilos (Nilus in Latin), transliterated to Nile.
The Edfu Text, found in the Horus Temple in Edfu, and the Papyrus of Hunefer tell the story of a “hill people” who became the first settlers of this region. “We came from the beginning of the Nile where god Hapi dwells at the foothills of the Mountains of the Moon.” The Mountains of the Moon are likely those that contain Lake Tana in ancient Abyssinia (modern day Ethiopia), the origin of the Blue Nile. This name may be traced back to a Greek ruler of late Egypt, Ptolemy, and the use of the name “Mountains of Selene,” the moon goddess of the Greeks.
Coupled with some of the floral and faunal evidence discussed below, Mariette’s discovery and the Dafnah tablet helped shift scholarly opinion as to where Punt was situated (including eventually that of Brugsch himself) away from a hypothesized Arabian location to the adjacent Horn of Africa.
The Palermo Stone contains the earliest hieroglyphic description of an ancient Egyptian expedition to the Land of Punt, during the Fifth Dynasty (Sci-News).
Cozzolino (1993) enumerates over 50 other hieroglyphic inscriptions relating to the Land of Punt, and a few additional engravings have subsequently been discovered. The earliest of these writings is the Palermo Stone. It informs us that an ancient Egyptian expedition to Punt brought back 80,000 measures of ‘ntiyw (a particular type of incense), among other items, during the thirteenth regnal year of Sahure (ca. 2445 BCE), the second Pharaoh of the Old Kingdom’s Fifth Dynasty. Unfortunately, the Palermo Stone does not specify where Punt itself was located. We do, however, have an idea of how Sahure’s men got there. In 2002, an inscribed block was found at the pyramid of Sahure in the Abusir necropolis, with two of its registers depicting the arrival of ancient Egyptian cargo vessels transporting goods from Punt. From this, we know that the Sahure expedition was carried out by sea rather than overland. Punt was therefore not a landlocked territory.
A Sixth Dynasty inscription provides an even clearer indication of the route that the ancient Egyptians took to get to Punt during the Old Kingdom. Phillips (1997) notes that Pharaoh Pepi II or Neferkare (2278/2269–2184/2175 BCE) dispatched one of his expedition leaders, Pepinakht, to retrieve the body of the official Anankhti, who had been killed by bedouins in the Eastern Desert (the “desert of the Asiatics”) while overseeing the construction of a ship intended for another commercial expedition to Punt. Thus, the particular water route that was favored by the ancient Egyptian rulers appears to have been via the Red Sea. This is confirmed by a later, Middle Kingdom rock inscription at Wadi Hammamat from the reign of Pharaoh Mentuhotep III or Senekhkere (r. 2004–1992 BCE) of the Eleventh Dynasty. According to the chief treasurer Henu, he was ordered by the king to build a vessel destined for Punt along the Red Sea littoral:
[My lord, life, prosperity] health[…] sent me to dispatch a ship to Punt to bring for him fresh myrrh from the sheiks over the Red Land, by reason of the fear of him in the highlands. Then I went forth from Koptos upon the road, which his majesty commanded me…
I went forth with an army of 3,000 men. I made the road a river, and the Red Land (desert) a stretch of field, for I gave a stretch of field, for I gave a leathern bottle, a carrying pole[…], 2 jars of water and 20 loaves to each one among them every day. The asses were laden with sandals[…]
Then I reached the (Red) Sea; then I made this ship, and I dispatched it with everything, when I had made for it a great oblation of cattle, bulls and ibexes.
Now, after my return from the (Red) Sea, I executed the command of his majesty, and I brought for him all the gifts, which I had found in the regions of God’s-Land. I returned through the ‘valley’ of Hammamat, I brought for him august blocks for statues belonging to the temple. Never was brought down the like thereof for the king’s court; never was done the like of this by any king’s-confidant sent out since the time of the god.
In 1971, the Egyptologist Kenneth Kitchen demonstrated the feasibility of such ancient travel down the Red Sea coast by charting an actual itinerary to get to Punt. This maritime gazetteer includes 80 possible anchorage points, as well as the intervening distances between them. It stretches from the Port of Sudan to northern Eritrea, a broad region that Kitchen suggests was coextensive with the Land of Punt.
So we know from the existing hieroglyphic texts that the ancient Egyptians preferred to reach Punt by water, and through the Red Sea specifically. We also know that this navigation was doable. The question is, what actual port did the ancient Egyptians use for these trading expeditions once they had finished constructing their vessels? A stela that was discovered at Wadi Gawasis (Wadi Gasus) provides an answer. Erected by Khentkhetwer, an official under the Twelfth Dynasty Pharaoh Amenemhet II or Nubkaure (r. 1922–1878 BCE), it contains an engraving that explicitly identifies Saww as the port where Khentkhetwer and his men arrived after their voyage to Punt. The Khentkhetwer stela inscription reads:
Giving divine praise and laudation to Horus[…], to Min of Coptos, by the hereditary prince, count, wearer of the royal seal, the master of the judgement-hall Khentkhetwer[…] after his arrival in safety from Punt; his army being with him, prosperous and healthy and his ships having landed at Sewew (Saww). Year 28.
Wooden cargo boxes labeled “wonderful things of Punt”, which were discovered at Mersa Gawasis, the ancient Egyptian port of Saww (Traveltoeat).
In 2004, archaeological excavations led by the Egyptologists Kathryn Bard and Rodolfo Fattovich at Mersa/Wadi Gawasis in Egypt identified this harbor as the old port of Saww, which the ancient Egyptians used during their expeditions to Punt. The excavators found a number of commodities at the site that may have been brought back from Punt, including fragments of carbonized ebony wood (Diospyros sp.) and obsidian. Since the latter volcanic glass does not occur naturally in Egypt, it was clearly imported from elsewhere. Lucarini et al. (2020) sought to identify the exact provenance of these artifacts, so they conducted a geochemical analysis comparing six obsidian fragments, which Sayed et al. had gathered from Mersa Gawasis some years prior, to those from various source areas in the Horn of Africa and Arabian peninsula (viz. sites in Eritrea, Ethiopia and Yemen). However, the scientists did not examine any obsidian culled from Djibouti or Somalia. Of the sites they did sample, Kusrale in Eritrea was found to be the most likely procurement location for five of the six analysed Mersa Gawasis obsidian artifacts. The other obsidian fragment appeared instead to have been obtained from the Dhamar Reda volcanic region in Yemen.
Obsidian artifacts recovered from excavations at Wadi/Mersa Gawasis, Egypt. Geochemical analysis conducted by Lucarini et al. (2020), which compared these fragments to other obsidian artifacts obtained from various source areas in the Horn of Africa and Arabian peninsula (viz. sites in Eritrea, Ethiopia and Yemen), revealed that Kusrale in Eritrea was the most likely procurement location for five of the six analysed Wadi/Mersa Gawasis obsidian artifacts. This supports the hypothesis that ancient Punt was centered in the Horn region (Lucarini et al. (2020)). (*N.B. The scientists did not examine obsidian fragments from Djibouti or Somalia. The conclusion they reach is therefore tentative.)
Moreover, the archaeologists working at Mersa Gawasis also discovered actual shipbuilding materials dating to the Middle Kingdom, such as anchors, timbers and huge steering oars, as well as 26 well-preserved coils of vessel-rigging rope that were lying on the floors of a cave. Most intriguingly, they came upon 43 cargo boxes from the reign of Pharaoh Amenemhat IV (r. 1990–1800 BCE). Two of these boxes were engraved with a package label, which had been recorded by the scribe Djedy. It included inscriptions for his name, a cartouche of Amenemhat IV and regnal Year 8, and the phrase “wonderful things of Punt” in hieroglyphics. The cargo boxes were made of sycamore wood and were all empty since their contents, which are believed to have included frankincense, were apparently unloaded into containers or bags for later transport via caravan across the Eastern Desert. Additionally, Bard found a limestone stela with hieroglyphic text on it that commemorates two royal maritime expeditions to Punt and Bia-Punt (“Mine(s) of Punt”) during the reign of Pharaoh Amenemhet III (r. 1831–1786 BCE).
Researchers exploring the ancient seafaring vessels at Wadi Gawasis would later find ceramic fragments inside. These sherds once formed pots, which were used to hold goods for transportation from Punt to Egypt. The scientists also analyzed the actual clay that was used to make this pottery. They discovered that it came from the eastern coast of Africa, further proving that this area was indeed part of the Land of Punt (Hoare (2020)). More specifically, of the main prospective locations for the ancient territory, this finding points to Djibouti, Eritrea and Somalia since Ethiopia is landlocked and Yemen is in the Arabian peninsula.
That the ancient Egyptians journeyed to and from Punt through a Red Sea route, and via the old port of Saww in particular, has thus been confirmed. What we shall now see is that the main landing point of these trading expeditions, at least during the New Kingdom, was in northern Somalia. As such, Punt was located in a more expansive area between Cape Guardafui and the Port of Sudan.
Flora and fauna of Punt
Mural from Pharaoh Hatshepsut’s temple showing the Puntites and Egyptians transporting frankincense trees from inland to the shore (Sayed (1989)).
Besides the route taken to get there, another key aspect in situating the Land of Punt is the flora and fauna of the various proposed locations for the ancient territory. Specific plants and animals, which are said to have been native to Punt, are depicted on Egyptian temple walls and murals. Some of these “wonderful things of Punt” were also brought back to Egypt as gifts and offerings. Combined, this leaves us with invaluable information as to what kind of habitat the Puntites actually lived in.
Plants
Primary distribution of the doum palm or Hyphaene thebaica in Africa (PROTA).
In 1858, Mariette discovered a wall in the funerary temple of the Pharaoh Hatshepsut (r. 1479–1458 BCE) at Deir el-Bahri, which depicts an Egyptian expedition to Punt during the queen’s reign. The temple reliefs show in detail the flora and fauna of Punt, as well as the Puntites themselves. Among the clearly identifiable plants are doum palms (Hyphaene thebaica), tree species that were regarded as sacred in ancient Egypt. Kitchen argues that on the Somali coast, the doum palm is restricted to the southernmost areas, far from the suggested Puntite nucleus in the north. In actuality, the doum palm grows throughout the Somali territories. The traditional gourd used by Somali pastoralists (who historically expanded from the north) is, in fact, crafted in part from doum palm fibers. Accordingly, the Plant Resources of Tropical Africa (PROTA) foundation describes the geographical distribution of Hyphaene thebaica as follows:
Hyphaene thebaica is distributed from Senegal and Gambia eastwards to Somalia, and is especially common between latitudes 8°N and 12°N. It also occurs in Libya, Egypt, Israel, the Arabian Peninsula and western India. Hyphaene thebaica is often planted. It was already cultivated in ancient Egypt, where it was considered sacred.
One of the main products that the ancient Egyptians traveled to Punt to obtain was ebony. Through hieroglyphic inscriptions, which indicate that the Egyptians themselves chopped down the plant while in Punt (“cutting ebony in great quantity”), ebony is known to have grown wild in the territory. The Puntites apparently did not import it from elsewhere for later resale to the Egyptians.
Barnett (1999) notes that analyses of plant specimens found in tombs have confirmed that the particular variety of ebony that was used in ancient Egypt is Dalbergia melanoxylon. This species is endemic to Eritrea, Ethiopia and Sudan alike, according to PROTA. In Somalia, ebony today has a limited distribution. Archaeological evidence, however, suggests that the plant was more abundant there too in Pharaonic times. On this likelihood, Ahmed Ibrahim Awale, a scholar who recently led excavations in northern Somalia (more on that below under Traces of ancient civilization), writes:
[Over] the past several millenia, so much has changed in the composition of vegetation in the Somali peninsula. The discovery of crocodile artifacts in Hargeisa valley suggest that a tropical riparian ecosystem existed in those areas. Therefore, it cannot be discounted that ebony, one of the chief exports from Punt, was sourced from the area. Diospyros spp., locally known as ‘Kolaati‘, is still found to a limited extent in riparian formations in Somalia.
Frankincense Terraces of Punt
The incense-yielding hills around Alula in northeastern Somalia, which correspond with the “Frankincense Terraces of Punt” (Sayed (1989)).
Of all the items that were exported from the Land of Punt to ancient Egypt, frankincense was by far the most important. Sayed (1989) notes that the Deir el-Bahri murals record Pharaoh Hatshepsut as specifically commanding her party “to fetch (as the texts say) ‘fresh incense’, and ‘frankincense living trees’ from ‘the frankincense terraces of Punt’.” The main purpose of that Egyptian voyage to Punt — the largest sojourn of its kind to the ancient territory — was, therefore, to retrieve the prized aromatic resin. Indeed, the very reason why Hatshepsut organized such a massive expedition was because she wanted her men to bring back live frankincense trees for later transplantation in Egypt. Her temple reliefs show that each of the 31 heavy incense trees required 4 to 6 men to transport them to the cargo ships, or 124 to 186 Egyptian and Puntite carriers in total. Since there were around 150 crewmen on the expedition’s five vessels (30 per ship), this would mean that the “Frankincense Terraces of Punt” had to have been situated near the shore.
View of Ras Filuk (Cape Elephas) from the Acannae/Akannai/Alula harbor (Sayed (1989)). According to the 1st century CE Periplus of the Erythraean Sea, Acannae is “where alone is produced the far-side frankincense, in great quantity and of the best grade.”
Sayed asserts that this incense-yielding locale, the “Frankincense Terraces of Punt,” was the northern Somali littoral. In particular, the northeastern section extending from Bandar Qasim to Alula. He bases this in part on historical texts, which hail this region as an early center of incense production and exportation. Chief among these old documents is the Periplus of the Erythraean Sea (Periplus Maris Erythraei), a travelogue written in the 1st century CE by an Alexandrian merchant. It indicates that a top-grade libanos peratikos or “incense from beyond the straits” (Bab el-Mandeb straits) was exported from ancient city-states that dotted this part of the Red Sea area, including Avalites, Malao, Mundus (Moundou), Mossylum, the Market and Cape of Spices, Pano (Panon), Opone (Opun) and Akannai (see toponym map below, on the left). The Periplus specifies that the laurel-grove of Akannai/Acannae is “where alone is produced the far-side frankincense, in great quantity and of the best grade.” Likewise, the historian and philosopher Arrian of Nicomedia testifies that the best frankincense of his day was exported from the same area around Ras Fiel/Ras Filuk (Cape Elephant/Cape Elephas), just off the Acannae harbor in present-day Alula.
In fact, various scholars hold that the classical market-town of Opone (Opun), located on the Ras Hafun peninsula in northeastern Somalia, is the etymological source of the toponym Punt. In this regard, G. W. B. Huntingford (1950) writes that “the name of the next cape south of Guardafui, Hafun, has descended through the Greek form Opone, from the ancient Punt.” Neville Chittick (1975) similarly asserts that “the possible early date of this [Hafun West] site would accord well with the relationship between the name Opone (Hafun) and the name Punt (pronounced Pwene?) suggested by Lewicki (1969, p. 38) citing F. Storbeck.” (*N.B. Also refer to Doresse in Lacroix (1998). For additional details, see What is the etymology of the term “Punt”?.)
Frankincense-growing areas on the northern Somali littoral and their hieroglyphic, classical and modern toponyms (Sayed (1989)).
According to Sayed, the ports of Malao, Mundus, Mosyllum and Akannai were the main hubs for incense exportation. He further notes that:
Comparing these locations with Hepper’s (op.cit. pl.XV) distribution of frankincense trees in north Somaliland today, we find an astonishingly close congruity. The areas where B. frereana (the best grade incense in Somaliland according to Hepper) grows [are] actually concentrated in those four locations, or at least — in the case of Malao — very near.
Sayed (1989) remarks that the ancient Egyptians imported two types of frankincense: a lower grade variety called sntr, and a higher grade variety known as ‘ntiyw or nty. According to inscriptions from the Sixth Dynasty travelers Harkhuf and Sebni, the lower grade sntr type was obtained from the Nile Valley or in Punt and was transported overland to Egypt. The higher grade ‘ntiyw incense was, on the other hand, exclusively acquired from Punt and was typically imported by sea.
Frankincense trees on the mountains overlooking Durba, northeastern Somalia (Chittick (1975)). These appear to be Boswelia carteri, a frankincense species that has been found within actual ancient Egyptian tombs (Lucas (1945)).
F. Nigel Hepper of the Royal Botanical Gardens reports that botanists have identified this ‘ntiyw variety with Boswellia frereana. Along with Boswellia carteri, he indicates that these are the incense types that are prevalent on the northern Somali littoral (cf. Sayed (2002)). Both species of frankincense have also been found in actual ancient Egyptian tombs (Lucas (1945)). This is pivotal since, according to Hepper, northern Somalia is the only area where Boswellia frereana grows in close proximity to the seashore, and on the requisite rocky hills to boot. Although such “terrace” land formations also occur in other parts of the Red Sea region, in Djibouti, Eritrea and Sudan, frankincense here grows instead at a minimum of 100 kilometers inland. The tree varieties that are found in this hinterland are likewise different from Boswellia frereana. Analogously, Chittick (1975) points out that a broad and arid plain lies between the sea and the steep mountains of the northwestern littoral of Somalia. It is therefore in the northeastern parts of the country, where the mountain ranges sit on the coast itself, that the most probable location of the “Frankincense Terraces of Punt” is to be found. Not coincidentally, this is also the very area where the ancient emporiums of Mossylum, Akkanai and Opone were all situated.
Animals
As with its flora, the fauna of Punt that is depicted on the Egyptian frescoes and described in hieroglyphic texts is native (though not entirely exclusive) to the Red Sea region. Among these animals is the giraffe, which today is only found in Africa. Superficially, this seems to rule out an Arabian location for the Land of Punt. A closer reading of the ancient testimonials, however, reveals that the giraffe was apparently also present in parts of the Arabian peninsula and Levant during the classical period. The ancient Greek historian Diodorus Siculus refers to the species as “camel-leopards”, and notes that it used to roam the area between northern Arabia and Syria (cf. Scott (2012)).
Depiction of an uncertain species of Puntite rhinoceros at the Hatshepsut temple (Meeks (2012)).
In an analogous vein, the representation of what may be a one-horned rhinoceros on the Hatsheptsut temple reliefs has been suggested as being indicative of an Indian location for Punt. This is because two-horned rhino species are today limited to Africa, whereas the one-horned rhino is restricted to the Eastern Himalayas. However, as Kenneth Kitchen (1971) observes, the rhinoceros figure in question, unlike the other animals depicted at Deir el-Bahri, is badly damaged, and this makes its identification difficult. Going by a similar “one-horned” rhino representation that was discovered at Kerma in Sudan, the depiction thus appears to have been an error. Kitchen writes:
That the beast seems only to have one horn (not two, as has the African rhino) is simply an error, analogous with that of one of the two Kerma representations of Middle Kingdom date (cf. Hilzheimer, ZÄS 67 (1931) 40), and with the stylized determinative of Louvre C. 14 accepted by Keimer (ASAÉ 48 [1948] 52 and fig. 5).
The archaeologist John Bimson also indicates that early Egyptian hieroglyphs included a pictogram of a one-horned rhinoceros. This, in turn, suggests that the species may have once inhabited the Nile Valley (cf. Sweeney (2006)). In short, the ultimate geographical origin of certain of the land-dwelling fauna of Punt is inconclusive.
Deir el-Bahri mural showing a Puntite village surrounded by Red Sea aquatic species, myrrh trees and other flora and fauna native to Northeast Africa (Edwards (1891)).
A rather different situation exists with the fish and other aquatic creatures that are illustrated on the same murals. Sayed (1989) points out that the Hatshepsut temple walls show marine species whose natural habitat is saltwater, including the lobster (palinurus). The body of water that is depicted therefore could not have been the freshwater Nile river, but instead more likely the Red Sea. Correspondingly, an analysis of the aquatic fauna on the Puntite temple reliefs by Eva Danelius and Heinz Steinitz found that the bulk of the specimens are indeed Red Sea varieties. As Kitchen (1971) notes, only a handful appear to be freshwater species, a fact which can be easily explained:
One factor largely discounted by Herzog (pp. 27, 55) is that of the fishes in the Deir el Bahri reliefs. These are, almost throughout, Red Sea/Indian Ocean fauna, with only two or perhaps three fresh-water species; see Eva Danelius and H. Steinitz, JEA 53 (1967) 15-24. If Hatshepsut’s expedition had reached Punt solely by travelling up the Nile, the overwhelming majority of Red Sea fishes is totally inexplicable. Why not solely Nile fauna, as in other Nile scenes? On the other hand, the Red Sea fauna fit a Red Sea route to Punt. The very few fresh-water fishes (a turtle; catfish, able to go in salt water, anyway; tilapia, dubious) could reflect the Nile part of the journey (Koptos-Thebes) or even fauna in Punt (River Baraka into Tokar Delta?), and pose an infinitely less problem.
A snippet of one of two secretary bird depictions on the Portico of Punt at Deir el-Bahri. The other bird figure is better preserved and displays the species’ distinctive head feathers, which allowed ornithologists to precisely identify it. Since the secretary bird is only found in Africa, and is a national emblem in Sudan, this finding supports a Northeast African location for Punt (Jadwiga Iwaszczuk).
Additionally, the walls of the Hatshepsut temple’s upper Portico of Punt feature bas-reliefs of a mysterious bird, which was among the animal species brought to Egypt from Punt. The bird was originally assumed to be a crane because only its rear could be seen. However, the Egyptologist Filip Taterka of the Polish Academy of Sciences found a well-preserved depiction of the same species on a nearby wall block. Thanks to its unique head feathers, Taterka was able to identify the animal as the secretary bird (Sagittarius serpentarius). This conclusion was also later confirmed by three ornithologists, who are experts on the secretary bird. Taterka’s finding is of particular importance vis-a-vis the debate on the whereabouts of Punt because the secretary bird only lives in open grassland in Africa. The species is endemic to Djibouti, Eritrea, Ethiopia, Somalia and Sudan alike; it actually serves as a national emblem in Sudan. According to Taterka, this fact strongly tips the weight of evidence in favor of a Northeast African location for Punt (cf. Foundation PAP). He states:
It was long thought that a bird that was difficult to identify was a crane, because at this spot the ornament was in poor condition—only the rump was visible.[…]
It is not without significance that today the secretary bird is the main element of the emblem of Sudan.
A Puntite man walking a Papio hamadryas baboon. Isotopic analysis of both bone and hair samples from baboon mummies that were brought to Egypt from Punt during the New Kingdom indicates that the specimens’ oxygen and strontium values most closely match those of baboons endemic to eastern Somalia and the Eritrea-Ethiopia corridor (Edwards (1891)).
The most definitive faunal data regarding where Punt was situated comes from baboons (Papio hamadryas). These are among the creatures that are depicted on the Hatshepsut temple walls at Deir el-Bahri, as well as on other ancient Egyptian murals. Baboon remains have also been found within actual tombs in the Valley of the Kings in Egypt. Moreover, in the Egyptian pantheon, the deity Thoth, who is associated with the moon cult, is often shown with the head of a baboon.
In 2010, a research unit led by Salima Ikram of the Egyptian Museum and Nathaniel Dominy and Gillian Leigh Moritz of the University of California analyzed hairs from two such mummified baboons, which had been kept at the British Museum. To determine the place of origin of the specimens, the scientists compared the baboons’ oxygen isotopic values with those of living baboon specimens from various hypothesized Puntite locations, including Eritrea, Ethiopia, Somalia and Yemen. Although the isotope data of one of the baboons was distorted, initial results from analysis of the other specimen indicated that its oxygen isotopic values matched closest with those of modern baboons from Eritrea and eastern Ethiopia. This prompted Dominy to posit that “Punt is a sort of circumscribed region that includes eastern Ethiopia and all of Eritrea”. He also suggested that the port of Massawa in Eritrea may have been the landing point of the ancient Egyptians’ expeditions to Punt since a baboon specimen from that harbor matched well with their ancient baboon mummy.
In 2015, the same Egyptian and American researchers conducted a more comprehensive isotopic study to confirm their preliminary findings. This time they compared both hair and bone samples, which they had extracted from two New Kingdom baboon mummies, with those of living baboons from the primary hypothesized locations of the Land of Punt. Analyzing both oxygen and strontium values, the scientists found that the closest matches were with specimens endemic to eastern Somalia and the Eritrea-Ethiopia corridor. They thus concluded that this area was the likeliest source of the baboons that were exported from Punt to Ancient Egypt:
The tandem origins of maritime trade and international diplomacy have roots in the Red Sea region. Graphic and epigraphic accounts of this trade often provide specific place names, or toponyms, with unambiguous geographic locations. Yet the location of one crucial polity, Punt (or Pwnt), remains uncertain. Punt was a major emporium of gold, electrum, and biological materials such as myrrh, ebony, ivory, short-horned cattle, leopards, and baboons (Papio hamadryas). The importance of these materials is reflected in the 1200-year duration of trade between Ancient Egypt and Punt (Vth-XXth Dynasties; ca. 2458-1163 BC). The recovery of mummified baboons from several New Kingdom tombs, which was a period of thriving trade with Punt, raises the possibility of using stable isotope analysis to source their provenience. Here we report the oxygen and strontium stable isotope composition of two P. hamadryas mummies from XXth Dynasty tombs. We also analyzed the hair and bone of modern baboons in 106 habitats spanning five hypothesized locations of Punt: (1) Eritrea-Ethiopia; (2) Mozambique; (3) Somalia; (4) western Uganda; and, (5) Yemen. Isoscapes based on kriging interpolation of hair keratin δ18O values and bioapatite 86Sr/88Sr ratios were produced and an index of similarity was calculated based on the geometric mean of the two kriged maps. Our results reveal a high likelihood match with eastern Somalia and the Eritrea-Ethiopia corridor, suggesting that this region was the source of Papio hamadryas exported to Ancient Egypt.
In 2023, Dominy co-authored an archaeogenetic study which compared the mitogenome of a Late Period (c. 800-540 BCE) baboon specimen retrieved from Gabbanat el-Qurud (“Valley of the Monkeys”), Egypt, with those of other baboons recovered from various hypothesized Puntite sites in Africa and the Arabian peninsula. The scientists observed that the Gabbanat el-Qurud baboon carried the G3-Y mtDNA haplogroup, a baboon mitochondrial subclade whose present-day distribution is focalized around Eritrea and eastern Sudan, including the Adulis vicinity. Along with other lines of evidence (discussed below) — which indicate that, centuries before the establishment of the Axumite kingdom, Adulis was a key part of ancient Punt — this discovery further reifies that the Red Sea area in Northeast Africa was indeed the location of the old Puntite civilization (cf. Grathwol et al. (2023)):
Adulis, located on the Red Sea coast in present-day Eritrea, was a bustling trading centre between the first and seventh centuries CE. Several classical geographers––Agatharchides of Cnidus, Pliny the Elder, Strabo––noted the value of Adulis to Greco-Roman Egypt, particularly as an emporium for living animals, including baboons (Papio spp.). Though fragmentary, these accounts predict the Adulite origins of mummified baboons in Ptolemaic catacombs, while inviting questions on the geoprovenance of older (Late Period) baboons recovered from Gabbanat el-Qurud (“Valley of the Monkeys”), Egypt. Dated to ca. 800–540 BCE, these animals could extend the antiquity of Egyptian-Adulite trade by as much as five mummified baboon from Gabbanat el-Qurud and 14 museum specimens with known centuries. To explore this possibility, we analysed complete mitochondrial genomes from a provenance together with published georeferenced mitochondrial sequence data. Phylogenetic assignment connects the mummified baboon to modern populations of Papio hamadryas in Eritrea and eastern Sudan. This result, assuming geographical stability of phylogenetic clades, suggests that present-day Eritrea, and by extension Adulis, was a source of baboons for Late Period Egyptians. It also establishes geographic continuity with baboons from the fabled Land of Punt (Dominy et al., 2020), giving weight to speculation that Punt and Adulis were essentially the same trading centres separated by a thousand years of history.
Other products of Punt
Gold
In various hieroglyphic texts, the ancient Egyptians refer to Punt by another name: Bia-Punt. This roughly translates as “Mine(s) of Punt”, which indicates the primacy of gold among the imports from the old territory. A Sixth Dynasty inscription belonging to the Pharaoh Pepi II is more explicit, as it demands “more than the mining region of Punt” (Sagrillo (2014)). Punt, or at least some of the districts under its control, was thus a center of gold production. This fact is of considerable value in helping to narrow down its location since gold was and is only mined in select areas around the world.
Geological formations in Northeast Africa. The old metamorphic (Precambrian) rocks are associated with gold-yielding areas. Eritrea is the only territory in Northeast Africa and the Arabian peninsula whose entire geological structure consists of these Precambrian formations. As such, it is the most probable location of the gold-mining district(s) of Punt, which is identified in hieroglyphics as Bia-Punt or the “Mine(s) of Punt” (Scientific American).
The first mention of the new toponym comes from the Sixth Dynasty traveler Harkhuf (ca. 2250 BCE). In inscriptions recorded at Aswan, this official asserts that he brought back products from “Bia-Punt.” According to Sayed (2002), this first reference to Bia-Punt appears to specifically concern the Nile area in northern Sudan. Sayed (1989, 2002), moreover, notes that the Harkhuf inscription is a clear allusion to gold, which the latter had presumably imported overland. In 1976-77, Sayed led a University of Alexandria expedition at Wadi Gawasis, where his archaeological team also found a Twelfth Dynasty stela “inscribed with a hieroglyphic text recording an order issued by King Sesostris I (Senusret I) to his vizier Antefoḳer to build ships to be sent to the region of Bia-Punt” (cf. Sayed (1978)).
Hence, Bia-Punt could be accessed through a water route. This implies that the region in question may have been coextensive with the Atbai desert in Sudan, which has long been a hub of gold mining. Other possibilities in the interior include the gold mines of western Ethiopia, as suggested by Eric Robson in his 2007 monograph In Search of Punt: Queen Hatshepsut’s Land of Marvels. Furthermore, Ibrahim (2013) reports that geologists have identified a zone in northwestern Somalia as potentially containing gold reserves. The Nubian Gold Corporation signed an agreement to prospect there, and the site has gold-quartz deposits with an estimated 13.5 gold parts per million. Since gold in Northeast Africa is associated with old metamorphic rocks — Precambrian geological formations that are found in all of these areas — Bia-Punt could conceivably have been situated anywhere within this traditional Puntite sphere. Eritrea would seem the most logical possibility, for it is the only territory in Northeast Africa and the Arabian peninsula whose entire geological structure consists of Precambrian rocks. On the other hand, Bressan (2013) notes that the geological formations of Yemen, Oman and other areas in Arabia are marked by recent sediments that are mostly bereft of gold. This makes the Arabian peninsula an unlikely location for Bia-Punt, the “Mine(s) of Punt.”
Slaves
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=Hamitic-type Nubians, Nilote (center-right).
In addition to gold, slaves were among the main exports that the Puntites sent to their ancient Egyptian trading partners. Most of these captives appear to have been of “Negroid” ancestral stock. This is suggested by the Grand Procession mural in Thebes at the tomb of Rekhmire, a vizier under Pharaoh Thutmose III. The painting contains several registers or levels, which depict various foreign envoys submitting tribute to the ancient Egyptians, including Puntite, Cretan and Nubian emissaries. The Grand Procession’s uppermost register shows a couple of reddish-skinned, orthognathous Puntite delegates, who strongly resemble the mural’s similarly copper-toned Egyptian, Cretan and Hamitic-type Nubian figures. These Puntites are flanked to the right by jet black-skinned, prognathous men, who, except for their attire, are physically identical to the Nilotes in the Nubian panel. As Kitchen puts it, “beside the so called ‘Hamitic’ type (like Parahu) 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 an Egyptian “Hamitic” type with a Nilotic “Negroid” type (Richardson (2003)).
Book of Gates mural in the Tomb of Seti I. From bottom left, clockwise: Reth (Ancient Egyptians, portrayed as copper-brown like the Puntites, Cretans and “red” Nubians); Aamu (Levantines, inhabitants of the Eastern Desert, portrayed as yellow-brown); Nehesu (Nilotes, the “black” Nubians); Temehu (Ancient Libyan group of European origin, portrayed as white; the original carriers of the paternal haplogroup R1b, they arrived directly from Europe (or indirectly from Central Asia) and were assimilated into the Nile Valley’s Afro-Asiatic-speaking populations (cf. Grugni et al. (2019))).
![Closeup of the original single panel in the Tomb of Seti I. The mural depicts (from left) “black” Nubian, copper-brown Egyptian, yellow-brown Levantine, and white Temehu figures. (*N.B. This is supported by archaeogenetic analysis. Haber et al. (2017) note that Bronze Age Levantine individuals from Sidon generally looked like their modern descendants, but had a darker complexion: "SNPs associated with phenotypic traits show that Sidon_BA and the Lebanese had comparable skin, hair, and eye colors[...] but with Sidon_BA probably having darker skin than Lebanese today from variants in SLC45A2 resulting in darker pigmentation".)](https://landofpunt.files.wordpress.com/2016/02/tomb-of-seti-i-single-panel-original-mural.jpg?w=300)
Closeup of the top three registers on the Grand Procession mural. The “Hamitic” Puntite figures on the top row resemble the copper-toned Cretans and Hamitic-type Nubians, and correlate with the “red” peoples described in the early inscriptions. The “Negroid” figures to the right of the Puntites are identical to the very dark Nilote on the bottom panel, and are in all likelihood the “black” peoples of the engravings (Hoskins (1835)).
Dimitri Meeks of the French National Centre for Scientific Research, like many other Egyptologists, believes that the conspicuously Nilotic folks on the Grand Procession represent the Puntites’ slaves. As such, they would have been accompanying their masters on this tributary voyage. The Egyptologist Stanley Balanda, however, argues that the “Negroid” individuals may instead have been diplomatic envoys too because “both races are being represented in the same way without distinctions being made to indicate any social or legal differences”. By his reckoning, the Nilotic figures would therefore have also inhabited some districts of Punt alongside the Egyptian-related Puntites. Meeks’ position on the slave origin of the “Negroid” individuals that are standing near the Puntites is compatible with an Arabian location for Punt (a geographic theory of which he is one of the main proponents). Balanda’s postulation is not, though, since there appear to be no indigenous populations in the Arabian peninsula, relict or otherwise, that have a similar phenotype (cf. Coon (1939)). On the other hand, both arguments are in line with a Northeast African locus for the Land of Punt.
Distribution of global population groups. The “Hamitic” Caucasoids comprise Afro-Asiatic speakers (Berber, Cushitic and Egyptian branches), ancestrally related peoples who inhabit North Africa and the Horn of Africa.
“Negroid” groups — Nilotic and Bantu populations known in Djibouti and Somalia as jareer or adoon and in Eritrea and Ethiopia as shanqilla or barya (terms denoting “negro”) — have long constituted the bulk of the slave class in the Horn. In antiquity, most of these slaves were captured from the surrounding areas bordering South Sudan and the Great Lakes region. However, there is textual evidence pointing to an early presence of two separate ancestral stocks in Northeast Africa; one with “Hamitic” affinities, and the other of apparent “Negroid” origin. (*N.B. On a population level, the Afro-Asiatic speakers of the Horn generally do not intermarry with jareer/barya (“negroes”), whether at home or abroad. This is due to traditional beauty standards, which place a low value on “Negroid” features. These aesthetic preferences are how the Afro-Asiatic-speaking populations have managed to maintain their distinctive physiognomies to a remarkable degree, despite being flanked by Niger-Congo/Nilo-Saharan/Khoisan communities with markedly different phenotypes. The Afro-Asiatic speakers’ reluctance to interbreed with “negro” groups also stems from a desire to preserve their own ancestral heritage (cf. Mwakikagile (2009), Gwyn (2013), Folklore Institute (2003)). Correspondingly, ancient DNA analysis has found that the early Cushitic settlers of East Africa were of North African ancestral stock, and that their modern descendants in the Horn share close affinities with them; see Ancient DNA from Ethiopia.)
This racial dichotomy was reported as far back as the kingdom of D’mt/Da’mat in Eritrea and northern Ethiopia, which flourished around 3,000 years ago during the pre-Axumite period. According to Robert Hoyland, who conducted archaeological excavations in the area in 2013, inscriptions attributed to the kings of that polity describe its rulers as lords of “Da’mat, its east and its west, its Sabaeans and its immigrants, its red people and its black people”. Tibebu (1995) likewise notes that “the distinction between the saba qayh (red men) and tsalim barya (dark slave) was also made during the Aksumite period”.
In that regard, in keeping with the Grand Procession and other ancient Egyptian murals, the Asiatic Society (1968) reports that epigraphs belonging to the Axumite King Ezana differentiate between “red” Nubians and “black” Nubians; the former correspond with Hamito-Cushitic peoples related to the Egyptians, and the latter are associated with Nilotes:
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.
A Somali queen, recalling the ancient Puntite figures depicted on the Deir el-Bahri and Thebes murals (French-Sheldon (1892)).
Thanks to the medieval chronicle of the Ethiopian Emperor Susenyos I, we are able to precisely identify the descendants of the “red” and “black” populations in the Horn. The royal court historian refers to the Afro-Asiatic-speaking non-slave populations as qayh (red) and the “shanqilla” groups as tsalim (black) (cf. Pankhurst (1976)). In short, whether the “Negroid” figures that are standing beside the Puntites on the Grand Procession mural represent emissaries or slaves, either possibility is consistent with a Northeast African center for the Land of Punt. This is because there was both an early presence in the area of distinct “Hamitic” and “Negroid” populations (as depicted on said fresco), and a well-established slavery tradition involving Nilotic captives. (For further details, see Who were the ancient Nubians?, If Nubians are ancestrally related to Egyptians, why do they now speak a Nilo-Saharan language instead of an Afro-Asiatic language?, and Are Sudanese “Arabs” really peninsular Arabs?.)
A slave of the queen, recalling the ancient “Negro” figures portrayed on the Grand Procession at Thebes (French-Sheldon (1892)).
Besides the foregoing, other ancient Egyptian inscriptions indicate that the Puntites also exported a number of Pygmy slaves. Harkhuf writes that during the second regnal year of the Pharaoh Pepi II (r. 2278–2184 BCE), he brought back a dancing pygmy as a royal gift for the child-king, which he had acquired from Punt by way of the kingdom of Yam in the Sudanese region. The traveler also avers that another pygmy had been imported from Punt at an earlier date, during the reign of Pharaoh Djedkare-Isesi of the Fifth Dynasty (r. 2414–2375 BCE). This exotic presence of Pygmy individuals in the Nile Valley is exemplified by Bes, the dward-god of ancient Egypt. According to the Egyptologist Amelia Edwards, the diminutive Bes and the goddess Hathor were “two of the principal deities worshipped by the Egyptians [who] had their divine origin in Punt” (see below for further discussion). Hart (2005), moreover, writes that the name Bes was also invoked by the ancient Egyptians as a catch-all term for their various dwarf-deities, who were unprepossessing in appearance but otherwise harmless:
The Egyptians had a number of monstrously formed dward-deities for which the name Bes was employed. The god in a plumed crown was normally bearded with his broad face surrounded by a lion’s mane and ears. His tongue often protruded in a playfully aggressive fashion. The body of Bes represented a band-legged dwarf wearing either a panther skin or a kilt, more Syrian than Egyptian, and a lion’s tail. Imitating for magical purposes the solar iconography of the god Horus of Behdet, Bes can be shown with his arms stretched out with a hawk’s wings suspended from them. The ‘sa’ sign meaning ‘protection’ is frequently carried by Bes.[…] He has his most crucial contribution to make to Egyptian life in his role as protector of childbirth in partnership with Taweret.
This begs the question, from where did the Puntites capture the Pygmy slaves to begin with? Perhaps somewhere in the central African hinterland, where they in fact have long been concentrated. Based on recent archaeological finds in an inland valley in northwestern Somalia (discussed below), Ibrahim (2013) argues that the Puntites had intermittently ventured even further in the interior, far beyond the boundaries of Punt and toward the Congo Basin and the gold mines of Mashonaland. There, they would have accessed slaves, additional gold reserves, and other commodities in the trans-regional trade networks for later barter with the ancient Egyptians. It would appear that such mercantile contacts persisted for centuries, as Leo Africanus testifies that the Sultanate of Mogadishu (Magadazo) originally controlled the gold trade at Sofala in present-day Mozambique (“this golden trade [at Sofala] was first in the power of the Moores of Magadazo”).
More parsimoniously, Lunde and Porter (2004) report that the so-called “Pygmies” brought back to Egypt might instead have been natural dwarves. This raises the possibility that these captives were simply short-statured individuals, and thus, plausibly drawn from the native hunter-gatherer communities of the Horn (as represented by the diminutive Chabo, Eyle and Ribi foragers) or Great Lakes regions (as represented by the small Hadza and Sandawe foragers). In support of these scenarios, Scozzari et al. (1999) observed a moderately high incidence of B-M60 haplotypes among individuals in northern and southern Egypt; Scozzari et al. (2014) also report that ~28% of Berbers in Egypt’s Siwa Oasis, a medieval hub of the slave trade, carry the B2a subclade (Table S7). However, this paternal lineage has not been found in ancient samples from either Egypt or Sudan (cf. this essay; Ancient DNA from Sudan). It instead today occurs at greatest frequencies among both the Hadza and Mbuti hunter-gatherers of southeastern and central Africa. Furthermore, genome analysis of modern Afro-Asiatic speakers in the Horn of Africa indicates that these individuals bear a predominant non-African ancestry (over 70%), with minor Sub-Saharan African admixture (~27%) that was in part derived from ancient East African foragers. This East African hunter-gatherer component is, however, rare among the contemporary Egyptian and Libyan samples (see Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa). Ergo, the presence of haplogroup B in Egypt is indeed likely a legacy of the captured “Pygmies,” who — considering the absence of this clade from the Nile Valley’s archaeogenetic record — seem to have been imported in significant numbers only after the Graeco-Roman period.
Bee hive-shaped houses
A mural at Deir el-Bahri shows a Puntite village next to a body of water containing Red Sea aquatic species (see above). This settlement consists of domes or bee hive-shaped huts, which are raised above ground on poles and accessible by ladder. Junker (1921) remarks that “Negroes” and mixed peoples, who are distinct from the “Hamitic” Puntites, can be seen standing near these houses:
It is significant that the first encounters with the Negroes of which we have any knowledge occurred through two quite different channels. When Queen Hatshepsowet, in the ninth year of her reign, i.e., 1482 b.C., sent a great fleet to the land of Punt, which included roughly the regions of Italian Eritrea and of British Somaliland, the expedition there met with genuine Negroes as well as half-breeds among the Hamitic Puntites. In the famous pictures of the rock temple at Der el-Bahri we see them moving about among the round huts built on piles. Whether they lived in a state of subjection to the real inhabitants of the country, or had settled beside them in the harbour district, or had merely come from the hinterland for purposes of trade, are matters of conjecture.
Balanda (2005) further states that, in Northeast Africa, such pile-dwellings are today restricted to the Nile Valley itself, for “houses on stilts were common in places subject to seasonal flooding such as regions inhabited by the Dinka and other tribes of the Sudan.” He argues that this particular scene, therefore, appears to depict a northerly part of the Land of Punt situated closer to Egypt, as opposed to the more distant sections of Punt which are clearly located in the Horn (namely, the Frankincense Terraces of Punt, the Mine(s) of Punt or Bia-Punt, and the Mountain of Punt). Kitchen (1971) reaches a similar conclusion:
Here, Herzog concentrates on pile-dwellings, one of the unique features of the Deir el Bahri scenes. While structures of this sort are known (past or present) in some form in many parts of the world, the only areas that prove relevant are S. Sudan and E. Africa. At present, those in the Sudan are between 12° N. and the Uganda border. There are some slender indications (tribal traditions; negritic skeletal remains) for the relevant Nilotic peoples having once lived further north than now – perhaps within part of Herzog’s Punt. He would dismiss Somaliland as unsuited and lacking any tradition or evidence.
Balanda and Kitchen are evidently right since their explanation accounts for both the uniquely built homes and the “Negro” figures portrayed at Deir el-Bahri. Moreover, it is supported by genetic research. Balemi (2018) reports that 55.55% of the Nilo-Saharan-speaking Berta individuals that he analysed bear the Eurasian F-M89 haplogroup. In Northeast Africa, the earliest instance of this Y-DNA lineage has been observed among Christian-period specimens entombed at sites on Meroë Island and the Nile’s 4th cataract (Yousif and Eltayeb (2009)). This is significant because it reveals that the Berta of the Ethiopia-Sudan border area — whom Koettlitz (1900) notes still build stilted huts very similar to those shown on the Deir el-Bahri mural — in the past had contact with and absorbed Eurasian-affiliated peoples (i.e., the Puntites). These are the exact conditions required to explicate the presence of both “Negroes” and mixed “Negro Puntite” folks dwelling near the “Hamitic Puntites” (or Puntites proper).
(*N.B. As of 2023, on the continent of Africa as a whole, the oldest instances of the Eurasian Y-DNA haplogroup F-M89 have been detected among Early Neolithic individuals excavated at Kahf Taht Al Ghar, an archaeological site located in the Maghreb, Northwest Africa. Genome analysis of these ancient specimens, who were agriculturalists, indicates that they bear European-related Anatolian Neolithic ancestry. By contrast, the medieval Kulubnarti individuals buried in Sudan mostly carry Levantine ancestry (which includes ancestry derived from the Middle Neolithic pastoralists of Skhirat, northern Morocco) as well as European-related Steppe ancestry and East Asian ancestry. Ergo, there are two separate ancient population sources that ultimately could have transmitted the Eurasian F-M89 clade, which many Berta individuals now harbor. The Kulubnarti Nubians appear to be the more probable candidate group considering their geographical proximity to the Berta. For our genome analysis of these medieval Kulubnarti Nubian specimens, see this. Also refer to here for a fuller discussion on the Middle Neolithic Skhirat individuals. The latter ancient pastoralists seem to be the ancestral population from whom the modern Cushitic, Ethiosemitic and North Omotic-speaking groups of the Horn and the contemporary Nubians of the Nile Valley inherited much of the Levantine ancestry they carry.)
Puntite short-sword
Another wall-painting found at the Pharaoh Hatshepsut’s temple in Deir el-Bahri depicts the Puntite chief Parahu wearing a sheath around his waist, which contains a particular kind of short-sword. This dagger is characterized by a straight double-edged blade design, topped by a hilt in the shape of a trident. According to Grottanelli (1947), among modern populations, only Somalis have traditionally manufactured a short-sword of this unique type.
Known as the bilao or bilawa, the dagger is evidently of great antiquity since old rock engravings featuring it have been observed at various archaeological sites in the Horn region, including Bur Eibe (Bur Eibi) in southern Somalia. The weapon figures at some of these locales are believed to have been carved at a time period more recent than the Puntite era. However, the short-swords portrayed at Bur Eibe are ancient, of sufficiently remote age to have been wielded by Parahu and his constituents in Punt.
Grottanelli (1947) writes:
The Somali[…] have long had, and still retain, a typical cutting weapon, a straight bladed dagger of a design peculiar to them alone (fig. I). That this weapon is traditionally old among them, is proved by the ancient Somali rock engravings discovered by Graziosi at Bur Eibi (fig. II), which faithfully reproduce the unmistakable shape of their hilts and straight double edged blades. The antiquity of this type of dagger would prove even greater if it were possible to identify it, as Revoil maintained, with that worn at the belt by Punt chiefs as depicted in the Deir-el-Bahri wall paintings. Be this as it may, it is a fact that the Somali departed so unwillingly from this particular type, that when they started forging swords for themselves they remained faithful to the same model, merely making the blade longer (fig. III). They adopted the Semitic word for “sword” (sayf or sef), but Reinisch’s texts show that they often continue to call this[…] bilawa or bilao, i.e. “dagger.” To this day, swords and sabres are very rare among them, as is the case with most Cushitic tribes.
Puntite leg-rings
On one of the wall panels at the Hatshepsut temple in Deir el-Bahri, the Puntite chief Parahu is shown with a series of yellow-colored rings adorning his right leg up to the middle of his calf. These seem to be leg bracelets, which are either made of gold or some other gold-plated metal. Although modern Cushitic-speaking males generally do not wear such leg rings, both ancient rock art and traditional accounts suggest that this was indeed once a custom of their Cushitic ancestors.
Hussein (2023) points out that an old cave painting at Laas Geel, an archaeological site in northwestern Somalia, depicts a pastoralist man wearing what appear to be stacked rings around one of his legs. These accoutrements are similar to the aforementioned leg bangles, which can be seen on the Puntite ruler Parahu. Describing this rock art, Gutherz and Jallot (2011) state:
The colours used for the human figures are the same as those used for the cows and the execution makes use of the same techniques. The thorax is often the only part which has resisted time. It is covered by a white or red ochre wide tunic, adorned with vertical stripes. The filiform arms outstretched in a gesture suggesting adoration, sometimes end with hands in the form of a fan. The legs are held together and clothed with “baggy” trousers. Rings possibly adorn the ankles, but there are no feet.
An oral history, as recounted to colonial administrators by a Karimojong Nilote elder in British East Africa, likewise asserts that in the later 1800s, a distinct pastoralist community used to inhabit parts of the Great Lakes region now occupied by Bantu/Nilotic/forager peoples. These herders have since been assimilated into the Oropom, a local hunter-gatherer community. Evidently, the vanished pastoralists were Cushitic peoples because they are described as having been “very light-skinned” with “pronounced slant eyes,” and their “men wore their hair long in a pigtail, like Indian women” — Cushites being the only folks in eastern Africa who have many members that adhere to a pastoral lifestyle and have a light complexion, oblique eye folds and lank hair. With regard to where ancient Punt was located, what is especially interesting about this testimony is the revelation that “both their men and women wore many bangles, sometimes stretching from their ankles to their knees and from their wrists to their armpits” (Wilson (1970)).
From the above account as well as the Laas Geel cave painting we may glean that wearing leg rings was a longstanding custom among Cushitic peoples. This tradition, though now essentially extinct, seems to have persisted to at least the 19th century.
Henna
During the Hatshepsut expedition to Punt, one of the key products that her party brought back with it to Egypt was henna (Lawsonia inermis) (IPA (1967)). A mural at Deir el-Bahri (shown to the right) depicts this queen’s men loading the plant onto their vessels alongside other cargo. The ancient Egyptians required the henna for cosmetic, medicinal and ritualistic purposes. They utilized it for personal beautification, as a fragrance to perfume their body oils, lotions and ointments, and also as an ingredient in the embalming process. Likewise, ancient texts convey that the early Greeks made similar use of henna. The physician Dioscorides of Anazarbus (c. 40 CE – 90 CE) refers to the plant as cyprus. This descriptor is reportedly the origin of the toponym Cyprus, as the ancient Cyprians once held a monopoly over trade in henna (Churchill (1875)).
Mural at Deir el-Bahri depicting ancient Egyptian men loading henna onto vessels for transportation from Punt to Egypt. Henna was and is ubiquitous in Egypt, where traditionally it has been used for cosmetic, medicinal and ritualistic purposes (Dreamstime).
In Egypt, actual mummies have been found with traces of the dye on their hands, feet, nails and hair. Grafton Elliot Smith, for instance, states that the 18th Dynasty official Henttawi had his hair dyed a bright reddish color. This practice seems to date to at least the predynastic era, for Guy Brunton writes that a Badarian old woman apparently had henna-dyed hair of a light brown-red hue (Lucas and Harris (1999)). (*N.B. Brothwell and Spearman (1963) also observed authentically blond ancient Egyptian individuals; their hair samples were not affected by either hair dye or cuticular damage.)
The custom of using henna to decorate the body was, therefore, of some importance in ancient Egypt. It is also still adhered to by modern Egyptians. Moreover, henna is a cultural staple of the Afro-Asiatic-speaking populations in the Horn of Africa. Among Somalis, Afars, Oromos, Saho, Beja, Bilen, Tigre, Jebertis, Argobbas and Hararis, the dye is used on a weekly basis for cosmetic adornment. More elaborate designs are reserved for special occasions such as wedding ceremonies, Eid celebrations and birthdays. Although usage of henna in the Horn is closely linked with Islam, the plant’s exportation from Punt suggests that, as in Egypt, this tradition is actually a holdover from remote times.
A Somali woman with henna designs on her arms. Henna application is an ancient custom among the Afro-Asiatic-speaking populations in Northeast Africa, a tradition which apparently dates to the Puntite era.
Mummified hand of a 19th Dynasty ancient Egyptian princess from Thebes. The appendage has darkened due to necrosis. However, it can be seen that the individual’s fingernails were painted red with henna.
Kohl
Kohl is another one of the “wonderful things of Punt” that were brought back to Egypt. The ancient Egyptians usually mined malachite and galena (lead sulphide) — the main elements used to make, respectively, the earlier green and later black varieties of kohl — at Sinai and the Eastern Desert, as well as in various areas in Upper Egypt. However, they would sometimes also import these cosmetic ingredients from the Arabian peninsula through Punt in the Horn region (Hardy et al. (2006)). During the New Kingdom, the Egyptians would procure kohl itself directly from Punt (Bard and Fattovich (2018)).
In ancient Egypt, kohl was liberally applied as eye-paint on both men and women. This has been gleaned from analysis of the contents of actual kohl pots and other cosmetic vessels, which were used to store the eye-paint. A dark residue has often been discovered on the inside of these containers (cf. Kalloniatis (2019)). Hassan and Hassan (1981) describe the creation process and storage of galena-based kohl by the predynastic Egyptians as follows:
The galena was ground on a cosmetic palette and applied as black eyeliner, equivalent to the kohl still widely used in Egypt. The ground galena was mixed with water and natural resins (Lucas 1948). The prepared galena paste was kept in shells, reeds, vessels, or wrapped in leaves. Another pigment used for eyelining in Predynastic and Dynastic Egypt is malachite. Sometimes both pigments were found together. Lucas (1948) noted that the black eye pigments used in ancient Egypt consisted of galena in 40 of 61 cases.
The receptacles used for storing kohl later evolved to small containers and eventually to tubes, pots and spoons during the dynastic era, with little sticks to apply the cosmetic to the eyes. Besides its obvious aesthetic function, scholars have proposed that in ancient Egypt, kohl may have also served a ritualistic or protective purpose insofar as its use was an attempt to symbolize the Eye of Horus (wadjet) (Mendoza (2017)). Likewise, the C-Group pastoralists of Lower Nubia, whom Oric Bates and George Andrew Reisner have identified as ancient Libyans, were known to apply the kohl eye-paint (E. S. Thomas (1926) notes: “the “C” Group people dressed in skins ; they used black eye paint and tatued their bodies, proving that they had light skins.” For additional details on this, see I have noticed that many rural Berber women have traditional face tattoos. Is this also common among other Afro-Asiatic speakers? And where did this practice originate?).
Ancient Egyptian figures depicted with the ubiquitous kohl eye-paint, from the Nebamun Tomb-Chapel c. 1350 BCE (British Museum).
Painted limestone statue of the ancient Egyptian Prince Rahotep, from the Mastaba of Rahotep at Meidum, Lower Egypt. This sculpture dates from the Old Kingdom’s 4th Dynasty (c. 2613-2498 BCE), showing the early usage of the kohl eyeliner in the Nile Valley.
The omnipresence of kohl in old Egypt is clearly reflected in dynastic period artwork. Monument walls and temple murals, such as at the Nebamun Tomb-Chapel, routinely show Egyptian figures wearing the eyeliner. Statues are likewise often depicted with painted eyes. This fact is especially interesting given that some ancient Puntite sculptures — which were recently excavated in northern Somalia; see discussion below — similarly feature contouring of the eyes, as if with kohl. Besides visual art, kohl is mentioned in the ancient Egyptian fable the Tale of the Shipwrecked Sailor. This story describes an encounter between a stranded Egyptian merchant and a talking snake, the latter of whom calls himself the “Prince of Punt.” The serpent offers to the trader a number of gifts to take back to Egypt, including kohl (Westling (1999)).
As with henna, kohl remains today in wide usage among Egyptians. The eye-paint is also a traditional fixture among the Afro-Asiatic-speaking populations in the Horn of Africa. This serves as yet another compelling reason why the ancient Land of Punt was almost certainly located in Northeast Africa.
A Somali woman with kohl eyes (indha kuul). Kohl is a commonly applied eye-paint among the Afro-Asiatic-speaking populations in Northeast Africa. The cosmetic’s usage dates from antiquity, for it was one of the “wonderful things of Punt” that were imported to ancient Egypt during the New Kingdom.
Appellatives
Hieroglyphic signs for brbrta, the primary ancient Egyptian ethnonym for the Puntites (AECR (1976)).
One of the most insightful clues as to the location of the Land of Punt involves the etymology of the word Berber. It has often been assumed — incorrectly — that the appellative originated with the ancient Greeks as a cognate of barbaros (“barbarian”). However, the first mention of the term actually dates earlier to the New Kingdom of Egypt (c. 1500 BCE), when it served as an ethnonym for the Puntites. Specifically, during the Hatshepsut expedition to Punt, the ancient Egyptians identified their Puntite counterparts as brbrta in hieroglyphic symbols. This is believed to have been an onomatopoeic imitation on the Egyptians’ part of the “bar” or “ber” sound that was apparently common in the Puntite language (cf. AECR (1976); Bowersock (2013)). In view of these hieroglyphics, the Egyptologist Ernesto Schiaparelli suggests that the Puntites inhabited a region coinciding with northern Somalia, Eritrea (then part of northern Ethiopia), and the Atbara zone in northeastern Sudan (AECR (1976)):
In the “Dictionnaire des noms geographiques contenus dans les textes hiéroglyphiques” by H. Gauthier, the Italian Egyptologist Schiaparelli is quoted as identifying the inhabitants of Punt during the Hatshepsut expedition with the people living in the northernmost part of Ethiopia, what is now Eritrea, and in two cities named Berbera. One is situated just north of Atbara, the town at the confluence of the Blue and White Niles, the other on the coast of Somaliland. Schiaparelli based his conclusion on the hieroglyphic signs which stood for the Puntites, namely [brbrta]. I believe that these signs brbr were an onomatopoeic attempt on the part of the Egyptians in the expedition to imitate the language of the people with whom they were dealing. The actual occurrence of brbrta as identifying the people of Punt, a historical people of ca. 1500 B.C., gives weight to the theory that it was here in the Egyptian New Kingdom that the word Barbar originated, rather than in the land of the Sumerians, or the Semites, or the Indo-Europeans.
Ancient Egyptian statue of the Pharaoh Sesostris I (Senusret I). According to a stela found at Wadi Gawasis, the king ordered an expedition to Punt. Pliny the Elder specifies that Sesostris’ party therein traveled as far as the cinnamon emporium of Mosylon (Mossylum), located in present-day northeastern Somalia (Aldred (1970)).
Schiaparelli also bases his argument on the aforementioned Periplus of the Erythraean Sea, a document which repeatedly alludes to “Berbers” living in these same areas. As a result, this territory was known to the ancient Greeks as “Barbaria” or “Barbara”, meaning the “land of the Berbers” (Huntingford (1980)). The Periplus indicates that there were Berber commercial settlements all along the Red Sea coast during the 1st century CE, with two such concentrations: one in the “Barbaria” in the Nile Valley around southern Egypt and northern Sudan, and the other in the “far-side” ports of the “other Barbaria” in the Horn (viz. “there are other Berber market-towns, known as the ‘far-side’ ports”). These Berbers/Puntites were therefore still trading in frankincense and other commodities in the southern part of their territory, just as they had over a millennium before in Pharaoh Hatshepsut’s time. This is confirmed by the Roman scholar Pliny the Elder. In his treatise Natural History, Pliny tells us that the Pharaoh Sesostris I — a ruler who, as seen on the Wadi Gawasis stela, ordered at least one expedition to Punt during his reign — led his men to the “far-side” Berber port of Mosylon (Mossylum), a cinnamon emporium located in the present-day Bosaso area in northeastern Somalia:
Beyond these is the Gulf of Abalites, the island of Diodorus, and other desert islands ; also, on the mainland, a succession of deserts, and then the town of Gaza, and the promontory and port of Mossylum, to the latter of which cinnamon is brought for exportation : it was thus far that Sesostris led his army.
Another key aspect of the Barbaria connection is the form of governance that the territory’s denizens were said to have adhered to. The Periplus indicates that the Berbers were divided into tribal communities, each ruled by its own chief. These independent city-states in the greater Barbaria were, in turn, overseen by a learned king or paramount chief named Zoscales (Zoskales):
On the right-hand coast next below Berenice is the country of the Berbers. Along the shore are the Fish-Eaters, living in scattered caves in the narrow valleys. Further inland are the Berbers, and beyond them the Wild-flesh-Eaters and Calf-Eaters, each tribe governed by its chief; and behind them, further inland, in the country towards the west, there lies a city called Meroe.[…]
These places, from the Calf-Eaters to the other Berber country, are governed by Zoscales; who is miserly in his ways and always striving for more, but otherwise upright, and acquainted with Greek literature.[…]
The voyage to all these farside market-towns is made from Egypt about the month of July, that is Epiphi. And ships are also customarily fitted out from the places across this sea, from Ariaca and Barygaza, bringing to these far-side market-towns the products of their own places; wheat, rice, clarified butter, sesame oil, cotton cloth, (the monache and the sagmatogene), and girdles, and honey from the reed called sacchari. Some make the voyage especially to these market-towns, and others exchange their cargoes while sailing along the coast. This country is not subject to a King, but each market-town is ruled by its separate chief.
As in the Berber period, the various Puntite districts were apparently governed by separate leaders. This is clear from Egyptian hieroglyphics, which Balanda (2005) notes repeatedly allude to the “chiefs” of Punt in the plural. For example, an inscription at Deir el-Bahri reads: “pitching tents for the king’s representative and his (the king’s) expedition to the myrrh terraces on both sides of the sea[…] in order to receive the chiefs of this land.” Likewise, an inscription at Sinai, which dates from the 36th regnal year of the Pharaoh Amenophis III (r. 1390–53 BCE), records an official indicating that: “I went forth by the sea coast (pr.n=j hr-gs wtd-wr) to announce the marvels of Punt to receive aromatic gums which the chiefs had brought (jn.n wrw) in their Khementy-boats as revenue from unknown lands.” The loosely centralized governmental structure of the Berbers/Puntites, therefore, also appears to have remained essentially unchanged since the New Kingdom.
(*N.B. From the Deir el-Bahri hieroglyphic text above, we also learn that there were “myrrh terraces on both sides of the sea,” which the ancient Egyptians visited during their sojourns in Punt. This statement appears to give some credence to Balanda’s assertion that the Puntites inhabited both the Horn of Africa and the neighboring parts of the Arabian peninsula. Other evidence to that effect includes linguistic analysis by Alexander Militarev, who identified a Cushitic substratum in the Modern South Arabian languages. Militarev, an advocate of a Levantine origin for the Afro-Asiatic language family, suggests that this signifies that a) Cushitic speakers originally dwelled in Arabia, in an area adjoining that of the speakers of the MSA languages, b) most Cushitic speakers later migrated to Northeast Africa, and c) the Cushites who stayed behind in the Arabian peninsula were assimilated by their Semitic neighbors (cf. Blažek (2013)). Since the ancient Himyarites occupied the Hadramout governate and southwestern areas in Yemen near the domain of the MSA speakers, it is relevant to note that “in the 5th century the Himyarites, in the south of Arabia, were styled by Syrian writers Cushaeans and Ethiopians” (Baynes (1878)). Additional evidence supporting Balanda’s contention includes genetic and anthropometric analysis: Non (2010) observed close mtDNA ties between various Cushitic/Ethiosemitic-speaking populations of the Horn and Yemenis from the Hadramout governate, and Billy (1988) reported that his Cushitic and Ethiosemitic-speaking samples shared anthropometric affinities with Yemenis.)
Hieroglyphic signs for khebsi, a secondary ancient Egyptian name for the Puntites, which they would often use during the later Ptolemaic era (Naville and Carter (1906)).
Édouard Naville relates that the ancient Egyptians also referred to the Puntites by another name, “the khebsi of the divine land.” This designation was apparently uncommon in earlier times, but subsequently gained currency during the Ptolemaic era, when it often appeared on hieroglyphic texts. Glaser links the appellation khebsi with that of Habesch, a toponym denoting the later territory of Abyssinia (Naville and Carter (1906)). Naville writes:
Another name of the Puntites is[…] “the khebsi of the divine land.” This name is somewhat rare in the old texts, but it is frequently found at a later period in the Ptolemaic inscriptions which speak of Punt. Brugsch translates the word khebsi, “those who cut or detach the incense from the trees;” Max Muller, “the curled men, or the men with long curls.” Brugsch already alludes to the assonance on which Glaser lays stress, between that word and that of Habesch, which is the name of the actual Abyssinia, and which used to stretch over a vaster territory. Whatever may be the meaning of the word, it seems to designate the real Puntite population of Hamite race, to distinguish it from the negroes who also inhabited the land of Punt.
Thus, the ancient Egyptians called the Puntites firstly brbrta (an umbrella term for Cushitic speakers of Northeast Africa) and then alternately brbrta and khebsi. The advent of the later designation, khebsi, coincides precisely with the time period when the Cushitic-speaking Agaw forebears of the Abyssinians are thought to have adopted Semitic tongues. It therefore appears that the Egyptians were trying to draw attention to the fact that, although the Abyssinians had switched languages, they, now styled as khebsi, were still ultimately of the same Cushitic Puntite stock as the Agaw and other brbrta.
A children’s coffin from Thebes, Upper Egypt, dating to the Roman Period. The structure is made of wood and is plastered. A portrait of two young females can be seen painted on the coffin’s interior. These individuals are dressed like, and have the appearance of, present-day Cushitic-speaking Agaw women and Abyssinian women (Agaws that adopted Semitic languages) of the Horn region (The Arts Society).
(For more details, see Why did the ancient Egyptians refer to the Puntites as brbrta if they were Cushitic speakers rather than Berber speakers? and What evidence is there that the Puntites spoke an Afro-Asiatic language?.)
Ancestral and spiritual homeland
Rock art at Laas Geel in northern Somalia, which appears to depict cow worshiping in honor of the ancient Egyptian deity Hathor (Phys).
The ancient Egyptians held a special reverence for the Land of Punt. Known to them as Ta netjer or Ta nuter (“God’s Land”), they regarded it as both their ancestral homeland and a spiritual center. Thus, whenever the ancient Egyptians depicted the Puntites on their temple walls, they consistently showed them as being similar to themselves in appearance and size (unlike other non-Egyptians, who were instead frequently caricatured). Punt was likewise always identified in the hieroglyphic texts without the determinative symbolizing a foreign territory.
The Egyptologist Gaston Maspero summarizes these conventions as follows:
The legends which seem to bring the ancestors of the Egyptians from the Red Sea coast have already been mentioned. They are closely connected with the worship of the Sky and Sun god Horus of Edfu. Hathor, his nurse, the “House of Horus,” the centre of whose worship was at Dendera, immediately opposite the mouth of the Wadi Hammamat, was said to have come from Ta-neter, “The Holy Land,” i.e. Abyssinia or the Red Sea coast, with the company or paut of the gods. Now the Egyptians always seem to have had some idea that they were connected racially with the inhabitants of the Land of Punt or Puenet, the modern Abyssinia and Somaliland. In the time of the XVIIIth Dynasty they depicted the inhabitants of Punt as greatly resembling themselves in form, feature, and dress, and as wearing the little turned-up beard which was worn by the Egyptians of the earliest times, but even as early as the IVth Dynasty was reserved for the gods. Further, the word Punt is always written without the hieroglyph determinative of a foreign country, thus showing that the Egyptians did not regard the Punites as foreigners. This certainly looks as if the Punites were a portion of the great migration from Arabia, left behind on the African shore when the rest of the wandering people pressed on northwards to the Wadi Hammamat and the Nile. It may be that the modern Gallas and Abyssinians are descendants of these Punites.
The Egyptologist Flinders Petrie, moreover, contends that the stimulus for the establishment of ancient Egypt’s first dynasties arrived from Punt in the south, a territory which he asserts was located on both coasts of the Red Sea, in the Horn of Africa and Arabian Peninsula:
On looking to the Egyptian representatives of the various races known to them, we see but one resembling the Egyptian high class race. The people of Pun (Fig. II), so admirably sculptured on the temple of Hatshepsut, are very closely like the high Egyptians. Further, the Egyptians call Pun “the land of the gods” ; and they do not appear to have made war on the Punite race, but only to have had a peaceful intercourse of embassies and commerce. It appears that Pun, or Punt, was a district at the south end of the Red Sea, which probably embraced both the African and Arabian shores. The name is connected with the Poeni, or Phoenicians, who appear to be a branch of that race. The Egyptians may then be another branch of the Punite race, and their earliest immigration into Egypt confirms this.
Petrie (1920) explains further that, since dynastic Egypt’s first Pharaoh Menes/Narmer hailed from the old city of This or Thinis in Upper Egypt — as did the Thinite Confederacy, a grouping of predynastic Egyptian nobles led by Menes — and because This would remain Egypt’s first capital until the Third Dynasty, when the seat of government would be moved northward to the newly-established city of Memphis, the impetus for the foundation of dynastic Egypt originally came from the south. In other words, this catalyzing influence ultimately penetrated the Nile Valley from the direction of ancient Punt:
Before Menes comes a dynasty of kings of This, and Menes is the Thinite who led his people to a new capital at Memphis. If the invading race had come in from the north, or from Suez, Memphis would have been naturally reached first, and their establishment so high up as This would be less likely [by “up” Petrie means toward the source of the Nile river, which is located well south of Egypt; therefore, traveling upriver here is traveling southward]. But the monarchy starting at This, in the middle of Egypt, points to the race having come into Egypt by the Koser road from the Red Sea. They must further have come from the middle or south end of the Red Sea ; as, if they were from the north end, they would have entered at Memphis. The first settlement being at This points then to an origin in the southern half of the Red Sea.
Correspondingly, George Rawlinson reports that hieroglyphic inscriptions variously describe Amun, the king of the ancient Egyptian gods, as the Hak or King of Punt. The sky deity Horus is also venerated as “the holy morning star which rose to the west of the land of Punt.” According to Petrie, all of Egypt’s earlier Pharaohs, in fact, claimed divine descent from Horus. The Egyptian priest Manetho (fl. 300 BCE) identifies these rulers as Thinite and Memphite in his lists. During the Fifth Dynasty, this tradition apparently changed because the Pharaoh Userkaf and other new kings, who Manetho identifies as Elephantine, each instead started to refer to himself as “Son of Ra” after the sun god Ra. Petrie (1920) writes:
[…] it has never been observed that no Egyptian king claims descent from Ra until this Vth dynasty. The earlier kings are always Horus kings, or Horus and Set united ; but no king calls himself “Son of Ra” until the new dynasty, who are here stated to be children of the god Ra, and to begin as his high priests at Heliopolis.
Moreover, a Ramesside monument inscription at the Sinai peninsula identifies the lunar god Thoth as the “Lord of Punt” (cf. Shaheen (1998)). This ancient text in full reads (Gardiner and Peet (1917)):
Beloved of Hathor, lady of the turquoise, beloved of Thoth, lord of Punt.[…]
Lord of the two lands, Usimarrēr Setepenrēc, beloved of Hathor, lady of the turquoise, lord of risings, Ramesses Meriamūn beloved of Thoth, lord of Punt.
As the historian Amelia Edwards further notes, Hathor, the greatest and one of the oldest of the ancient Egyptian female deities, was in fact styled as the “Lady of Punt”:
The Egyptians entertained an extreme reverence in the abstract for the Land of Punt, which apparently formed part of a larger district known generally as Ta-nuter, or the Land of the Gods. Hathor and Bes, two of the principal deities worshipped by the Egyptians had their divine origin in Punt, and Hathor was adored under a special form as “The Lady of Punt.” Bes, in his grotesque features and general characteristics, is clearly a barbaric divinity, and is occasionally represented as nursing or devouring the large cynocephalus apes depicted in the wall-sculptures of Dayr-el-Bahari as indigenous to the Land of Punt. The Egyptians appear to have cherished a vague tradition of their own origin as natives of Ta-nuter at some extremely remote period ; and it is interesting to note that the curved beard characteristic of these natives of the Land of the Gods is a special attribute of divinities as well as of deified personages in Egyptian art.
Another sobriquet that the ancient Egyptians used to identify the goddess Hathor was “Mistress of Punt” (cf. Redman (2021)). The respect they held for her, and for Punt generally, is perhaps best expressed by Pharaoh Hatshepsut herself. Hieroglyphic inscriptions attributed to this monarch remark (Manley (1996)):
It is the sacred region of God’s Land; it is my place of distraction; I have made it for myself in order to cleanse my spirit, along with my mother, Hathor… the lady of Punt.
Given the above, we can better situate ancient Punt by examining whether or not the various hypothesized Puntite locales once had an established spiritual cult devoted to deities from the Egyptian pantheon. To this end, the Roman scholar Pliny the Elder refers to the area near present-day Bulhar in northwestern Somalia as the “Port of Isis” (Isidis Portus), so named after the Egyptian goddess Isis. In the original Latin text, Pliny states that “in eum Troglodytis myrrha confertur” (Österreichische Leo-Gesellschaft (1941)). This phrase translates into English as “myrrh of Troglodytes is poured into him.” Ergo, in antiquity, the Port of Isis appears to have held a ritual significance. Certain pastoral-themed rock art in northern Somalia, such as at the Laas Geel site, likewise seems to depict worshipers in ceremonial attire honoring the deity Hathor, the presiding “Lady of Punt” and figurative “mother” of Queen Hatshepsut (cf. Ibrahim (2013)).
Analysis of toponyms or place-names provides additional, confirmatory evidence that northern Somalia was a spiritual hub in ancient Punt. According to Ibrahim (2013), the traditional name for the northern Somali territories, waaqoyi, is etymologically derived from waaq, an appellation connoting both the pre-Abrahamic religion of the Cushitic peoples and its sky-god. As such, waaqoyi literally translates as “God’s Land” (Ta netjer) — the very name which the ancient Egyptians used to refer to the Land of Punt:
The Somali word Waaqooyi or Waqooyi has two meanings: a) North direction; and b) the northern mountainous areas facing the Gulf of Aden. Since Waaq (in Somali) and Waaqoo (in Oromo) are equivalent to God, Waaqooyi is undoubtedly the Land of gods! Because of the past sacredness of those northern mountainous areas, their relation to Waaq can be fully justified. Many other place names in the northern Somali coast and hinterland can be identified with Waaq such as Waaqeel (a plain near Gacan Libaax Mountain), Waaderiaya (on the coast between Mait and Las Qoray), Waaqderi (near Las Anod) and Waaq Dhambala (the Estuary where Waheen seasonal watercourse enters the sea).
It could be as simple as that! Waaqooyi — a word which is as ancient as the Somalis — means ‘the land of gods.’ Therefore, the Land of Punt could be no other place than Waaqooyi (the northern Somali coast).
(For additional details, see Were the ancient Egyptians related to the Puntites?.)
Biological affinities
Egyptologists have long acknowledged that whichever geographical location truly conforms with the Land of Punt, it should have populations that share close biological ties with the ancient Egyptians; or, at the very least, with the ancient Egyptians’ lineal descendants, the modern Egyptians. This rules out Mozambique and Uganda since their Bantu and Nilotic majorities are fairly recent settlers, and these groups also do not have any significant affinities with the ancient and modern Egyptians. The Afro-Asiatic (Hamitic-Semitic) speaking populations of the Horn, Sudan, Maghreb and the Arabian peninsula, on the other hand, are in an altogether different position. Anthropological studies have confirmed that they share close physical and genetic ties with Egyptians as a whole. This perhaps should not come as a surprise, for the ancient Egyptians were Afro-Asiatic speakers too.
![A Somali man of "pure" Cushitic type, also showing little Sub-Saharan African influence. Notice the soft-textured close-cropped hair. Clark indicates that "during [Amenhotep II's] reign and earlier, in that of Tuthmosis III, an intermediate 'bobbed' hairstyle appears, and thereafter Puntites have close-cropped hair similar to that of the chief of Punt under Hatshepsut."](https://landofpunt.files.wordpress.com/2023/08/somali-man-cushitic-194a.jpg?w=529)
Morphology
As seen at Deir el-Bahri, among other archaeological sites, the ancient Egyptians consistently show the Puntites on their temple walls as being very similar to themselves in physical type — just as though these groups were sibling populations. The Puntites are depicted as moderately tall and of gracile build, with “Caucasoid” features and reddish-brown skin; they also frequently wear their hair long. John Desmond Clark writes:
The Puntites are depicted in several Eighteenth Dynasty scenes. Typically, the men have dark reddish skins and fine features; characteristic negroid types are not shown, although they occur amongst depictions of riverine southerners (of Wawat, Kush, Irem, etc.). Other Puntite features are also not found amongst other southerners. Long hairstyles are typical for Puntites until the reign of Amenhotep II; during his reign and earlier, in that of Tuthmosis III, an intermediate ‘bobbed’ hairstyle appears, and thereafter Puntites have close-cropped hair similar to that of the chief of Punt under Hatshepsut. A long or medium dressed goatee is found at all periods
— Peoples of Cushitic origin. Top row, from left to right: Somali man, Beja man, Afar man, Oromo man. Bottom row, from left to right: Somali woman, Agaw woman, Afar woman, Oromo woman. Modern Cushitic peoples with the cad (“light” or “olive-skinned”) phenotype and mariin (“red-brown”) phenotype have in all essential aspects retained the physiognomy of their Cushitic ancestors. For this reason, they closely resemble the Cushitic Puntite figures depicted at Hashepsut’s temple and other ancient monuments in Egypt.
Such external morphological traits are relatively common among the Afro-Asiatic-speaking populations on either side of the Red Sea (see, for example, the anthropometric studies Leguebe (1981) and Billy (1988) below). For our purposes, then, the skeletal characteristics of these groups are more useful in locating Punt.
In terms of height, the Egyptologist Édouard Naville states that the Puntite chief Perahu (Parahu) is depicted on the Deir el-Bahri temple walls as “a tall well-shaped man.” This portrayal concurs with the estimated stature of an ancient individual excavated at Heis (Xiis), a site in northwestern Somalia corresponding with the old “Berber” port of Mundus (discussed further below). Measurement of the total length of this adult male skeleton (c. 21 to 46 years in age), which was exhumed from Tomb 75, produced a height of around 1.68 m (cf. González-Ruibal et al. (2022)). When adjusted for flesh covering the bones in the living and minor osteological shrinkage from dehydration, this gives a stature close to the averages of 1.729 m and 1.723 m, respectively, which Boughey (1971) reported for his Dir male sample from northwestern Somalia and Puccioni (1931) reported for his Darod male sample from northeastern Somalia.
Craniometric relations between various global populations. The Afro-Asiatic-speaking groups of the Horn of Africa cluster with Egyptians from all periods, from the predynastic era through to the dynastic and modern epochs. This concurs with the theory that Punt, a brotherly civilization to ancient Egypt, was located in the Horn (Kemp (2006)).
In 1975, G. P. Rightmire analysed the skulls of ancient Egyptian male and female individuals (Egyptian E, Gizeh) and Cushitic specimens of the Pastoral Neolithic, comparing the crania of these Afro-Asiatic speakers with those of various earlier Khoisan and Bantu samples (including Rwanda). He observed that most of the Egyptian and Cushitic individuals were distinct from the Sub-Saharan African samples, with the Cushitic Baharini, Makalia I, and Elmenteitan F1 specimens in particular showing close ties with their Egyptian counterparts. As Rightmire put it, “there is little doubt about Baharini as Egyptian female.” Analogously, Kemp (2006) found that the ancient and modern Egyptians are craniometrically closest to other Afro-Asiatic speakers and Nubians inhabiting Northeast Africa. They are also more distantly related to populations in the Near East, but again share no significant affinities with the ancient and modern “Negroid” populations in Africa. Spradley (2006) compared the skulls of recent populations from northern Somalia and Egypt with those of various Sub-Saharan African groups and recently mixed African Diaspora populations, including African Americans (who are descended from enslaved Niger-Congo speakers from West Africa, with ancillary European and Native American admixture). She similarly observed that the individuals from “Somalia and Egypt are closest to one another.” Likewise, Terrazas Mata and Benavente (2013) report that their Horn of Africa and Dynastic Egypt samples craniometrically cluster together, separately from their Niger-Congo, Nilo-Saharan and Khoisan-speaking samples. The scientists attribute this shared affinity between the Horn and Egyptian groups to common descent from ancestral Afro-Asiatic-speaking populations, whom they suggest ultimately arrived from the Middle East. In the same vein, Adel et al. (2021) note that:
Craniofacial features are considered one of the most unique features of populations with different ethnic backgrounds. The Egyptians present facial features close to those of Northeast Africans, Mediterranean Asians, and Europeans, all of them sharing Caucasian ancestry.
The biological ties between the Afro-Asiatic-speaking groups in Northeast Africa are, in fact, so well established that researchers have moved on to exploring which specific ancient “Hamitic” series in the region they share the most immediate affinities with (e.g. Batrawi (1946); Mukherjee (1955); Billy (1977); Billy (1981b); Rösing (1990)). G. Billy (1975) summarizes these findings thusly:
During the dynastic era, this last [Upper Egyptian] variety covered a wide central zone of the Nile valley, stretching well beyond towards East Africa, as shown by the similarity which persists with the present-day Ethiopian populations (Tigre) or even Somali. By virtue both of its diffusion and its perenniality, they deserve to be assimilated to the basic population type of the Egypto-Nubian complex.
Osteological relations between Galla, Somali and Tigre populations and other “Hamitic” series from early Northeast Africa as well as the ancient Near East. Batrawi (1946) notes that “the affinities of the Galla and Somali to the A-group, C-group and Meroitic populations of Lower Nubia : these are very close, and they may suggest the extension of the Predynastic Upper Egyptian type over a very wide area in north-east Africa. It cannot be said with any certainty, however, whether the people of that type existed over all that area since Predynastic times, or whether they were pushed from their earliest known home in Egypt southwards” (Batrawi (1946)).
Altogether, this highlights the close relatedness of the Puntites and ancient Egyptians, ancestral populations whose apparent descendants have remained biologically proximate.
In addition to the preceding, Egyptologists have noted a sporadic occurrence of blondism in the Land of Punt. This can serve as a helpful hint as to where the territory was situated, for blond individuals were relatively uncommon in the ancient world. The Hatshepsut expedition murals show the Egyptians being received by a chief of Punt, the aforementioned Parahu/Perahu, who Édouard Naville (1906) indicates is depicted as “flaxen” or blond-haired. Naville writes:
The Punite is a tall, well-shaped man, of a type which certainly belongs to the Caucasian race ; his hair is flaxen, and is divided into well-made plaits ; his nose is aquiline, his beard long and pointed ; he wears only a loin-cloth, with a belt, in which a dagger is fixed. The left leg of the chief is covered with a bracing of rings which seem to be of metal. The Punites are representatives of the Hamitic race, of that red race to which the Egyptians also belong, and which gave its name to the Erythraean Sea. They are painted red, but with the red of Horus, which is not exactly like that of ancient Egyptians.
Sidamo women wearing traditional headbands. Once again, we can observe a conspicuous similarity with the headband donned by the Puntite chieftess Ati.
Sculpture of the ancient Egyptian Princess Nofret (c. 2613-2498 BCE, 4th Dynasty, Old Kingdom). Like their Puntite relatives, ancient Egyptians were also known to wear traditional headbands (Egyptian National Museum).
Sweeney (2006) argues that this precludes both the Horn and Southern Arabia as prospective locations for the Land of Punt since there are few blond individuals today among the Afro-Asiatic-speaking populations in these areas. He asserts that the blondism points instead to an Indo-European source centered in the Near East; either the Indo-Iranian mariyanna elites of the Levant, or the Phoenicians (whom he proposes may have acquired such an element through intermarriage in western Europe). However, ancient blond individuals did, in fact, exist in the Nile Valley itself. Archaeologists working in burial sites associated with the Meroitic culture have unearthed a number of clearly blond specimens. Janssen (1978) reports that “330 graves were excavated in cemetery 221 (Meroitic) and a proportion of blond individuals of Caucasoid type found.” Hrdy (1978), also analysing Meroitic remains, notes that many ancient individuals buried at Semna South in Sudanese Nubia had blond or red hair. He suggests that this “probably points to a significantly lighter-haired population than is now present in the Nubian region.” Archaeogenetic analysis indicates that this blondism was likely inherited from earlier European settlers in Nubia. A Meroitic period individual (dated to c. 350 BCE to 350 CE) excavated from the Kwieka site in northern Sudan was found to bear the Eurasian mtDNA haplogroup H2a, and, moreover, this ancient mitogenome showed ties with Northern European mitochondria (cf. Breidenstein (2023)). Given these finds, the minor incidence of blondism among the Puntites appears to reflect a direct Meroitic strain in this population and, indirectly, an Indo-European one (for empirical evidence of such an influence, see Batrawi diagram above). This too is in agreement with a Northeast African locus for Punt.
(*N.B. Lazaridis et al. (2022) conducted a comprehensive analysis of phenotypic traits borne by ancient individuals exhumed in Europe and Asia, and report that blond hair was most common among their early European samples (viz. 62.5% in Early Medieval Germany, 42.9% in Pre-Christian Iceland, 40% among ancient Saxons of England, and 40% among the Motala hunter-gatherers of Sweden; cf. Supplementary Materials). By contrast, the scientists did not observe instances of blond hair in any of their examined ancient Levantine samples, except for a low frequency of 14.3% among specimens from Chalcolithic Israel — a situation clearly ascribable to foreign influx since no blond individuals have been found among both earlier Levantine samples from Mesolithic and Neolithic Israel and later Levantine samples from Middle-to-Late Bronze Age Israel. In brief, archaeogenetic analysis has confirmed that the blondism documented among the Meroites and ancient Egyptians is almost certainly affiliated with early European populations, as are the instances of red hair in the Nile Valley (refer to Ancient DNA from Sudan). For details on these ancient peoples, who introduced the Steppe genome component to Northeast Africa, see Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa.)
Genetics
Y-DNA and mtDNA
Along with comparative morphology, genetic analysis provides much information on the likely location of ancient Punt. Examination of uniparental lineages, which include both Y-DNA (paternal) and mtDNA (maternal) haplogroups, has revealed strong ties between the Afro-Asiatic-speaking populations in the Horn and Nile Valley.
Phylogenetic tree of the E1b1b paternal haplogroup. Notice how the V32 subclade that is frequent in the Horn immediately descends from V12, the most common lineage today among southern Egyptians (Eupedia).
Trombetta et al. (2015) observed that around 74.5% of southern Egyptians are haplogroup E1b1b-V12* or E3b1a1 carriers (cf. Supplementary Table 7). According to Hirbo (2011), this clade is next most prevalent among Cushitic-speaking populations in East Africa (Garreh=74.1%, Gabra=58.6%, Borana Oromo=50%). E1b1b-V12 is also immediately ancestral to the V32 subhaplogroup that constitutes the main paternal lineage among ethnic Somalis from Somalia (80%) and Tigre individuals in Eritrea (60%) i.e., in the core ancient Puntite area. (*N.B. While the Tigre nowadays speak a Semitic tongue, their language contains a North Cushitic (Beja) substratum. This suggests that they were originally Cushitic speakers, who later adopted a Semitic language from South Arabian settlers in the Eritrean highlands. Accordingly, Hirbo (2011) reports that his sampled Beja individuals overwhelmingly belong to the E1b1b paternal haplogroup (100%). Hassan et al. (2008) further indicate that most of their Beja male samples, like the Tigre, fall under the clade’s V32 subhaplogroup.)
Antonio et al. (2019) identified E1b1b-V12 in two Imperial and Medieval Roman individuals, remarking that “this haplogroup is present at high frequency across present-day North Africa, especially in Egypt (up to 74.5%)” (cf. Supplementary Materials). Moreover, Sirak et al. (2021) observed a prevalence of the Y125054 subclade of V12 among Christian-era Nubian individuals from Kulubnarti in Sudan. These specimens also had considerable West Eurasian ancestry, which the scientists propose was likely derived from Egypt. Prendergast et al. (2018) further indicates that an ancient Cushitic individual (ca. 3350-3180 BP) excavated at Cole’s Burial, Kenya, a site associated with the Pastoral Neolithic cultural complex, bears the CTS3282 sublineage of V32. As of 2021, this is the earliest reported instance of E1b1b-V32 in the archaeogenetic record. (For more details, see I have heard that the V12 and V32 subclades of the paternal haplogroup E1b1b, which are today common among Afro-Asiatic speakers in Northeast Africa, are old Arabian lineages. Is this true?.)
Similarly, mitochondrial analyses by Stefflova et al. (2011) and Boattini et al. (2013) found that the maternal haplogroups that are common among the Cushitic and Semitic-speaking Afro-Asiatic populations in the Horn are also frequent among the Afro-Asiatic speakers in Egypt. These shared clades include the M1 haplogroup, whose earliest occurrence has been detected among Epipaleolithic Iberomaurusian specimens excavated at Taforalt (Loosdrecht et al. (2018)) and early Neolithic individuals buried at Ifri n’Amr or Moussa (Fregel et al. (2018)), both located in Morocco. Loosdrecht et al. (2018) note that the M1 and U6 mtDNA lineages are “mostly confined to present-day populations in North and East Africa” and that “U6 and M1 have been proposed as markers for autochthonous Maghreb ancestry, which might have been originally introduced into this region by a back-to-Africa migration from West Asia.” Stevanovitch et al. (2004) also report that M1 is now particularly common in Egypt’s Gurna Oasis, suggesting an ancestral connection between the Gurna population and other Afro-Asiatic-speaking communities in the Horn of Africa.
Autosomal DNA
As discussed on Ancient DNA from Ethiopia, the Afro-Asiatic-speaking populations in Northeast Africa also appear to share the same autosomal DNA (auDNA) signature. Hodgson et al. (2014) observed that ethnic Somalis, Afar, Amhara, Tigray and Oromos are defined by a West Eurasian-affiliated ancestral component, which they refer to as “Ethio-Somali”. Dobon et al. (2015) found that roughly the same ancestral element defines Egyptian Copts, Beja, Afro-Asiatic-speaking Ethiopians, Sudanese Arabs, and many Nubians (evidently, individuals descended from the Hamitic Kasu or “red Nubians”). By contrast, Hodgson et al. discovered that the Omotic-speaking Ari are defined by a separate, Nilo-Saharan-affiliated ancestral component, which the researchers call “Ethiopic”. Thus, the Ethio-Somali/Coptic and Ari components appear to correspond rather closely with the “red” (“Hamitic”) and “black” (“Negroid”) ancestral populations, respectively, that are depicted on the ancient temple walls and mentioned in the hieroglyphic inscriptions as having inhabited Punt.
Phylogenetic clustering of ancestral components based on genetic distance (Fst). The Cushitic genome component, which typifies modern Cushitic, Ethiosemitic and North Omotic-speaking populations in the Horn of Africa, groups nearest to the Berber, Southern European and other West Eurasian components (Shriner et al. (2016)). This is consistent with the “Ethio-Somali” ancestral component, which Hodgson et al. (2014) similarly found had a predominant non-African affinity based on genetic distance (cf. Supplementary Text S1). The Omotic component, which peaks among the Ari and other South Omotic speakers, groups instead closest to the Nilo-Saharan, Niger-Congo and other Sub-Saharan components. This is consistent with Hodgson et al.’s “Ethiopic” component, which the researchers found had a nearest genetic distance to the Nilo-Saharan, Niger-Congo and other Sub-Saharan components (Hodgson et al. (2014), Supplementary Text S1).
The foregoing was also confirmed by López et al. (2021), who compared the genomes of the Afro-Asiatic-speaking groups in Ethiopia with those of other global populations, both ancient and modern. The scientists identified a West Eurasian ancestral component among the local Afro-Asiatic speakers, which they suggest is Egypt-related and traces back to the inhabitants of the Land of Punt:
Ethiopians in the southwest, typically NS speakers plus a few non-NS speaking groups (Chabu, Dasanech, Karo), are more related to Bantu and Nilotic speakers relative to AA speakers in the northeast that instead show more ancestry related to Egyptians and West Eurasians (Fig 3, Fig S12). The inferred timing and sources of admixture related to Egypt/W.Eurasian-like sources, starting around 100-125 generations ago (~2800-3500 years ago; Fig 3, Fig S12) as in previous findings (Pickrell et al., 2014; Pagani et al., 2012), is consistent with significant contact and gene flow between the peoples of present day Ethiopia and northern Africa even before the rise of the kingdom of D’mt and interactions with the Saba kingdom of southern Yemen which traded extensively along the Red Sea (Currey, 2014; Phillipson, 2012). This timing is also consistent with trading ties between the greater Horn and Egypt dated back only to 1500 BCE, when a well-preserved wall relief from Queen Hateshepsut’s Deir el-Bahari temple shows ancient Egyptian seafarers heading back home from an expedition to what was known as the Land of Punt (SI Section 1A).
An Ababda Beja girl.
In 2022, Osman and Jonasson for the first time studied the genomes of Northern Somali individuals, who were born in the northeastern Puntland region of Somalia (where most hail from the Majerteen Darod clan). The scientists conducted a Principal Component Analysis, which compared their Northern Somali cohort with other global samples from Sub-Saharan Africa, North Africa, the Middle East, South Asia and East Asia. They observed that some of the Northern Somali individuals clustered with the Oromo samples, whereas a few other outliers grouped near the Ethiopian Jew samples. However, the majority of the Northern Somalis were located in a position parallel to but separate from the North African/Arabian cluster. This suggests that, on average, Northern Somalis from the Puntland region bear a predominant non-African ancestry, which is comparable in frequency to that carried by North African individuals. Such a finding agrees with the theory that the Land of Punt, a sibling civilization to ancient Egypt, was located in the Horn region (cf. Fig. S1).
As of 2023, Northern Somalis from the northeastern Puntland region of Somalia have not been analysed for Y-DNA haplogroups. However, in connection with the autosomal DNA findings above, Trombetta et al. (2015) report that the Afro-Asiatic-speaking Eritrean samples in their dataset mostly belong to the E1b1b lineage, with many falling under the V32 subclade (observed total E1b1b percentages for Afro-Asiatic-speaking Eritreans: Tigre=100%; Saho=98.9%; Tigray-Tigrinya=71.9%; cf. Supplementary Table 7). This is instructive because, after Northern Somalis from Puntland, Cushitic and Ethiosemitic-speaking Eritreans have the next highest reported frequency of Eurasian ancestry in the Horn of Africa (e.g. refer to the Tigray-Tigrinya samples in Hodgson et al. (2014), Supplementary Text S1; Llorente et al. (2015)). Hence, the parts of the Horn that are thought to have formed the nucleus of the old Punt civilization (northern Somalia and Eritrea) are still peak areas for Eurasian ancestry. Likewise, the Afro-Asiatic speakers here that have been studied primarily bear Egyptian-related uniparental lineages; either the V32 subclade, which descended from V12 (the most common paternal haplogroup today in Upper Egypt), or, as in the case of most Saho individuals, the V22 subclade that has been observed among ancient Egyptian mummies (see ancient DNA discussion below).
(For further details, see Which Afro-Asiatic-speaking population of the Horn region has the most non-African ancestry?.)
Ancient DNA
Coffins of the ancient Egyptian aristocrats Nakht-Ankh & Khnum-Nakht. The Two Brothers were found to belong to the M1a1 haplogroup, a signature maternal clade among Afro-Asiatic-speaking populations in Northeast Africa (Drosou et al. (2018)).
Ancient DNA analysis provides the most direct genetic evidence that the Land of Punt was located in Northeast Africa.
In 2013, a research unit led by Rabab Khairat of the University of Tübingen completed the first genetic study utilizing next-generation sequencing techniques to gauge the ancestral lineage of an ancient Egyptian. The scientists extracted DNA from the heads of five Egyptian mummies dating from the Late Dynastic to Ptolemaic periods (806 BCE–124 CE). They found that one of the mummified individuals belonged to the I2 mtDNA haplogroup. This discovery is especially interesting for a number of reasons. Firstly, the I maternal clade is closely associated with the spread of Indo-European speakers because the lineage has been detected among various ancient cultures on the European Steppe. This implies early migrations from this area into Northeast Africa; either directly from Europe or indirectly through Asia. Secondly, haplogroup I is today quite rare globally and exceeds 5% in few populations. The clade is by far most common among the Rendille and other Cushitic-speaking remnant groups inhabiting the Great Lakes region, where it has been observed at frequencies as high as 23% (cf. Castrì et al. (2008)). Thirdly, the basal or ancestral I* haplogroup has only been identified in three persons worldwide. Of these individuals, two are from Somalia and the other is from Iran (Olivieri (2013)). Lastly, I2 (formerly known as N1e) is a subclade of N1. Yatsishina et al. (2021) indicate that the haplogroup N is the most frequently occurring maternal lineage among the ancient Egyptian mummies they examined at the Kurchatov Institute (2 out of 3 specimens or ~67%). Similarly, Kılınç et al. (2016) report that N1 is the most commonly borne maternal haplogroup among the central Anatolian Neolithic individuals they studied (5 out of 9 or 55.56% of the examined specimens). This mitochondrial lineage is nowadays also rare but likewise peaks in frequency among Afro-Asiatic speakers in the Horn. (*N.B. The Rendille proper (who are known as asil or original Rendille) are distinct from the Ariaal (or assimilated Rendille). The Ariaal are instead people of Nilotic Samburu origin and “Negroid” appearance, most of whom still speak their native Nilo-Saharan language.)
In 2018, a research unit led by Konstantina Drosou of the Manchester Institute of Biotechnology again used next-generation sequencing to examine the DNA of ancient Egyptian individuals. The samples were culled from a tomb at Deir Rifeh and were dated to the Middle Kingdom (1985 BCE–1773 BCE), making them the second oldest Egyptian specimens so far to be genetically analyzed. They comprised the mummies of Nakht-Ankh and Khnum-Nakht, a pair of 12th Dynasty aristocrat siblings, who shared a mother but had different fathers. The Two Brothers were found to belong to the M1a1 haplogroup. As mentioned above, this maternal lineage is today most common among the Afro-Asiatic-speaking populations inhabiting the Horn of Africa and Nile Valley. (*N.B. The oldest Egyptian specimen thus far analyzed, the 4000-year old mummy Djehutynakht, Overlord of the Hare (15th) nome of Upper Egypt, belongs to the U5b2b5 mtDNA haplogroup. This is an early European maternal lineage, which has been found among Early Bronze Age Sardinian individuals (cf. Olivieri et al. (2017); Marcus et al. (2020)). It is also borne today by Berbers in Egypt’s Siwa Oasis. Ergo, we see here again that the population affinities of the earliest Egyptians are not Semitic (in the modern sense), much less Nilo-Saharan/Niger-Congo/Khoisan (cf. Loreille et al. (2018)).)
Interestingly, Schuenemann et al. (2017) also observed both the M1 and I mitochondrial clades as well as the E1b1b-V22 paternal haplogroup in their analysis of later New Kingdom to Roman Period (c. 1388 BCE–426 CE) Egyptian samples from Abusir el-Meleq. The researchers, moreover, identified some J clade bearers. However, it is unlikely that older Egyptian individuals from the Early Dynastic (c. 3000 BCE–2650 BCE) and Predynastic (before 3000 BCE) epochs belong to this Y-DNA lineage. This is because, although haplogroup J arrived in Egypt from the Middle East, the earliest reported example of this clade in the Levant/Arabia is quite recent, being first attested there in a 3700 year old (c. 1680 BCE) Bronze Age specimen (Haber et al. (2017)). Furthermore, the nobleman Nakht-Ankh, who is the oldest Egyptian individual to have his Y-DNA successfully identified, appears to belong to the H2m clade (cf. HaploTree; Open Genomes). In the paleogenetic record, H2 is a patrilineage mainly affiliated with populations bearing Anatolian Neolithic ancestry; particularly Neolithic Europeans (e.g. Cassidy et al. (2020)). The H2 paternal clade has also been detected in an individual linked with the Pre-Pottery Neolithic B, a culture in the Levant dating from an era marked by the arrival of new settlers carrying Anatolian Neolithic ancestry (Lazaridis et al. (2016), Table S6.1). In addition, Gad et al. (2020a) affirm that the ancient Egyptian aristocrat Yuya, the maternal great-grandfather of the Pharaoh Tutankhamun, belongs to the G2a clade. Like the H lineage, the Y-DNA haplogroup G is primarily associated with early populations bearing Anatolian Neolithic ancestry. The oldest instance of this G paternal clade in Africa has been detected among Early Neolithic specimens exhumed at Kahf Taht Al Ghar, an archaeological site located in the Maghreb. This tentatively suggests that the ancient Egyptian nobleman Yuya’s haplogroup G may have introgressed from a northwestern, likely Iberian, source population. (For further details, see Is it true that the ancient Egyptians, like the Minoans, descend from Anatolian Neolithic farmers? and If Muslim Egyptians have other admixture elements besides the ancient Egyptian component, does this mean they are mostly of non-Egyptian origin?.)
(*N.B. Gad et al. (2020b) report that the 18th Dynasty Pharaoh Amenhotep III (b. 1411 BCE), his son Pharaoh Akhenaten, and grandson Pharaoh Tutankhamun carried the R1b haplogroup, a typical European paternal lineage (also see Gad et al. (2020a)). iGENEA specifies further that they belonged to the clade’s M269 subhaplogroup, which now accounts for over half of all Y-DNA lineages in western Europe. What’s more, according to DNA Consultants, the 20th Dynasty Pharaoh Ramesses III (b. 1217 BCE) and his son Prince Pentawere (“Unknown Man E”) bore the V22 sublineage of the E1b1b haplogroup. The V22 subhaplogroup is today most common among the Cushitic-speaking Saho population inhabiting Eritrea (cf. Trombetta et al. (2015), Supplementary Table 7). Ramesses III and Pentawere were initially thought to have belonged to the E1b1a clade, an error caused by an incomplete analysis of their Y-STR markers: Whit Athey’s Haplogroup Predictor, the tool that Hawass et al. (2012) indicate they used to determine the rulers’ paternal lineage, does not have an option for GATAH4, one of the markers that Ramesses III and Pentawere were originally analysed for; when GATAH4 and the other Y-STR markers are inputed into the Nevgen Haplogroup Predictor, these men are instead assigned to the E1b1b-V22 clade. Additionally, Yatsishina et al. (2021) examined ancient Egyptian mummies at the Kurchatov Institute, and found that the specimens likewise carried the E1b1b-V22 and R1b-M269 haplogroups (see Ancient DNA from Sudan for hair morphology analysis conducted on the Kurchatov Institute’s ancient Egyptian mummies). All in all, this suggests that ancient Egyptian individuals may have borne the common North African E1b1b lineage, as well as R1b and Anatolian Neolithic-associated haplogroups (e.g. the T, H and G clades) before the haplogroup J first entered the Egyptian gene pool. It is known that the J clade was introduced to the Semitic-speaking areas of the Levant and Arabia during the Bronze Age, by outsiders from the Caucasus/Iranian plateau who carried the Caucasus Hunter-Gatherer genome component. However, when exactly this western Asian admixture spread to the Egypt vicinity is uncertain. Morez (2023) proposes that this introgression first occurred during the Second Intermediate Period (c. 1650-1550 BCE), an epoch marked by the Hyksos invasion of Egypt, since genomic analyses of an earlier, Old Kingdom Egyptian individual from Nuerat (dated to c. 2868-2492 BCE) did “show a strong genetic affinity of this sample to Levantine Natufians.” By contrast, “the Nuerat sample did not carry the Caucasus Hunter-Gatherer genetic component that started to spread across West Asia ~4000 years ago and is widely spread in present-day populations.” Likewise, genome analysis of Middle Neolithic pastoralists excavated at the Skhirat site in Morocco — ancient specimens that bear Levantine ancestry and who are believed to have spread westward from the direction of the Nile Valley — also did not detect any Caucasus Hunter-Gatherer ancestry. This suggests that, at that early point in time, this component indeed had not yet been introduced into the Egypt area (cf. Simões (2023a); Simões (2023b)).)
Ancient DNA analysis has revealed that the Egyptian Pharaoh Amenhotep III, his son Akhenaten, and grandson Tutankhamun bore the R1b haplogroup, the most common paternal clade among modern Europeans. Maternally, Pharaoh Tutankhamun belonged to the European Neolithic-associated K lineage, which he inherited from his mother “The Younger Lady”, his grandmother Queen Tiye (“The Elder Lady”), and his great-grandmother Thuya (Gad et al. (2020a)). The oldest examined Cushitic settler in East Africa was likewise found to belong to the K mtDNA haplogroup, in agreement with the idea that the Horn region was coextensive with the Land of Punt (Prendergast et al. (2018)).
(For more details, refer to our study Autosomal STR Analysis of the Ancient Egyptian Amarna Royal Family, Pharaoh Ramesses III, and Unknown Man E (Prince Pentawere. Also see Is it true that the ancient Egyptian Amarna royal family shares closest genetic affinities with Sub-Saharan African populations?.)
Additionally, Schuenemann et al. analyzed phenotype alleles carried by several of their ancient Egyptian samples. They reported that two of the individuals bore the derived allele of the SLC24A5 gene, a variant that is associated with lighter skin coloration (this mutation has also been found among ancient Cushitic pastoralists in East Africa; cf. Wang et al. (2020), Table S7). However, the specimens carried ancestral alleles at the Europe-specific SLC45A2 locus and other pigmentation-related genes. Altogether, this suggests that they had a light brown complexion. These findings are consistent with notarized contracts from the Pathyrite and Latopolite nomes. In these legally-binding documents, almost all of the ancient Egyptian signatories indicated that they had a “honey-coloured complexion”, with a “long face”, “straight hair” and a “straight nose” (Van Dorpe (2004)).
An excerpt of notarized contracts from the Pathyrite and Latopolite nomes, wherein almost all of the ancient Egyptian signatories indicated that they had a “honey-coloured complexion”, with a “long face”, “straight hair” and a “straight nose”. This is consistent with ancient DNA analysis, which has detected that the examined Egyptian specimens bore the derived allele of the SLC24A5 gene (Van Dorpe (2004)). This mutation is associated with lighter skin pigmentation, and has also been found among the early Cushitic settlers of East Africa (Wang et al. (2020), Table S7).
In 2020, Ester Oras of the University of Tartu and colleagues examined two ancient Egyptian child mummies, which likewise dated from the Late/Graeco-Roman Periods. They found that one of the individuals bore the HV haplogroup, an mtDNA lineage that nowadays occurs most frequently among Afro-Asiatic speakers; particularly Egyptians from El-Hayez oasis (Kujanova ́ et al. (2009)), Somalis from Somalia (Musilová et al. (2011)), and Asni, Bouhria, Figuig and Siwa Berbers (Coudray et al. (2009)).
Urban Christian et al. (2021) conducted a comprehensive archaeogenetic study of ancient Egyptian individuals. The samples were culled from six different archaeological sites across Egypt, a time transect spanning 4000 years of Egyptian civilization. According to the researchers, the analyzed specimens carried mtDNA lineages similar to those previously identified at Abusir. This, in turn, indicates that almost all of the examined ancient Egyptian individuals bore derivatives of the Eurasian M and N maternal lineages.
In summary, genetics strongly relates both the modern and ancient Egyptians with contemporary Afro-Asiatic-speaking groups elsewhere in Northeast Africa. This perfectly concords with Puntite origins for the latter populations. (*N.B. For detailed analysis, refer to Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa. Also see Ancient DNA from Sudan for discussion on the newly-analysed Middle Neolithic individuals excavated at Skhirat, an archaeological site in northern Morocco. These ancient herders, who originated from the Levant, appear to be among the first Afro-Asiatic-speaking populations in Africa. They also seem to have been responsible for introducing pastoralism to the continent.)
Traces of ancient civilization
Stone ruins, inscriptions and coins
One of the principal challenges in locating the Land of Punt has been the absence of artifacts that could be definitively identified with an ancient Puntite civilization. Perhaps the closest thing to that were two v-shaped arm-clamps made of ivory, which were found in an Old Kingdom tomb at Shellal in Upper Egypt. Some writers initially proposed that the person buried within the grave may have been a Puntite envoy. However, the Egyptologist David O’Connor later demonstrated that the man was, in fact, an Upper Nubian emissary since an Upper Nubian figure is shown wearing a very similar armlet on the causeway of Pharaoh Sahure’s mortuary temple in Abusir (cf. Wilkinson (2002)).
Aside from this false positive, various early scholars and colonial officials discovered traces of an ancient civilization in the Horn; one apparently distinct from and older than the Axumite Kingdom. The explorer Georges Révoil encountered many stone ruins while in northern Somalia, as well as inscriptions in a mysterious writing script.
On this lost orthography, the Ministry of Information indicates that:
An important point which is often lost sight of is that the ancient Somalis had evolved their own script systems which existed for a considerable period in their history. Convincing historical evidence in this respect is the numerous inscriptions and rockpaintings on cave-walls, on granite rocks, old coins etc., that are found to this day in various parts of the country.[…]
An interesting point, however, is that this script system was apparently based on vowel sound, not a Word-Picture writing as in ancient Egypt.
The foregoing rules out the known Sabaean and Himyarite writing systems since they and other ancient Semitic scripts, except Ge’ez, have no vowels (for details on Sabaean finds in northern Somalia, see Prioletta et al. (2021) and Hussein (2023)). Likewise, it seemingly disqualifies the Libyco-Berber orthography, for it too is an abjad (examples of this ancient script, known as Tifinagh, have recently been found in Somalia). The above also precludes Meroitic hieroglyphs because they were derived from ancient Egyptian hieroglyphics. However, Meroitic cursive remains a possibility; especially since, as an abugida, it has inherent vowel representation. In this regard, the scholar Muhammad Shamsaddin Megalommatis (2021) notes that “even today both [the Afar and Somali] languages retain ample Ancient Cushitic vocabulary that was written in Meroitic hieroglyphic and cursive writing before 2000 years. The Somali word ‘boqor’ (king) is identical to the title of the Cushitic kings of Meroe: ‘Qore’.”
Another plausible candidate is Linear A, the undeciphered second script used by the ancient Minoans of Crete (the other two writing systems being Cretan Hieroglyphs and Linear B). Several of the characters etched on the cave walls and granite rocks in northern Somalia, where most of the examples of the old Somali orthography have been found, can clearly be identified with Linear A symbols. Among these are the cross-filled circle or encircled cross (represented as the ka syllabogram in Linear A, such as on the Tablet ARKH 3a discovered at Arkhanes), the three open circles (similar to Linear A characters carved into Tablet HT 31, found at Haghia Triada), as well as various signs which Ester Salgarella of St. John’s College — who created the SigLA online database for Linear A — refers to as being “more like Chinese ideograms.” The cross-filled circle sign has also been observed on rock art in other areas of the Horn, which are inhabited by Cushitic peoples. Among these archaeological sites are Harurona in Ethiopia and Abka in Sudan. Traditionally, this cross-in-circle symbol has been associated with the ancient C-Group culture, which flourished in the Nile Valley during the Middle Kingdom. It is a common motif embossed on C-Group artifacts, and likewise still features prominently in Coptic Egyptian magic (cf. Bachechi (2005)).
During excavations of one of the tumuli at Salweyn (Salwine), Révoil found ceramic fragments of apparent Macedonian origin. These objects and other recovered items, which include Ptolemaic and Roman era artifacts, are presently kept at the Musée de l’Homme in Paris (cf. Desanges (1992)). Révoil was also able to discern a definite ancient Egyptian and Classical influence in the local culture and populace, notably in terms of physiognomy, customary law, attire, weaponry and vocabulary. These findings inspired him to declare that Auguste Mariette was indeed right, and that he had in fact stumbled upon the Punt of antiquity. The Geographical Society of Marseilles explains:
Being detained by bad weather near Hais, in a small creek named the Salwine, M. Revoil landed to get ballast, and found on the banks of the creek a great number of tumuli, similar in every respect to those which he had met with in all parts of the country, and had photographed in the course of his expedition. For the first time, and not without danger, he succeeded in partially excavating one of these tumuli, and brought to light a tomb and close alongside some remains denoting the former existence of a very advanced civilisation, including some superb enamels, fragments of pottery from Samos, and a mask, indicative of a Greek colony. Taking the observations of Dr. J. C. Prichard, who described a type, in connection with the information he obtained himself in regard to the existence of a white Galla race living farther south on the borders of Webi, M. Revoil put forward the suggestion that the present Somali race bears the marks of the existence of a white colony, very probably Macedonian, and that this colony, according to the traces he met with in the interior, has been preserved almost intact in this white tribe of Gallas. M. Revoil found besides in their idiom, arms, and dress, strong arguments in support of his opinion, which he further corroborated by an important series of profile photographs.
The scientist Johannes Maria Hildebrandt would likewise remark that:
We know from ancient authors that these districts, at present so desert, were formerly populous and civilised. I also discovered ancient ruins and rock-inscriptions both in pictures and characters. These have hitherto not been deciphered.
Over the ensuing years, Ernest-Théodore Hamy, George Andrew Reisner, Grafton Elliot Smith and many other prominent Egyptologists would echo these sentiments on similar grounds, insisting that Punt was to be found in the Horn.
In a remarkable coincidence, the Alexandrian merchant Cosmas Indicopleustes records a Greek inscription, which he had seen on a stela in Adulis, Eritrea. The basalt slab was apparently erected there by the Egyptian king Ptolemy III Euergetes (246-221 BCE) to commemorate his conquest of various kingdoms in the Near East. Phillips (1997) suggests that this signifies that Adulis itself and environs were at the time under the control of Ptolemaic Egypt:
The location of the stela observed by Kosmas, at Adulis, suggests the king had ordered it to be erected somewhere around there, possibly in gratitude for providing the means to conquer (he boasts) virtually all of Western Asia; it could hardly have been brought there by chance. This suggests, in turn, that Adulis (or the nearby place where it originally was erected) and by extension the area immediately inland, was sufficiently controlled by Egypt that its king could erect a self-laudatory stela there.
During the early 1920s, C. W. Hayward also uncovered a hoard of old coins at Port Dunford, a site in southern Somalia that is thought to correspond with the Periplus’ ancient market-town of Nikon. H. Mattingly later published these numismatic finds in 1932. Among the pieces were 22 Ptolemaic mints (c. 3rd-1st century BCE), 6 of Imperial Rome, 46 of Byzantium (primarily 4th century CE), 6 of Mamluk Egypt, and 7 of Ottoman Egypt.
In 1975, the archaeologist Neville Chittick more fully explored some of the visible ancient structures at Ras Hafun (the Periplus’ Opone) and other areas in northeastern Somalia. He described a number of heretofore obscure ruins, including stone platform monuments near Alula and sherds of unglazed Roman pottery at Damo (the Market and Cape of Spices described in the Periplus). At the Hafun Main Site, on a low ridge close to the shore, Chittick found many ceramic fragments, including a steatite sherd and blue-glazed ware of likely Parthian derivation. He also discovered an alabaster vase lying on the surface, which is reportedly similar to pottery finds from the Timna area in southern Arabia. Additionally, over a strip of land stretching out around two hectares, Chittick observed numerous stone artifacts. No actual Puntite material objects, though, had yet been excavated, and most of the Egyptian imports that had been discovered were of Ptolemaic age or younger. Consequently, supporters of an Arabian or Levantine location for Punt would often cite this archaeological void as a primary reason why the ancient territory could not have been situated in the Horn. As Sweeney (2006) concluded (somewhat prematurely), “nor was there in Eritrea or Somalia, in the time of Hatshepsut or Thutmose III, any civilization or culture of the type portrayed at Deir el Bahri that the Egyptians could have traded with[…] this topic [is] of fundamental importance to the whole debate”.
Puntite artifacts
In 2013, the scholar Ahmed Ibrahim Awale led excavations at Gol Waraabe, a site in Hargeisa valley in northwestern Somalia, where his archaeological team unearthed what appear to be the first actual artifacts belonging to the Land of Punt. As discussed in greater detail on The Mystery of the Land of Punt Unravelled – book review, the recovered objects include stone statuettes and bowls of considerable antiquity. The human figurines, in particular, bear an uncanny resemblance to those produced in ancient Egypt and by other early Afro-Asiatic-speaking populations in the Sudan area. These commonalities range from similar headdress styles, oval face shape, keen facial features and Puntite beard (Osiris beard), to analogous cobra and vulture headdress emblems, and crook and flail symbolism. Ibrahim describes the busts’ likeness thusly:
our Puntite personages tell a different and more reliable story[…] The oval face, thin lips, aquiline nose and the slender feature are all typically Somali.[…]
Of the many convincing features to be seen in our Puntite artifacts, I want to present the below figurine (fig. 6) which carries two symbols of authority among ancient Egyptians. The crook (heka) and the flail or flabellum (nekhaka), are two of the most prominent items in the royal regalia of ancient Egypt.[…] Now let us compare it with the next picture (fig. 7) which stands for King Tutankhamun. Both figurines display the two sticks. Again both of them carry on their heads the cobra snake and the vulture as a symbol of the deities Wadjet and Nekhbet. The royal cobra (uraeus) was worn by the pharaohs over their brows. It was thought to spit fire at the pharaoh’s enemies. Another great similarity of the two figurines is the divine osird beard which in death, the kings were frequently portrayed wearing it.[…]
Another common similarity between the Puntites and Egyptians is the headgear which is clearly seen in the below Puntite statuette found in Hargeisa. Again the cobra snake and vulture are mounted on their forehead while the osird beard is attached to the chin.
Most of the statuettes that were dug up appear to depict male personages. However, at least one is manifestly that of a woman. The bust in question (shown above) has the same standard “Caucasoid” features and oval face shape, but her head is more narrow and her nasal bridge and visage are exaggerated in length. This figure is also wearing a headpiece that looks very similar to the cap-crown donned by the Egyptian female Pharaoh Nefertiti (r. 1370–1330 BCE), which suggests that she may represent a Puntite queen. Other conspicuous similarities between the excavated Puntite statuettes and ancient Egyptian figurines include the practice of artificial cranial deformation, which was evident on certain other busts, as well as worship of deities from the Egyptian pantheon, like Sobek. On the apparent artificial cranial deformation, Ibrahim (2013) states:
Head elongation was a common feature among Egyptians. But in comparison with the Puntites found so far — many of them showing a vertical elongation and even sometimes made into the form of a crocodile — the Egyptians had a somewhat horizontal elongation. The strange cranium formation may not have been natural but rather something induced and born from constant applying of external force by nursing mothers. Cranial deformation was a cultural preference among many races around the world and was probably performed to signify group affiliation, or to demonstrate social status.
Furthermore, Ibrahim (2013) remarks that one of the exhumed Puntite sculptures “has a cross-shaped hair left unshaven on the scalp whereby a band of hair links one ear to the other, and another band from the forehead to the back of the head.” Door-Harder and Vogt (2012) write that in the traditional Coptic culture of Egypt, when a child is to receive his/her first haircut or has attained a given age, the child often has his/her hair shorn in a similar cross-shaped pattern. Hence, we see here once more a notable connection between the excavated Puntite statues and the Egypto-Nubian civilization of the Nile Valley.
Illustration of Oromo/Galla men, with one figure wearing a traditional cone-shaped hat. Note the close resemblance between this headdress, the conical headdress on the excavated Puntite statue, and the hedjet of ancient Egypt.
Although Ibrahim did not manage to date the artifacts, particular headdress styles present on several of the figurines give us a rough idea of when they were likely made. For starters, one of the sculptures, an anthropomorphic statue of a male, is wearing what appears to be the hedjet or White Crown of Upper Egypt. This headdress resembles the cone-shaped hat traditionally worn by Oromo/Galla men in the Horn region (see illustration above). The hedjet predates the unification of Upper Egypt and Lower Egypt by the Pharaoh Narmer (Menes), the last king of the predynastic period and founder of ancient Egypt’s First Dynasty (c. 3100 BCE). Several of the male Puntite statuettes are also donning the nemes, a headcloth topped on the brow by the cobra (uraeus) and vulture (nekhbet) emblems. The nemes was the most commonly depicted royal headdress in ancient Egyptian art from the Old Kingdom onwards (cf. Russman (1997)). Ergo, these Puntite busts seem to have been hewn either during the predynastic epoch or in the Early Dynastic period, though later manufacture cannot be definitively ruled out.
Aerial view of three of the Puntite statues excavated at Gol Waraabe. The bust in the center is the same one as shown above from the front (BBC).
In 2018, Robert Kluijver, curator of the Museum of Contemporary Ancient Arabia, announced the discovery of a second batch of ancient statues in northwestern Somalia. Kluijver apparently procured the artifacts from a private seller, who had come across the objects while digging on his farm near Berbera. The excavated materials include realistic busts of a white hue, which appear to have been made of limestone; abstract figures of a reddish color, possibly crafted from sandstone; and blocks of stone containing inscriptions. According to Kluijver and specialists who he showed the objects to, the Berbera statues (like those previously found in Hargeisa valley) do not correspond with any of the known styles in ancient Yemeni art, nor do they resemble the figurines produced by the Axumites. He estimates that the artifacts are 2000 years old. This would make them contemporaneous with the ancient “Berber” city-states described in the Periplus of the Erythraean Sea, including the northwestern port of Malao. The latter market-town was located in the vicinity of present-day Berbera.
Among the recovered artifacts, the limestone busts feature most of the artistic elements first seen in the Gol Waraabe statues: stoic figures with long faces, orthognathous profiles and narrow, moderately projecting noses. At least one of the statues also appears to have had the Osiris or Puntite beard attached, though this Pharaonic appendage has since broken off. Kluijver postulates that this figure may therefore represent a prince. With regard to the inscriptions, they are in an unknown script. Kluijver speculates that the writing may be an early form of Arabic or Phoenician. However, it is more probable that these engravings are yet another instance of the mysterious ancient Somali orthography, which Révoil and others encountered in northern Somalia.
Statuette of the ancient Egyptian Pharaoh Thutmoses III, excavated by Carl Peters at the old stone ruins in Zimbabwe (Rhodesia). According to Ibrahim (2013), this territory was a sourcing area for the Egyptian-related Puntites, where they would often sojourn to obtain gold and other trading items. The Puntites may therefore have left the object behind during one of these commercial expeditions (Peters (1902)).
It is interesting to note at this juncture that in 1899, during his final expedition to the old stone ruins in Zimbabwe (formerly Rhodesia) — a territory which Ibrahim (2013) suggests was a distant sourcing area for the Puntites, where they would periodically sojourn to obtain gold and some exotic trading products — the explorer Carl Peters disclosed that he had found a similar ancient Egyptian statue. This sculpture (shown on the right) was inscribed with what appeared to be hieroglyphic symbols. Shortly afterwards, Peters submitted the object to the Egyptologist Flinders Petrie for inspection and appraisal. Petrie confirmed that “the figure is certainly genuinely ancient.” He noted further that on the statuette’s chest was an engraved cartouche of the Pharaoh Thutmoses III (Tahutmes III), the nephew of the Queen Hatshepsut. Petrie opined that the item therefore probably belonged to a courtier of this 18th Dynasty ruler (Keane (1901)). Peters’ discovery was met with ambivalence among the intelligentsia of his day. Although most pundits recognized the authenticity of the artifact itself, they expressed uncertainty and puzzlement as to how in the first place such a statue had wound up in its remote place of excavation (cf. Keane (1901); The Independent (1901)).
Around the same time period, J. H. Patterson (1907) reported the finding of comparable Egyptian-related antiquities in Mombasa, a coastal town in Kenya. Before the arrival of Bantus/Nilotes from the interior (c. 10th century CE), this area was part of the Iron Age territory of Azania or Po-Pa-Li, which was the realm of the Cushitic Azanians (see Who were the ancient Azanians?):
The town of Mombasa[…] is supposed to have been founded about A.D. 1000, but the discovery of ancient Egyptian idols, and of coins of the early Persian and Chinese dynasties, goes to show that it must at different ages have been settled by people of the very earliest civilisations.
(For more details, refer to Could the ancient “Puntite” statues that were excavated in Somalia actually be forgeries?.)
Legacy
A Galla woman (Sergi (1901)).
Considering the importance of the Land of Punt within the ancient Egyptian ethnocultural complex, the territory left behind a substantial legacy. Quite understandably, it has figured prominently in most anthropological and linguistic reconstructions that attempt to better understand the ancestral relations between the Afro-Asiatic-speaking populations in Northeast Africa. This scholarly tradition is exemplified in the work of Flinders Petrie, leader of the Archaeological Survey of Egypt. The pioneering Egyptologist maintained that the dynastic Egyptians may have originated from an ancient “Hamitic” population centered in the Horn:
The Gala are the remnant of an ancient Hamitic people who appear to have come from North-east Africa, now Somaliland, the region which is most probably to be identified with the land of Punt. It seems, also, that from the same stock which produced the Gala came the dynastic Egyptians, as I have suggested (ANCIENT EGYPT, 1926, p. 10). This is attested, among other things, by the proofs of the Gala origin of the XIIth dynasty (ANCIENT EGYPT, 1924, pp. 38-42).
In addition, Punt has been commemorated in state insignia, literature and philately. The government of Somalia accordingly indicates in its official documentation that:
Somalia was called «Land of Punt» by the Pharaohs, who believed it to be the land of the gods they then used to worship[…]
Ethnically, the Somali people belongs to the Hamitic group, which is further subdivided into:
a) the so-called «Northern Hamites», i.e. the inhabitants of North Africa, namely Libya, Tunisia, Algeria and Morocco;
b) the so-called «Eastern Hamites», i.e. the inhabitants of Somalia and Egypt.
Along with the Egyptian government, the colonial authorities in Italian Somaliland also issued stamps in honor of Pharaoh Hatshepsut’s expedition to Punt during the New Kingdom. In 1998, an autonomous region in northeastern Somalia, Puntland, was likewise named after the territory.
Perhaps the most salient legacy of Punt is the impact that the ancient land has had on the Eritrean national consciousness. It has sometimes been hypothesized that the seeds of the independence movement in Eritrea date back to the medieval kingdom of Medri Bahri, which was a distinct polity from the Solomonid dynasty based in northern Ethiopia. However, textual evidence suggests that these roots actually trace their origin much earlier, to the Puntite era. Because the king Zoscales held sway over Adulis in Eritrea, which in later centuries became the main port of the Axumite kingdom, certain scholars have conflated him with one Za-Haqale, a ruler of Axum. This association is unlikely, though. As George W. B. Huntingford remarks, the Abyssinian king list where Za-Haqale appears was composed retroactively, during the Middle Ages, and is generally apocryphal (e.g. several of the kings on the list are said to have reigned for over a hundred years, which biologically-speaking is improbable). He, moreover, states that “Zöskalés has often been identified with one Haqlé or ZaHaqlé of Conti Rossini’s king-list C, though this name is found only in this list. But these lists are most untrustworthy, and there is nothing in the PME to suggest that Zöskalés was king of Aksum.” Huntingford therefore proposes that Zöskalés just governed the coast on behalf of the king of Axum, to whom he would submit tribute. Even so, Lion Casson notes that this theory is doubtful since the Periplus Maris Erythraei (PME) or Periplus of the Erythraean Sea specifically refers to Zoscales as a king rather than as a subordinate ruler:
A number of commentators hold that [Zoskales] was but a local ruler, in charge of the coastal area as a subordinate of the king of Axum[…] But this view overlooks the fact that the author of the Periplus calls such subordinates tyrannoi (see under B 2:1.9-10), whereas he clearly refers to Zoskales as a king (5:2.19: basileuei, “rules as king”; 6:2.33: “for [the] king”.
Furthermore, as already seen, the Periplus only indicates that Zoscales was a paramount chief in Barbaria; not in Axum per se. It also differentiates between Adulis and “the city of the people called Auxumites.” The former was a three-days’ journey from the interior town of Coloe (present-day Qohaito), whereas the latter was a five-days’ journey from there. Epiphanius of Constantia further bears witness to this when he explicitly distinguishes between the Adulites and Axumites. Casson writes:
There is a third possibility — that Zôskalês was king not of Axum but of an independent realm centered on Adulis and embracing the coastal areas to the north and south. In a paper presented at the Colloque de Strasbourg, 24-27 juin 1987: L’Arabie préislamique, G. Fiaccadori, on the basis of the later history of Adulis and its environs, argued that the area previously must have been independent; it follows that Zôskalês would have been its ruler at the time the Periplus was written. Epiphanius of Constantia (4th century A.D.), in a list of peoples living along the African shore of the Red Sea in the third century A.D., includes Azomitorum cum (A)dulitibus, “the Axumites along with the Adulitans” (Corpus Scriptorum Ecclesiasticorum Latinorum 35, p. 478; cf. Desanges 346); this would indicate not only that Axum and Adulis were separate but that they long remained so.
That Adulis (Aduliton) was, at its inception, not yet under Axumite control is additionally confirmed by Pliny the Elder. The Roman scholar avers that the city was founded by ancient Egyptian deserters, who had moved southwards and established a new settlement thereabouts:
Five days journey from Ptolemais is Aduliton, a city built by the Egyptian deserters.
Zoscales thus reigned from a seat at Adulis, which, like the rest of Barbaria, was originally independent from Axum. Correspondingly, the dominant paternal haplogroup among the Tigre, who are the present-day inhabitants of the Adulis vicinity, is the same Egyptian-affiliated E1b1b-V32 clade that is common elsewhere in the former Berber/Puntite area. In fact, due to their Cushitic origins, most Ethiosemitic speakers (especially those inhabiting Eritrea) belong to the E1b1b paternal haplogroup: Tigre (up to 100%), Tigray-Tigrinya (up to 72%), Amhara (45%) (cf. Trombetta et al. (2015), Supplementary Table 7). The Semitic-mediated J lineage is instead more frequent toward the southern interior in Ethiopia, where it climaxes among North Omotic-speaking populations: Shekecho (52%), Kefa (38%), Yem (32%) (Plaster (2011)). This clade reaches its next highest prevalence in Ethiopia’s erstwhile Axum region, in agreement with a Sabaean origin for the separate Axumite empire. Having said that, there is evidence that at least some Pharaonic cultural elements did penetrate the latter kingdom; likely via Adulis or Meroë. The 18th century explorer James Bruce reported having witnessed stelae in Axum that were engraved with figures of the ancient Egyptian deity Horus (cf. van de Walle (1953)). Although these stone slabs unfortunately no longer exist, the revelation that they once did serves to further highlight the bonds between the Afro-Asiatic-speaking populations in the Horn and Nile Valley.
An African American man bearing a close resemblance to an ancient sculpture from the Ife culture. This likeness underlines the West African ancestral origins of African Americans.
The way forward
In conclusion, a holistic examination of the data on Punt emphatically locates it in Northeast Africa. That is, the hieroglyphic, botanical, craniometric, genetic, cultural and geological evidence indicates that the territory was situated in a broad region encompassing northern Somalia, Djibouti, the Eritrea/Ethiopia corridor and northeastern Sudan, with the ancient Egyptians visiting or writing about different parts of this expansive area at different times.
For starters, the names which the ancient Egyptians used to designate their Puntite counterparts (viz. brbrta and khebsi) both associate the latter with Cushitic peoples. The famous 18th Dynasty Egyptian expedition to Punt organized by the Pharaoh Hatshepsut, moreover, clearly alighted on the incense-bearing hills of Alula (Acannae) in northeastern Somalia, as this is the only place where the higher grade of frankincense known as ‘ntiyw (Boswellia frereana) grows on “terrace” land formations near the seashore (i.e., these are the “Frankincense Terraces of Punt” alluded to on the edifices at Deir el-Bahri). Likewise, Pliny the Elder informs us that the Pharaoh Sesostris I (Senusret I) ordered an expedition to the port of Mossylum. We know from the Periplus of the Erythraean Sea that this cinnamon emporium was located in the present-day Bosaso area in northeastern Somalia. What’s more, the Puntite tributaries shown on the Grand Procession at Thebes likely arrived from Eritrea or Ethiopia since, other than Nubia, these are the only locales today where Egyptian-related peoples (viz. Cushitic, Ethiosemitic and North Omotic speakers) and Nilotes like those depicted on the mural can still be found living in close proximity to each other. Furthermore, isotopic and paleogenetic analyses of ancient baboon mummies as well as geochemical analysis of obsidian fragments, all of which were brought back from Punt to Egypt, indicates that Eritrea is the most probable area from where these specimens and artifacts were imported. (*N.B. The scientists did not examine obsidian fragments from Djibouti or Somalia. The conclusion they reach is therefore tentative.) Eritrea is also the most convincing location for Bia-Punt (“Mine(s) of Punt”), the gold-mining district(s) of Punt, as it is the lone territory in Northeast Africa and the Arabian peninsula whose entire geological formation consists of old metamorphic (Precambrian) rocks, which are associated with gold-yielding areas. Additionally, a 26th Dynasty stela recovered at Dafnah, close to the Egyptian Delta, states that “when rain falls on the mountain of Punt, the Nile floods” — a clear allusion to the northern Ethiopian highlands, near the source of the Blue Nile at Lake Tana.
With the above established, archaeological excavations on a larger scale must hereafter be conducted in order to begin to understand ancient Punt’s history. Who, for instance, are the kings and queens that appear to be represented on the Puntite statuettes, which were exhumed at Gol Waraabe and near Berbera? For how long did these nobles reign and under what circumstances? What was their royal order of succession and was it hereditary? Was Adulis their original seat? Or were they alternately, at different times, domiciled in Alula and other cities within the greater Barbaria?
Ransom (1978) indicates that Pharaoh Hatshepsut drew inspiration from a monument that her expedition had observed while in Punt, and used that shrine as a basis for her own temple at Deir el-Bahri:
When Hatshepsut returned, she built a temple patterned after the one she had seen in Punt. She even referred to the construction as building a “Punt”; and the reliefs on one wall were devoted to describing the trips to Punt. The temple was called “The Most Splendid of Splendors” and the remains are still located at Deir el-Bahari near Thebes. Many comments have been made concerning the apparent fact that the architecture does not fit the standard traditional Egyptian style.
According to Trigger et al. (1983), Hatshepsut’s party constructed another temple during their two or three month sojourn in Punt itself, and dedicated that shrine to the queen and the deity Amun. All this considered, more extensive digging in the Puntite area may eventually also yield the remains of monuments that are similar, if not identical, to those that were erected in ancient Egypt.
11 Saturday Mar 2023
Tags
A-Group, Afro-Asiatic, Ala111Thr, ancient DNA, Ancient Egypt, Badarian, biogeographical analysis, Brenda Stoessiger, C-Group, Capsian, Carleton Coon, Cushitic, Daniel Stiles, EDAR, Ethiopia, G. P. Rightmire, Hamitic, Harold C. Fleming, Iberomaurusian, Indus Valley, Kiffian, lactose tolerance, Linearbandkeramik, Mediterranean, Meroe, Mota, Nella Puccioni, Omotic, Out-of-Africa, Pastoral Neolithic, Richard Leakey, Robert Gayre, Samuel George Morton, Savanna Pastoral Neolithic, Semitic
Scientists often regard Ethiopia as the cradle of humanity. In the 1960s, an archaeological expedition under Richard Leakey discovered skeletal remains at the Omo-Kibish sites near the Omo river. Since these fossils show some modern features and have been tentatively dated to almost 200,000 years before present, many paleontologists argue that they represent the earliest anatomically modern humans (Homo sapiens sapiens) found so far. Other scholars dispute this association, remarking that the skulls also possess archaic characteristics; particularly the Omo II specimen, which has retained the robusticity of earlier, non-modern hominids.
Various evolutionary theories on the origins and dispersal of modern humans (Groucutt et al. (2015)).
The “Out-of-Africa” (OOA) or “Recent African Origin” (RAO) theory emerged as one of several competing hypotheses seeking to explain the prehistoric peopling of the world. Although initially born out of a hoax, complete with false dating, the OOA model gained popularity in the 1990s with the development of the human paternal (Y-DNA) and maternal (mtDNA/mitochondrial DNA) phylogenetic trees. Researchers observed that the deepest or oldest uniparental lineages on the respective trees, paternal haplogroup A and maternal haplogroup L0, are mainly restricted to small Khoisan groups inhabiting Southern Africa. They also found that most populations outside of Africa carry younger maternal lineages that were prehistorically derived from the L3 macroclade (“Eurasian Eve”). Although widely diffused both within and outside of Africa, this haplogroup today has its greatest diversity in Ethiopia. Consequently, L3 is assumed to have been in the area for at least several millenia i.e. long enough to have evolved various sublineages, and thus likely to have originated in the region. Ethiopia is therefore often considered the most probable starting point of the suggested Out-of-Africa colonization.
For these and other reasons, ancient DNA from human fossils in the area holds the potential to greatly improve our understanding of global prehistory.
Mota
In 2015, a genetic research team led by M. Gallego Llorente and E. R. Jones managed to successfully extract ancient DNA from a human skeleton found in Mota Cave, located in the Gamo highlands of southwestern Ethiopia. The Mota remains belonged to a middle-aged male hunter-gatherer, and were radiocarbon dated to around 4,500 years before present:
Low contamination rates were observed, confirming the authenticity of the extracted DNA. Further examination of the fossil’s Y-DNA and mtDNA assigned him to the paternal haplogroup E1b1 and the maternal haplogroup L3x2a, respectively:
To more closely gauge Mota’s population affinities, the scientists also ran a principal component analysis comparing his DNA against that of various contemporary Ethiopian populations from different ethnolinguistic groups. The specimen was most similar to the South Omotic-speaking Ari and the Khoisan-speaking Sandawe populations:
The close association between Mota and the modern Ari was also supported by f3 statistical analysis, which showed that they formed a clade unto themselves.
In order to ascertain whether Mota harbored any West Eurasian ancestry like modern Ethiopian populations, the researchers ran an admixture analysis using the Yoruba and Druze as the African and West Eurasian reference samples, respectively, against which Mota’s DNA and that of other contemporary populations was compared. The results suggested that Mota lacked any West Eurasian ancestry. This was also supported by the fact that the specimen did not carry the derived SLC24A5 (Ala111Thr/rs1426654) allele linked with lighter skin pigmentation, nor any lactase persistence variants, nor apparently any Neanderthal-associated alleles.
So what can be concluded from this ancient DNA analysis? Are the findings significant, or is Mota simply an early Ari individual and little beyond that?
The results are quite illuminating in that they suggest, among other things, that:
Limitations of biogeographical analysis
Before proceeding further, let us briefly note some of the principal limitations of biogeographical analysis, such as that around which the Mota study is centered. BGA/admixture testing is at its core speculative, for it is based on probability. That is, such analysis estimates where or in which geographical area given stretches of DNA known as SNPs likely originated according to which reference populations today carry those SNPs at highest frequencies. This is problematic for a number of reasons, some quite obvious:
Besides the points above, perhaps the biggest limitation of biogeographical analysis is the fact that it can only at best capture a small fraction of an individual’s total ancestry i.e. that contained within the genetic tree, not that within the exponentially larger genealogical tree. The Genetic Genealogist explains:
In reality, everyone has two family trees. The first is a Genealogical Tree, which is every ancestor in history that had a child who had a child who had a child that ultimately led to you. Every decision made by every person in that tree contributed to who and what you are today.
However, not every person in that tree contributed a segment of your DNA sequence (because of random inheritance, as discussed above). As a result, we have a second family tree – a Genetic Tree – which is a tree that contains only those ancestors who contributed to our DNA. No one has yet been able to construct their Genetic Tree, but soon it will be a reality thanks to advances in genetic sequencing and comparison such Relative Finder. These tools are using relatedness between people living today to deduce the inheritance of DNA from people who have been dead for centuries.[…]
The Genealogical Family Tree contains ALL of your biological ancestors[…] The Genetic Family Tree contains a small subset of your biological ancestors[…] Due to the nature of the Genealogical versus the Genetic Family Tree, entire populations, ancestors, and ethnicities are regularly lost entirely from your DNA! [They] therefore would not be detected by a DNA test.
Uniparental markers and Out-of-Africa
The Mota specimen’s paternal haplogroup E1b1 is today relatively rare, and is mainly restricted to a few Afro-Asiatic speakers in Ethiopia and environs. It has been found amongst 18% of Ethiopian Jews, 11%-12.8% of Oromos, 11% of Iraqw, 6%-10.4% of Amhara, 10% of Ethiosemitic speakers generally, and in 18.2% of Ethiopians as a whole. Given the clade’s close association with Afro-Asiatic-speaking populations in Ethiopia, it appears that Mota’s ancestors obtained the E1b1 haplogroup through contact with early Afro-Asiatic male settlers in the area.
Mota’s L3x2a maternal lineage points to a similar, if more ambiguous, affiliation. It too is today mainly concentrated among some Afro-Asiatic speakers in East Africa, and is also present in Egypt and among Yemeni Jews. How exactly Mota’s hunter-gatherer culture acquired the haplogroup is uncertain, for Babalini et al. (2002) found a comparably-aged L3 carrier in their ancient DNA study of early human specimens from the Fezzan in Libya (dated to around 3,000-1,500 BCE). Fernández et al. (2014) likewise identified an even older L3-bearing individual in their examination of skeletal remains from Pre-Pottery Neolithic B (PPNB) sites in the Near East. Dated to between 8,700-6,600 BCE, the PPNB specimen predates Mota and the Fezzan individual by over four thousand years. This highlights the early global distribution of the L3 haplogroup, if not an ultimately Middle Eastern origin for the clade, as Farrell et al. (2013) propose:
Here we present the first high-coverage whole genome sequences from a Middle Eastern population consisting of 14 Eastern Province Saudi Arabians. Genomes from this region are of interest to further answer questions regarding “Out-of-Africa” human migration. Applying a pairwise sequentially Markovian coalescent model (PSMC), we inferred the history of population sizes between 10,000 years and 1,000,000 years before present (YBP) for the Saudi genomes and an additional 11 high-coverage whole genome sequences from Africa, Asia and Europe.
The model estimated the initial separation from Africans at approximately 110,000 YBP. This intermediate population then underwent a long period of decreasing population size culminating in a bottleneck 50,000 YBP followed by an expansion into Asia and Europe. The split and subsequent bottleneck were thus two distinct events separated by a long intermediate period of genetic drift in the Middle East. The two most frequent mitochondria haplogroups (30% each) were the Middle Eastern U7a and the African L. The presence of the L haplogroup common in Africa was unexpected given the clustering of the Saudis with Europeans in the phylogenetic tree and suggests some recent African admixture. To examine this further, 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.
These results are consistent with the hypothesis that modern humans populated the Middle East before a split 110,000 YBP, underwent genetic drift for 60,000 years before expanding to Asia and Europe as well as back-migration into Africa. Examination of genetic variants discovered by Saudi whole genome sequencing in ancestral African populations and European/Asian populations will contribute to the understanding human migration patterns and the origin of genetic variation in modern humans.
Thus, the prehistoric “Out-of-Africa” colonization, assuming one occurred at all, does not appear to have emanated from the Horn. The finding that the L3 haplogroup likely originated in and spread from the Middle East points instead to some other center of evolution. (*N.B. For the latest evidence on the suggested non-African origin of the mtDNA macrohaplogroup L3, see Cabrera (2022) and Cabrera et al. (2017).)
North Omotic vs. South Omotic
Kawo Tona, the last King of the Wolayta. Prior to assimilating Mota-related foragers, most Omotic speakers had a Cushitic physiognomy like that of this late ruler. Nowadays, this phenotype is mainly confined to North Omotic speakers.
The Omotic branch of Afro-Asiatic is divided into two subgroups: North (also known as Nomotic) and South (or Somotic). South Omotic consists of the Ari, Dime, Hamar, Gayil and Karo languages. Of these South Omotic idioms, the first three are sometimes collectively known as “Aroid”.
Anthropologists and linguists working in the Ethiopian region have long observed a marked physical and linguistic cleavage between, on the one hand, the speakers of the Aroid Omotic languages, and on the other, the non-Aroid Afro-Asiatic-speaking populations. Harold C. Fleming, who coined the term “Omotic” and helped establish the validity of the phylum as an independent branch of the Afro-Asiatic family, remarks that the Ari are generally “Negroid” in physiognomy, in contrast to the “Ethiopid” northern Cushitic and Ethiosemitic groups:
The Ari peoples have been described extensively by scholars of the Frobenius tradition, especially A. Jensen, Eike Haberland, and others, as well as observed and described more informally by various anthropologists (e.g., Herbert Lewis, Jean Lydall, Ivo Strecker, myself) and Ethiopian government officials. One of our colleagues, Ayyalew Mitiku, is an Amhara who grew up among the Ari and speaks their language fluently. Among all these observers there is a consensus that the Ari and many of the Banna and Hamar represent a variety of African Negro, rather than an Ethiopid or “Afro-Mediterranean” variety. However, the Ari variety of Negro is distinctive and is not very much like the neighbouring Nilotic or Surmic peoples to the west and south; nor particularly like the Bantu of East Africa. Some resemblance is noted to the Koman of the Ethiopia-Sudan border areas and some Sudanese populations. Above all this aspect of the Ari, while it has earned them the label of “blacks” or “Shanqillas”, strikes many of us as being the appearance of a distinctive kind of African, a population that has evolved on its own to a considerable extent.
Due to the Ari’s aberrant morphology and the divergent nature of the Aroid languages themselves, Fleming further indicates that Ari and the other Aroid languages were regarded by earlier scholars as non-Afro-Asiatic idioms possessing Nilo-Saharan affinities:
The three main branches of my Somotic, viz, Dime, Hamar and the Ari cluster are what [Lionel Bender] calls Aroid; they are so different from the rest of Omotic that older German and Italian scholars thought of them as ‘Negroid’ or Nilotic or the like. It was a struggle to get them accepted as a branch of Omotic or West Cushitic. All you have to do is work on a Dizoid (Maji, Na’o, Shako) language and then on Dime or Galila (Ari), as I did, to see that they are very different in phonology and morphology. Yes, there has been some borrowing across the Omo river. Dime informants will tell you that “Maji” used to rule them; the borrowings are not massive, however. Dizoid and ‘Aroid’ simply do not belong in the same moiety in opposition to the rest of Omotic.
Correspondingly, in the ethnological literature, the traditional explanation for the Ari’s differing physiognomy and culture as well as the divergent nature of their South Omotic language is that they are descendants of Nilo-Saharan-speaking peoples (the Pre-Nilotes), who settled in the parts of Ethiopia bordering South Sudan around 4,000 years ago. These Pre-Nilote groups are then thought to have interbred with early Omotic populations from the Sahara and gradually adopted the latter’s Afro-Asiatic languages, thereby giving rise to the Ari and related groups. Per the Centre National de la Recherche Scientifique:
Negroid people from the Sudan, speaking languages ancestral to those classified as the four branches of the Nilo-Saharan family settled in West and South West Ethiopia about the third millenium B.C. (Pre-Nilotes). They became ancestors of peoples living on the western border and intermixed with Omotic speakers to form the Ari, Basketo, Dimi and Gimira-Maji groups of tribes. The languages spoken in Ethiopia in the third millennium B.C. are derived from a single stock (Hamito-Semitic or Afro-Asiatic) originating in the Eastern Sahara.
Dermatoglyphic affinities of Afro-Asiatic and Nilo-Saharan-speaking populations in Ethiopia. Fingerprint patterns, which are genetically inherited, indicate that the North Omotic-speaking Shinasha share ties with the other sampled Afro-Asiatic-speaking individuals (Cushitic-speaking Oromo and Ethiosemitic-speaking Amhara and Tigray) rather than with the Nilo-Saharan-speaking individuals (Berta). This again suggests that the Shinasha and other North Omotic speakers directly descend from the original Omotic settlers of the Horn (Yohannes and Bekele (2015)).
Genomic analysis of various populations in Africa and Europe. The North Omotic-speaking Shekkacho sample has a similar level of West Eurasian ancestries (Afro-Asiatic & Iberian-related components) as the other Afro-Asiatic-speaking populations and Nubian groups in Northeast Africa. Although bearing some Mota-related admixture like the Ethiosemitic-speaking samples (Amhara & Tigray), the Shekkacho primarily belong to the Afro-Asiatic ancestral component, which peaks among Cushitic-speaking Somalis. This element is analogous to Hodgson et al. (2014)‘s “Ethio-Somali”, an inferred ancestral component that the scientists demonstrated via a battery of tests to be essentially West Eurasian. The foregoing supports the view that the earliest Omotic settlers in the Horn region were, like the ancient Cushitic pastoralists in the Great Lakes area, transplants from North Africa. As also observed by López et al. (2021), the North Omotic-speaking Shekkacho, Shinasha and Wolayta thus represent pockets of Omotic speakers who retained much of their original Afro-Asiatic ancestry, whereas this ancestry was largely diluted in South Omotic-speaking areas through intermixture with and assimilation of local hunter-gatherers. Since Omotic is regarded by most linguists as the first diverging branch of the Afro-Asiatic language family, this analysis holds implications for the population affinities of the Proto-Afro-Asiatic speakers (Gopalan et al. 2019)).
Genetic differentiation
Besides morphology and language, the postulation that the Pre-Nilote forebears of the Ari (who are presumably represented by the Mota specimen) were of a different ancestral stock than the progenitors of the Afro-Asiatic-speaking Ethiopian groups is strongly supported by genetic studies on these contemporary populations.
For starters, researchers have observed a high frequency of the derived SLC24A5 allele (Ala111Thr or rs1426654) among various Cushitic- and Semitic-speaking Afro-Asiatic populations in Ethiopia. This mutation is closely associated with lighter skin pigmentation, and is believed to have originated in or near West Asia. Almost 60% of Ethiopian Jews and ethnic Somalis from Somalia carry the variant. et al. (2015) found a similarly high percentage of the allele among the North Omotic-speaking Wolayta. By contrast, only 12% or so of the South Omotic-speaking Ari ironworkers possess the Ala111Thr polymorphism. Since the Mota specimen does not harbor the mutation, this suggests that the ancestors of the Afro-Asiatic-speaking groups in the Horn were responsible for having introduced the allele into the Ari gene pool. This, in turn, implies that these early Afro-Asiatic settlers were of a lighter complexion than the Ari’s Mota-like forebears.
Lactase persistence (LP) frequencies among various Afro-Asiatic-speaking Ethiopian populations and adjacent groups (Jones et al. (2015)).
Further evidence that the Mota specimen was clearly not ancestral to the local Afro-Asiatic-speaking populations can be seen through comparative lactase persistence (LP) analysis. While Llorente et al. observed that Mota does not carry any lactose tolerance alleles, such LP mutations are found at high frequencies among the region’s main pastoral groups. To this end, Jones et al. (2015) note that over 60% of Beja, Afar and Borana Oromos are lactase persistent, carrying several different LP variants. In their large study of lactose tolerance in eastern Africa, Tishkoff et al. (2007) likewise find widespread lactase persistence among not only the Afro-Asiatic-speaking populations, but also among certain Nilo-Saharan and Niger-Congo pastoralist groups (like the Datog Nilotes) that are known to have absorbed some early Southern Cushites in the Great Lakes region.
Various analyses showing negligible-to-low autosomal DNA affinities between the Ari and adjacent Afro-Asiatic-speaking populations (Vandorp et al. (2015)).
Most tellingly, the Afro-Asiatic-speaking populations in the Horn appear to have generally different autosomal DNA (auDNA) signatures than the Ari. Hodgson et al. (2014) observed that ethnic Somalis, Afar, Amhara, Tigray and Oromos are defined by a West Eurasian-affiliated ancestral component, which they refer to as “Ethio-Somali”. On the other hand, the Ari are defined by a separate, Nilo-Saharan-affiliated ancestral component, which the researchers call “Ethiopic”. The Wolayta, despite possessing comparatively greater Mota-like admixture, were again found to be closer overall to the Cushitic- and Semitic-speaking populations than to the Ari. Dobon et al. (2015) further confirmed the existence of a defining West Eurasian ancestral element among the Afro-Asiatic speakers in Northeast Africa. Vandorp et al. (2015), in turn, reaffirmed the lack of close ties between the Afro-Asiatic-speaking populations and the neighboring Ari.
Ethnolinguistic distribution of maternal lineages in the Horn, Nile Valley, Sahara, Maghreb, Great Lakes and the Arabian peninsula (Boattini et al. (2013)).
Similarly, Boattini et al. (2013) observed that the mtDNA of the South Omotic-speaking Dawro-Konta and Hamer in southern Ethiopia, who are closely related to the Ari, is tied to that of Nilotic and Bantu populations in the Great Lakes region as well as the Ongota (who are believed to have once spoken a Nilo-Saharan language). On the other hand, the maternal lineages of the Cushitic- and Ethiosemitic-speaking groups of the Horn cluster instead with those of other Afro-Asiatic-speaking populations in the Nile Valley, Yemen and Sahara.
Altogether, this is consistent with the aforementioned tradition that the Ari’s “Shanqilla” ancestors (i.e. Mota) were originally Nilo-Saharan speakers, who at some point in antiquity adopted an Omotic language from early Afro-Asiatic-speaking settlers.
Afro-Asiatic ancestral population
In their study of the Mota remains, Llorente et al. note that contemporary Ethiopian populations (specifically, the Afro-Asiatic speakers) have substantial West Eurasian affinities that the Mota specimen does not appear to possess. They therefore conclude that that ancestry likely arrived after Mota’s lifetime, and originated from a Sardinian-like population most similar to the early Neolithic farmers who settled Europe (according to f3 statistical analysis).
In lieu of modern reference populations, the researchers ran an admixture test using the Mota specimen and a Neolithic Linear Pottery culture or Linearbandkeramik/LBK sample (represented by the Stuttgart specimen) as the baseline African and West Eurasian samples, respectively, against which the DNA of the Ari and that of the neighboring Afro-Asiatic-speaking populations was compared. This alone constitutes something of a breakthrough in biogeographical analysis, where disparate, admixed modern groups are instead frequently used as stand-ins for ancient populations (e.g. Yoruba and Utah denizens versus early Africans and West Eurasians). Nonetheless, there remain several problems with utilizing these particular Mota and Linearbandkeramik samples as the ancient proxy groups. Perhaps the most conspicuous issue is that, despite their antiquity, neither the Mota fossil nor the makers of the Neolithic LBK industry are representative of the ancestral Afro-Asiatic speakers in the Horn. We already examined above why that is vis-a-vis the Ari and their apparently Nilo-Saharan-speaking Mota progenitors, so let us now turn to the Linearbandkeramik agriculturalists.
While the LBK farmers may very well have prehistorically contributed some genes to the populations in Northeast Africa (and/or elsewhere on the continent), they do not appear to have been responsible for the majority of the West Eurasian affinities that are found today in the Horn. There are a number of reasons why we can be confident of this. Firstly, although the mtDNA haplogroup N1a, which is quite common today among the Afro-Asiatic-speaking populations in the Horn, is found at high frequencies among Neolithic LBK samples from Central Europe, the particular 7,000 year old Stuttgart sample from Germany that Llorente et al. utilized in their own analysis belongs to the T2 clade (cf. Lazaridis et al. (2013)). This latter maternal haplogroup is, by comparison, relatively rare in Northeast Africa. Unfortunately, since the Stuttgart sample is that of a woman and females do not inherit a Y chromosome, it is unclear whether the situation is the same with regard to the paternal side of things. Haak et al. (2010) did, however, successfully type three Neolithic LBK males for their Y-DNA. These individuals were assigned to haplogroups G2a3 and F*, both of which are also uncommon in Northeast Africa.
Another clue that the Linearbandkeramik agriculturalists are not the main source of the West Eurasian ancestry in the Horn is the fact that local rock art, such as at Laas Geel in northern Somalia, clearly depicts both herders and their domesticated cattle prior to the ~3,000 year old date that Llorente et al. propose as the arrival period of the LBK people. Those cave paintings also have stylistic precedents in the Arabian Peninsula (as explained further here, under ecology, rock art and genetics). Moreover, the Stuttgart woman, like the Mota man, was apparently unable to digest lactose. Lazaridis et al. remark that she lacks any alleles associated with lactase persistence. This in itself is not especially unusual for members of non-herding populations (whether farmers or hunter-gatherers) since they often do not drink cow milk and thus their bodies have no need to continue producing the lactase enzyme passed the breast-feeding infant age. Nevertheless, the finding is at odds with the aforediscussed high frequency of lactose tolerance among the pastoralist Afro-Asiatic-speaking groups and also contrary to the great diversity of the LP alleles present in the region.
The ~3,500 year old Cairn 4 burial excavated by Daniel Stiles, containing the 190 cm/6’4″ skeleton of an early Cushitic male settler (Stiles and Munro-Hay (1981)).
Aside from differing genetic markers and pastoral rock art, archaeology provides the most obvious indication that a West Eurasian-affiliated population — one likely distinct from the LBK farmers — co-existed in the region at around the same period as the Mota man and his kinfolk. In the 1980s, at various oases on the eastern margin of the Chalbi desert, the anthropologist and archaeologist Daniel Stiles excavated a number of cairns belonging to the Cushitic-associated Savanna Pastoral Neolithic (Stone Bowl Culture). The burials yielded the remains of tall individuals of “Caucasoid” physical type, with the oldest such skeleton, a 190 cm/6’4” male, radiocarbon-dated to around 3,500 years before present. By contrast, Llorente et al. estimate the roughly coeval Mota specimen’s stature at only 154.8 cm/5’1” i.e., a whopping 35.2 cm/13.9 inch difference in height! As explained in detail on The Elongated African fallacy (under physiognomy and exotic influences), indirect evidence has also established that, unlike Mota, these early Cushitic settlers possessed both the Ala111Thr allele for lighter skin pigmentation and a lactase persistence mutation as well as non-kinky (cymotrichous) hair texture.
Approximate difference in height between the Savanna Pastoral Neolithic Cairn 4 specimen and the coeval Mota hunter-gatherer specimen (MrInitialMan.com).
Thus, with the discovery, biological examination and dating of the Mota fossil, we now have concrete evidence of the existence of at least two genetically and physically distinct populations residing in the region during the Neolithic: one population ancestral to the Afro-Asiatic-speaking groups and linked with West Eurasians, and the other population ancestral to the Aroid groups and possibly linked with Nilo-Saharans (though the Mota specimen’s diminutive stature, his hunter-gatherer lifestyle, and his and the Ari’s genetic ties with the Sandawe may point to deeper Khoisan relations).
Unidentified East Eurasian elements
In addition to unrepresentative reference populations, another problematic aspect of the Mota biogeographical analysis is its intrinsic assumption that two proxy groups — an African sample and a West Eurasian sample — are sufficient to accurately capture the ethnic composition of the populations in Northeast Africa and elsewhere on the continent. A closer, multidisciplinary look at the broader data, however, points to the presence of at least a third ancestral element; one with apparently East Eurasian affinities.
Morphology and ancient testimonies
Craniometric relationships between various ancient and modern populations (Brace (1993)).
Almost two centuries’ worth of craniometric and anthropometric studies contradict the suggestion in the Mota paper that the Afro-Asiatic-speaking populations in the Horn may have been ancestrally formed through interbreeding between an early Sardinian-like population (the LBK makers) and a Mota-like population. In actuality, rather than morphologically clustering somewhere halfway between these two ancient reference groups — as one would expect for peoples that are supposedly 50% European and 50% African — the Afro-Asiatic speakers show few of the osteological characteristics of actual biracial populations. They instead appear “skeletally Mediterranean” (see, for instance, Brace (1993) to the right; also Kemp (2006) and the other studies here).
Sidamo men of relatively “pure” Cushitic type, closely resembling many Oromos.
The anthropologist Carleton Coon observed this firsthand in his detailed examination of physical types in the Horn, “The Mediterranean Race in East Africa”, a chapter in his influential 1939 work The Races of Europe. By analyzing the Sidamos of southern Ethiopia in particular, who are the actual product of recent intermixture between Cushitic peoples and adjacent Pre-Nilotes, he was able to see what exact morphological changes such hybridization produces. Coon thus concluded that the local “Hamitic” populations were essentially “Caucasoid” and that “Negroid” influence, while also present, was on the whole minor.
Furthermore, Coon was able to identify an additional “non-Negroid” ancestral element, which was especially important among Somalis, Afars/Danakils, Agaus and other Cushitic groups. This swarthy “Veddoid” component, he postulated, may have arrived in antiquity from the Indus Valley by way of Southern Arabia along with the zebu cattle (Bos indicus):
Later than the development of highland agriculture in East Africa was the introduction and diffusion of pastoral nomadism. The cattle complex, with its elaborate set of social restrictions and of social differentiation on the basis of wealth in herds, was introduced from India by way of southern Arabia, along with the humped zebu, at some none too distant period, probably as late as the first millennium B.C.[…]
Our survey of the metrical characters of the inhabitants of the Hamitic racial area has brought several facts to light; the agricultural population of the Ethiopian highlands, both indigenous and imported from Arabia, belongs to a tall, dolichocephalic to mesocephalic, leptoprosopic, moderately leptorrhine race, which is Mediterranean in metrical position and cannot be distinguished, on the basis of the more commonly taken measurements, from blond and brunet Mediterraneans of Europe and North Africa. The Somalis, on the other hand, belong to an extreme racial form; extremely linear in bodily build, extremely narrow-headed and narrow-faced, with a special narrowness of the jaw. The relationship of the Somalis, on metrical grounds, is with some of the peoples of India as much as with the Mediterraneans elsewhere. The leptosome tendency, and the narrowness of the face, remind one of the same tendency found among the mixed Bedawin group of the Hadhramaut. It cannot be attributed to negro-white mixture, for that phenomenon, as witnessed among the Sidamos, has produced a heaping of characters, resulting in an enlargement of both sagittal and lateral diameters of the face, in some cases in excess of either the Hamitic white or the negroid parent. Upper face height and nose height are especially affected. The Somali face and nose are not long, they are merely narrow.
Royal effigy of the Meroitic King Asharramon (Fig. 191) and heads of ancient Egyptian commoners of similar “Austral-Egyptian” type (Morton (1854)).
Coon’s assertion is nothing new, for various early Greek, Roman, Egyptian and Brahmin writers, including Strabo and Philostratus, inform us of ancient colonies of Indus Valley peoples in Northeast Africa. For example, Bahadur (1917) notes that “Eusebius states that Ethiopians emigrating from the River Indus settled in the vicinity of Egypt [Meroë].” Nilus similarly relayed to Apollonius Tynaeus that “the Indi are the wisest of all mankind. The Ethiopians [Meroites] are a colony from them: and they inherit the wisdom of their forefathers”.
Herodotus (c. 440 BCE) indicates that different populations — both light-skinned and dark-skinned — lived in ancient Aethiopia, an area in Africa roughly corresponding with the territory to the south of Egypt and Libya. Among these inhabitants were (cf. Herodotus: The Histories):
Other anthropologists have likewise proposed that an ethnographical tie exists between early Afro-Asiatic-speaking populations in Northeast Africa and the makers of the Indus Valley civilization. Following detailed examination of the skulls and murals of the ancient Meroites in the Nile Valley, Samuel George Morton asserted that “the Austral-Egyptian or Meroite communities were an Indo-Arabian stock, engrafted on the primitive Libyan inhabitants”. He further explained that, while such an influence was also present among commoners in Egypt, it was strongest in Nubia, including among the ruling class:
I observe, among the Egyptian crania, some which differ in nothing from the Hindoo type, either in respect to size or configuration. I have already, in my remarks upon the ear, mentioned a downward elongation of the upper jaw, which I have more frequently met with in Egyptian and Hindoo heads than in any other, although I have seen it occassionally in all the races. This feature is remarkable in two of the following five crania (A, B), and may be compared with a similar form from Abydos[…]
It is in that mixed family of nations which I have called the Austral-Egyptian that we should expect to meet with the strongest evidence of Hindoo lineage; and here, again, we can only institute adequate comparisons by reference to the works of Champollion and Rosellini. I observe the Hindoo style of features in several of the royal effigies; and in none more decidedly than in the head of Asharramon (Fig. 191), as sculptured in the temple of Debod, in Nubia. The date of this king has not yet been ascertained; but, as he ruled over Meroe, and not in Egypt, (probably in Ptolemaic times [B. C. 200-300],) he may be regarded as an illustration of at least one modification of the Austral-Egyptian type[…]
Another set of features, but little different, however, from the preceding, is seen among the middling class of Egyptians as pictured on the monuments, and these I also refer to the Hindoo type. Take, for example, the four annexed outlines (Fig. 192), copied from a sculptured fragment preserved in the museum of Turin. These effigies may be said to be essentially Egyptian; but do they not forcibly remind us of the Hindoo?
The Badarian factor
A typical Badarian male skull (Stoessiger (1927)).
Of all the ancient populations in Northeast Africa, the Badarians of Upper Egypt were most often regarded as the likeliest link between the local Afro-Asiatic-speaking groups and the Indus Valley peoples.
The Badarian culture flourished between 4400-4000 BCE. It consisted of small villages of semi-nomadic agropastoralists, who kept domesticated animals, cultivated grain, produced the first glazed objects, and also used metal. The Badarians interred their dead in cemeteries on the outskirts of their living area. Within the actual graves, the deceased were laid out in the fetal position, facing the setting sun toward the west. They were buried with some clothing items, pottery, jewelry and a fertility idol. Despite its brief existence, the culture is considered archaeologically important since Badarian sites have yielded the earliest evidence of agriculture in Egypt.
In Coon’s examination of Badarian skeletal remains, he observed that they were on the whole quite similar to those of other predynastic Egyptians. He described the Badarians as essentially “Mediterranean” in the anthropological sense, possessing cranial affinities with both the Afro-Asiatic-speaking populations of the Horn (although the Horn groups are ultimately closer to the later Naqadans) and the Dravidian populations of southern India:
From the type site, Badari, come the earliest skulls of a definitely Egyptian group which have yet been discovered. These Badarians lived about 4000 B.C., after the climate had become considerably drier than it was in Tasian times, so dry, in fact, that in many cases the skin and hair of their dead have been naturally preserved. The skin was apparently brunet white, while the hair was black or dark brown in color, thick, of fine texture, and usually wavy in form.
Although the Badarians, like the Tasians and Merimdians, still hunted and fished to enhance their larders and vary their diet, they lived primarily by agriculture and by herding cattle and sheep. Unlike the Merimdians they raised no pigs. By hammering copper they were entering the transition from the Neolithic to the Metal Age. They navigated the Nile in ships, whose shapes are revealed by pottery models, but we cannot be sure that they sailed them. These Badarians were undoubtedly newcomers to Upper Egypt who displaced the Tasians and perhaps other predecessors.[…]
The Badarian type represents a small branch of the Mediterranean racial group. The head is unusually high in comparison to the other dimensions, and the facial skeleton is in the absolute scale unusually small; the mandible is small, narrow and light. Its mean male bicondylar diameter is the smallest known, while the bigonial diameter of 91.6 mm. is also extremely low.[…]
Morant shows that the Badarian cranial type is closely similar to that of some of the modern Christians of northern Ethiopia, who incidentally do not show negroid characteristics in the skull, and also to the crania of Dravidian-speaking peoples of southern India. One might add that living Somalis show a close approximation to this physical type in most respects, and the extremely narrow jaw in which the Badarians seem to reach a world extreme may be duplicated among both Somalis and the inhabitants of southern India. In Europe, the closest parallel to the Badarian type is found among modern Sardinians, but this is not as close as their relationships to outer and later Egyptians.
Coon indicates that the Badarians seem to have eventually been absorbed by the ensuing predynastic Egyptians of Naqada. He also asserts that a Badarian strain persists in the Horn. Despite the prevailing “Mediterranean” element, this lingering influence, he suggests, manifests itself in various ways; particularly through attenuation of the distal or remote segments of the limbs (i.e., small wrists, hands, ankles and feet), as is common among the populations of southern India:
The bodily build of the African Hamites is typically Mediterranean in the ratio of arms, legs, and trunk, but the special attenuation of the extremities among the Somalis is a strong local feature, which finds its closest parallels outside the white racial group, in southern India and in Australia.
In her comprehensive study of Badarian skeletal remains, the anthropologist Brenda Stoessiger (1927) confirmed that the Badarians were closely related with other predynastic Egyptian series, but also bore relations with Dravidian populations in the Indian peninsula. She attributes these bonds to parallel population movements, westward into Northeast Africa and eastward into South Asia, from a common center in or near the ancient Caucasus:
Anthropologists have frequently drawn attention to the similarity in the appearance of the skulls of the Hindu races of India and the Early Egyptians. In a paper “Sur l’Origine de l’Ancienne Race Egyptienne,” published in the first volume of the Mémoires de la Société d’Anthropologie de Paris, pp. 410-422, Pruner-Bey notes this similarity only to reject it; each race, he says, has a skull, small and oval in shape; the body and extremities are for both races beautifully proportioned; but there is a marked difference in the fleshy parts, the Ancient Egyptian resembling the modern Berber, while the Hindu is bronze to bistre in colour. Finally he dismisses the idea of direct relationship on the grounds of linguistic differences, an argument which would scarcely now-a-days be advanced.[…]
From skull measurements alone it would be difficult to choose between the primitive Indian and Egyptian series as the group to which the Badarians are closer. Unfortunately it is not possible to carry the analogy further and find low coefficients between the later Egyptian series and the Indian series as the types diverge in different directions.[…]
This study confirms the conclusions based on cultural and topographical evidence that the Badarian skulls are early Predynastic Egyptian but if anything more primitive in type than the other series of this period, though the mean direct measurements differ very little from them. The early and late Predynastic types however do show a significant difference which, if we may assume the races to be divided by a period of four or five thousand years or more, may be accounted for by slight evolutionary changes or a gradual infusion of races.
How far do these results confirm Sir Flinders Petrie’s theory of a Caucasian origin? When we compare the Badarian race with others outside Egypt, it is not the Mediterranean or any Negro type which it resembles most closely but the primitive Indian, the Dravidian and the Veddah. Thus they do not oppose the suggestion of a common origin in the Caucasus from a race sending one branch westward to Egypt and Europe and another south-eastward to India. To confirm this, however, we should need series of ancient skulls from Palestine, Persia, and Western India.
Genetic and phenotypic oddities
According to Coon (1939), ethnic Somalis (over 86% among northern Darod individuals), Afars and other lowland Cushitic groups possess significantly higher rates of non-kinky hair texture than do the Amhara (40%), Tigray and other highland Abyssinian Semitic speakers. He asserts that the Sidamo, despite their elevated South Omotic admixture, also have a higher rate of non-kinky hair texture than do the Abyssinians. Puccioni (1931) reported comparable findings for the Somali clans in Somalia. Charpin and Georget (1977) similarly observed that only 3.8%-4.9% of the Issa Somalis and the Afars of Djibouti had hair in the kinky/ulotrichous class (cheveux crépus).
Besides the osteological work, ancient testimonials, zebu cattle complex and hair morphology, there are a couple of other indications that an East Eurasian ancestral component likely exists among the Afro-Asiatic-speaking groups in Northeast Africa. Narasimhan et al. (2019) report a high prevalence of the Afro-Asiatic-linked E1b1b paternal haplogroup among Late Bronze Age/Early Iron Age individuals from the Swat Valley in northern Pakistan, specimens which also exclusively bore M and N mtDNA derivatives. This points to ancient ties between this area, Mesopotamia, the Levant, the Arabian peninsula, North Africa and the Horn of Africa (cf. Table S1-S5). More specifically, the presence of haplogroup E1b1b in South/Central Asia this early in time likely reflects eastward incursions by Neolithic and/or Bronze Age Levantines (the original Elamo-Dravidian settlers, perhaps?). These newcomers possibly interbred with local populations, before eventually returning to the Afro-Asiatic fold along with their newly-acquired East Eurasian genes. Such early connections between Afro-Asiatic-speaking populations and groups bearing East Eurasian ancestry are likewise supported by Witas et al. (2013), who analyzed Early Bronze Age to Roman era specimens excavated at Tell Ashara (Terqa) and Tell Masaikh (Kar-Assurnasirpal) in Mesopotamia. The researchers found that these ancient individuals (dated to 2500BCE-500CE) belonged to the mtDNA clades M4b1, M49 and/or M61, maternal lineages that are today concentrated and believed to have evolved in the Indian subcontinent. Various autosomal DNA analyses have also observed an East Eurasian component in their Somali samples (e.g. Kidd et al. (2011); Kidd (2011b); Truelsen et al. (2017); Pereira et al. (2017)). Other factors supporting this association are the elevated frequency of the B blood group among Ethiopian Somali males (44.44%) and Afar males (33.33%) (cf. Abegaz (2021)) as well as among Beni Amer Beja (31%; Corkill (1949)) — a serological system that is also common among Egyptians (24.1%; Swelem et al. (2018)) and Riverain & Shaigiya Sudanese “Arabs” (25% and 24%, respectively; cf. Corkill (1949)) and which globally peaks among populations in South Asia (Kocak et al. (2017)); the high mean number of private alleles that are found today in certain Cushitic groups (cf. Babiker et al. (2011)); as well as the presence in these populations of unusual genetic variants of apparent East Eurasian origin (e.g. the EDAR gene’s derived allele, which is associated with hair thickness among populations in eastern Asia; around 12.5% of ethnic Somalis bear the mutation, whereas it is largely absent in Subequatorial Africa, the Maghreb, West Asia and Europe).
(*N.B. Our genome analysis has also detected an East Eurasian component borne by our Cushitic, Ethiosemitic and North Omotic-speaking samples from the Horn. For specifics on this East Asia-associated element, see Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa.)
Genome analysis of modern Emirati individuals detecting the presence of a South/Central Asian component. Researchers have usually ascribed this element among contemporary Arabians to recent admixture with persons from South/Central Asia. However, the absence of the Ancestral South Indian component among Afro-Asiatic speakers in general, coupled with the presence of a South/Central Asian component among some Afro-Asiatic-speaking individuals, suggests deeper associations, perhaps linked to the dispersal of the original Elamo-Dravidian speakers (Elbait et al. (2021), Figure S2).
Ancestral composition of modern populations in South and Central Asia. The Onge or Ancestral South Indian genome component is today widely distributed across this region. However, it has not been observed among the Afro-Asiatic-speaking populations in Africa. This suggests that the South/Central Asian inferred element, which in some analyses has been detected among modern Cushitic, Berber and Arabic speakers, was either a) derived from an area in South/Central Asia, such as Balochistan, where the Ancestral South Indian component is not found at appreciable frequencies, or b) introduced early on by South/Central Asian settlers, such as the makers of the Indus Valley civilization, at a time period when the latter had not yet interbred with the original bearers of the Ancestral South Indian component (Lazaridis et al. (2016)). (*N.B. Distance analysis on the Vahaduo Admixture JS program indicates that, besides the recent IND_Great_Andamanese_100BP sample on Eurogenes’ official Global25 datasheet, the Onge share the nearest genetic affinity with the LAO_Hoabinhian specimen (see here). Ergo, the latter ancient individual is currently the best available stand-in for “pure” Ancestral South Indian ancestry.)
Final observations and recommendations
Another interesting facet of the Mota analysis concerns Llorente et al.’s assertion that they were able to detect West Eurasian affinities in every African population that they examined. While such ancestry is not unexpected for certain groups like the Maasai, Sandawe, and Khwe and Nama Khoi (who are known to have assimilated some early Cushitic pastoralists), it is less clear how isolated hunter-gatherer populations like the Xun/!Kung San and Mbuti Pygmies would have acquired such an influence. It is tempting to suggest that the spread of haplogroup E may have had something to do with this. However, the fact that the Neolithic LBK farmers apparently did not carry the paternal clade rules them out as the disseminating ancient population. So does the fact that the Stuttgart LBK woman possesses the derived SLC24A5 allele for lighter skin pigmentation, whereas most African populations, other than the Afro-Asiatic-speaking groups in the Horn and North Africa, do not.
Given the foregoing, better potential reference populations for future ancient DNA tests would be:
For related archaeogenetics, see Ancient DNA from Sudan. Also stay tuned for new aDNA work from Egypt.
*Update #1
In January 2016, Llorente et al. published an erratum online pertaining to their Mota study. The announcement (summarized here by Scientific American) indicates that there was a software-related oversight on the researchers’ end, which caused them to unknowingly overlook some chromosomal affinities that do apparently exist between the Mota specimen and their West Eurasian reference sample. The scientists also write that the proposed large ancient migration from West Eurasia was instead mainly confined to East Africa:
Erratum to Gallego Llorente et al. 2015
The results presented in the Report “Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent“ were affected by a bioinformatics error. A script necessary to convert the input produced by samtools v0.1.19 to be compatible with PLINK was not run when merging the ancient genome, Mota, with the contemporary populations SNP panel, leading to homozygote positions to the human reference genome being dropped as missing data (the analysis of admixture with Neanderthals and Denisovans was not affected). When those positions were included, 255,922 SNP out of 256,540 from the contemporary reference panel could be called in Mota. The conclusion of a large migration into East Africa from Western Eurasia, and more precisely from a source genetically close to the early Neolithic farmers, is not affected. However, the geographic extent of the genetic impact of this migration was overestimated: the Western Eurasian backflow mostly affected East Africa and only a few Sub-Saharan populations; the Yoruba and Mbuti do not show higher levels of Western Eurasian ancestry compared to Mota.
We thank Pontus Skoglund and David Reich for letting us know about this problem.
First off, the researchers are to be commended for their professionalism; both for having the courtesy to admit to a mistake (which happens) and then promptly trying to redress that, and for having the vision to conduct such an ancient DNA analysis in the first place.
That said, what are the implications of the Mota erratum? Not much since, as explained above, Mota was not an adequate African proxy to begin with. His uniparental lineages already pointed to gene flow from an early Afro-Asiatic-speaking settler group(s), as is the situation with his contemporary Ari relatives. This introgression is now also supported by Llorente et al.’s revised analysis, which has apparently detected minor West Eurasian ancestry in Mota’s genome.
A better and older African reference population than Mota would, therefore, perhaps be Schepartz (1987)’s hunter-gatherer sample from the Gogoshiis Qabe Rockshelter in southern Somalia (8,100-5,400 BP). Sellers (2008) compared these skeletons’ dental metric affinities to those of modern ethnic Somali pastoralists and Final Paleolithic Nubian hunter-gatherers, Sudanese agriculturists (3,400-1,200 BP) and Sudanese intensive agriculturists (1-1,500 BCE), and found that the ethnic Somali individuals were much more closely related to the more recent Sudanese intensive agriculturalists (viz. the Meroitic, X-Group and Christian period populations) and Sudanese agriculturalists (viz. the A-Group and C-Group populations) than to the Gogoshiis Qabe Rockshelter hunter-gatherers and Final Paleolithic Nubian hunter-gatherers. As explained on Ancient DNA from Sudan, this is because ethnic Somalis and other Afro-Asiatic-speaking populations that today inhabit the Horn did not actually evolve there. Their ancestors — ancient Cushitic speakers; not the Gogoshiis Qabe Rockshelter hunter-gatherers or Mota or the LBK makers — arrived instead from the Sahara and Nile Valley. The latter area is the likely Afro-Asiatic urheimat or “original homeland”.
For a comprehensive overview of these early “Hamitic” settlers in the Horn, see Punt: an ancient civilization rediscovered.
*Update #2
It is 2020, a good four years since our last update, and much has happened in the world of palaeogenetics. We now have genomic data from both the Iberomaurusians in Northwest Africa and early Southern Cushitic settlers in East Africa, as well as more extensive testing of ancient Egyptian specimens of the Nile Valley. These ancient DNA analyses, amplified by studies on modern groups, have gone a long way towards helping to clarify existing and past population relationships and inheritance patterns.
Anatolian Neolithic component & ancestral deconvolution
As discussed above, various SNP-based autosomal DNA analyses have suggested that the modern Afro-Asiatic speakers in the Horn of Africa (and to a lesser extent the Afro-Asiatic speakers in North Africa) harbor two distinct ancestries: one non-African and the other African.
Non-African ancestral composition of the Oromo. This NAF component is estimated to be comprised of 85% Anatolian Neolithic and 15% CHG for the Oromo, Amhara and Wolayta, and 92% Anatolian Neolithic and 8% CHG for the Somali (Molinaro et al. (2019), Supplementary Information).
Researchers have hypothesized that this non-African component is associated with either Neolithic Europe (as represented by the LBK culture), the Mesolithic Levant (Natufian culture), or the Neolithic Levant (Pre-Pottery Neolithic culture). According to Molinaro et al. (2019), when the whole genome of the Cushitic and Semitic-speaking individuals is analyzed (that is, when both the non-African and African components are examined together), their West Eurasian ancestry appears to be primarily derived from the Neolithic Levant. However, when the non-African component (NAF) is isolated through a process of ancestry deconvolution, it instead appears to be largely composed of the Anatolian Neolithic component, with minor admixture from the Caucasus Hunter-Gatherer (CHG) component. Among populations in Ethiopia, Molinaro et al. estimate this non-African component ratio at 85% Anatolian Neolithic and 15% CHG for the Amhara, Oromo and Wolayta, and 92% Anatolian Neolithic and 8% CHG for the Somali.
What are the merits of this biogenesis scenario?
Meshwesh, one of the ancient tribes of Sea Peoples. Note the braided hairstyle, which is still maintained by some Afro-Asiatic speakers in Northeast Africa (Salimbeti).
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 view that the early Cushitic settlers in the Horn of Africa descended from a population bearing Anatolian Neolithic ancestry (Schaefer et al. (2021)).
Lactase persistence allele frequencies of Afro-Asiatic-speaking populations in the Nile Valley and East Africa. The Northeast African-affiliated G-13907 variant peaks among the Beni Amer Beja (25%), the Arabian-affiliated G-13915 variant climaxes among Somalis (50%), and the South Cushitic-affiliated C-14010 variant is prevalent among the Yaaku (53.6%) (Tishkoff et al. (2010)).
It is supported by the lactase persistence (LP) alleles that are presently distributed in Northeast Africa. Tishkoff et al. (2010) and Hassan et al. (2016) note that the local Afro-Asiatic-speaking populations have high frequencies of various LP mutations, especially the pastoralist Cushitic groups. Of these derived alleles, the most commonly borne ones are: G-13907, which is centered in the Nile Valley and Horn of Africa and associated with cattle pastoralism; G-13915, which is centered in Arabia and associated with camel pastoralism; and C-14010, which is centered in the Great Lakes region and also associated with cattle pastoralism. The first two of these lactose tolerance variants are prevalent among North Cushitic and East Cushitic groups (G-13907 climaxes among Beni Amer Beja at 25%, and G-13915 peaks among Somalis at 50%), whereas C-14010 is typical of Southern Cushitic groups (the earliest occurence of this allele is among ancient Cushitic specimens of the Pastoral Neolithic; cf. Prendergast et al. (2018)). A fourth LP allele, T-13910, is most common in Europe and is also found among Tuareg Berbers, Mozabite Berbers and Fulani groups. According to Enattah et al. (2008):
The European T-13910 and the earlier identified East African G-13907 LP allele share the same ancestral background and most likely the same history, probably related to the same cattle domestication event.
Kulichová et al. (2017) indicate that the T-13910 LP allele was likely spread by pastoralists originating from outside Africa. Furthermore, Marcus et al. (2020) suggest that such population movement(s) from Europe into Africa also introduced both the R1b-V88 paternal haplogroup and Sardinian-related ancestry (Anatolian Neolithic). Given the shared ancestral background of the T-13910 and G-13907 lactase persistence variants, this implies that the Cushitic groups likewise inherited the G-13907 allele from ancestors bearing the Anatolian Neolithic component.
Haplotype network of lactase peristence (LP) alleles in Africa, Europe, the Middle East and Central Asia. The Northeast African G-13907 variant (located on haplotype 21) and the European T-13910 variant (located on haplotype 24) are associated with an A haplotype background (Liebert et al. (2017)). According to Enattah et al. (2008), the fact that these LP alleles share the same ancestral background denotes that they have “most likely the same history, probably related to the same cattle domestication event.”
Hodgson et al. argue cogently that genome-wide dating methods based on linkage disequilibrium are strongly biased in favour of recent admixture events.
Such recent admixture events did affect the current Ethiosemitic speakers, when they shifted from speaking their original Cushitic Agaw languages and adopted South Semitic languages. In the process, they acquired the paternal haplogroup J through intermarriage with Sabaeans (33% among the Amhara). They also have haplogroup A from admixture with South Omotic/hunter-gatherer populations: 17%-20% of Amhara, up to 36% of Tigrinya, and as high as 41% of Ethiopian Jews (cf. Wood et al. (2005), Appendix A; Scozzari et al. (2014), Table S7; Cruciani et al. (2002), Table 2; Gebremeskel (2018), Table 3.7).
Y-chromosome haplotype frequencies in select African populations Cruciani et al. (2002), Table 2). Most of the Ethiopian Jew individuals (41%) bear the haplotype I, which corresponds with the paternal haplogroup A. In Northeast Africa, this archaic African lineage is most common among Omotic and Nilo-Saharan-speaking populations. This suggests that the ancestors of the modern Ethiosemitic-speaking groups in Ethiopia significantly admixed with neighboring hunter-gatherers. Later periods of interbreeding with haplogroup J carriers from Arabia would dilute that elevated forager admixture, in the process raising the Ethiosemitic speakers’ overall West Eurasian ancestry to a level similar to that of their Cushitic-speaking neighbors (cf. López et al. (2021); their Cushitic-speaking Agaw sample has comparable West Eurasian ancestry as their Ethiosemitic-speaking samples despite the fact that the Agaw have sustained less recent gene flow from the Arabian peninsula).
Inferred ancestral components among the Afro-Asiatic-speaking groups of the Horn of Africa and other global populations. After correcting for linkage disequilibrium bias, most of the Cushitic and Ethiosemitic-speaking populations have non-African ancestry estimated in the 60%-70% range, similar to southern Moroccans. This non-African ancestry in the Horn largely belongs to the “Ethio-Somali” component, a West Eurasian ancestral element that peaks among ethnic Somalis (Hodgson et al. (2014), Table S6). (*N.B. The Sudanese sample above consists of Nilotes. It does not include Nubians or Sudanese “Arabs”, who instead share affinities with the Afro-Asiatic-speaking populations in Northeast Africa (see Dobon et al. (2015)).)
Pairwise FST estimates show the non-African affinities of the Ethio-Somali ancestral component. The Ethio-Somali component has the lowest pairwise FST values with the Maghrebi and Arabian ancestral populations (0.074, 0.083). FST values are on average lower between the Ethio-Somali component and non-African populations (mean=0.097) than with sub-Saharan African populations (mean=0.126) (Hodgson et al. (2014), Supplementary Text S1).
After correcting for this linkage disequilibrium bias, Hodgson et al. found that the non-African ancestry proportions for most of the Afro-Asiatic-speaking samples (Cushitic and Ethiosemitic speakers alike) are in the same 60%-70% range as southern Moroccans (Eaaswarkhanth et al. (2020) also observed a similar level of West Eurasian ancestry among many of the Kuwaiti Arab individuals they analyzed). Most of this estimated non-African ancestry in the Horn belongs to the “Ethio-Somali” component, a West Eurasian ancestral element that peaks among ethnic Somalis. The Ethio-Somali component correlates with ancient Cushitic ancestry, and is most closely related to the Maghrebi and Arabian ancestral components. Conversely, the African ancestry in the Horn primarily belongs to the “Ethiopic” component (also known as the “Omotic” component). This ancestral element correlates with indigenous hunter-gatherer or Mota-like ancestry, and peaks among the Ari Blacksmiths. The Ari today speak an Omotic language, which their forager progenitors are thought to have adopted from early Afro-Asiatic-speaking settlers in the Ethiopian highlands (Hodgson et al. (2014), Table S6). (*N.B. An earlier linkage disequilibrium-based study by Pickerell et al. (2014) posited that the West Eurasian ancestry in East Africa was only around 3000 years old and linked with the Semitic-speaking founders of the Kingdom of D’mt. Thanks to ancient DNA analysis, we now know that this biogenesis scenario is incorrect. Most of the West Eurasian ancestry in the region is actually older, higher in estimated frequency (especially when linkage disequilibrium bias is corrected for), and — as already strongly suggested by archaeological excavations — associated instead with the ancient Cushitic settlers of the Pastoral Neolithic (cf. Skoglund et al. (2017), Prendergast et al. (2018), Wang et al. (2020)).)
Afar:
69% Non-African ancestry (of which 43%=Ethio-Somali, 12%=Arabian, 14%=Others)
31% African ancestry (of which 8%=Ethiopic, 20%=Nilo-Saharan, 3%=Others)
Amhara:
65% Non-African ancestry (of which 34%=Ethio-Somali, 16%=Arabian, 15%=Others)
35% African ancestry (of which 16%=Ethiopic, 16%=Nilo-Saharan, 3%=Others)
Ari Blacksmith:
2% Non-African ancestry (of which 1%=Ethio-Somali, 1%=Arabian, 0%=Others)
97% African ancestry (of which 94%=Ethiopic, 1%=Nilo-Saharan, 2%=Others)
Ari Cultivator:
19% Non-African ancestry (of which 17%=Ethio-Somali, 1%=Arabian, 1%=Others)
81% African ancestry (of which 63%=Ethiopic, 10%=Nilo-Saharan, 8%=Others)
Ethiopian Somali:
64% Non-African ancestry (of which 53%=Ethio-Somali, 3%=Arabian, 8%=Others)
36% African ancestry (of which 8%=Ethiopic, 22%=Nilo-Saharan, 6%=Others)
Morocco S:
69% Non-African ancestry (of which 44%=Maghrebi, 8%=European, 17%=Others)
31% African ancestry (of which 22%=Niger-Congo, 4%=Nilo-Saharan, 5%=Others)
Oromo:
55% Non-African ancestry (of which 32%=Ethio-Somali, 12%=Arabian, 11%=Others)
45% African ancestry (of which 21%=Ethiopic, 19%=Nilo-Saharan, 5%=Others)
Somali:
66% Non-African ancestry (of which 57%=Ethio-Somali, 1%=Arabian, 8%=Others)
34% African ancestry (of which 6%=Ethiopic, 23%=Nilo-Saharan, 5%=Others)
Tygray:
68% Non-African ancestry (of which 35%=Ethio-Somali, 16%=Arabian, 17%=Others)
32% African ancestry (of which 12%=Ethiopic, 17%=Nilo-Saharan, 3%=Others)
Wolayta:
44% Non-African ancestry (of which 27%=Ethio-Somali, 7%=Arabian, 10%=Others)
56% African ancestry (of which 35%=Ethiopic, 15%=Nilo-Saharan, 6%=Others)
Although Molinaro et al. and Hodgson et al. differ on when that non-African ancestry arrived in Africa, these estimates nevertheless accord well with Molinaro et al.’s theory because the ancient Libyans (the immediate ancestors of the Maghrebans) were among the Sea Peoples.
Other examples:
The European explorer Bertram Thomas pictured with some “pure” Arabs. The deeply pigmented, nearly black color of these individuals approximates that of the earliest Semites. This is because, prior to contact with European-related peoples bearing Anatolian Neolithic ancestry and peoples from the Iranian plateau/Caucasus bearing Caucasus Hunter-Gatherer ancestry, the ancestral Semites (who descended from the Natufians) did not bear any alleles associated with lighter coloration.
(For further details, see Are Ashkenazi Jews and Sephardic Jews really of the same ancestral origin as Mizrahi Jews, Palestinians, Lebanese, peninsular Arabs, Assyrians, Mahra and other Semites? and Why do so many Ashkenazi Jews and Sephardic Jews have afros?.)
Y-DNA affinities of populations in the Middle East, Africa and Europe. Ashkenazi Jews cluster with Syrians, Palestinians, Lebanese and other Semitic-speaking groups due to their shared Levantine ancestral origins (Hammer et al. (2000)). (*N.B. Along with their mtDNA haplogroup frequencies, the Y-DNA haplogroup frequencies of Ashkenazi and Sephardic Jews can be used to infer how much Middle Eastern ancestry and European ancestry these individuals carry. Since around 80% of Ashkenazi and Sephardic Jews bear Middle Eastern-affiliated Y-DNA lineages compared to about 20% who bear European-affiliated Y-DNA lineages, and around 20% of Ashkenazi and Sephardic Jews bear Middle Eastern-affiliated mtDNA lineages compared to about 80% who bear European-affiliated mtDNA lineages, we may surmise that, on average, Ashkenazi and Sephardic Jewish individuals carry roughly 50% Middle Eastern-associated ancestry and 50% European-associated ancestry. This corroborates genome analyses on Ashkenazi and Sephardic Jews, which have similarly found that these individuals are of Middle Eastern origin but harbor substantial European admixture (e.g. Atzmon et al. (2010)). For further details on Stefflova et al. (2009)‘s mAUTO formula, which we have used here to calculate the autosomal DNA of Ashkenazi and Sephardic Jews, see Can the Y-DNA and mtDNA of a population be used to infer its autosomal DNA?.)
In terms of autosomal DNA, Ashkenazi Jews and Sephardic Jews cluster in between other Middle Easterners and Europeans. This is a reflection of the fact that their Semitic male ancestors interbred with native women in southern Europe.
Principal Component Analysis of Askhenazi Jews and contemporary and medieval samples from the Middle East and Europe. Individuals buried at the medieval Jewish cemetery at Erfurt, Germany, form two distinct genetic clusters: one group pulls toward modern Lebanese and the other group pulls toward modern north Italians and Greeks. Ashkenazi Jews, like some Erfurt individuals, are positioned in between these two clusters. This again reflects the fact that the Semitic ancestors (mostly males) of Ashkenazi Jews and Sephardic Jews interbred with natives in southern Europe (mostly females) when they left their Levant homeland — an admixture process that was apparently still in its early stages as recently as the Middle Ages (Waldman et al. (2022)). This is also supported by linguistic analysis, which indicates that the Yiddish tongue that many Ashkenazi Jews now speak is a creole of their Levantine ancestors’ original Semitic Hebrew language and the German language of their European host community (much like how Jamaican patois is a creole of the native Niger-Congo languages of the West African ancestors of Jamaicans and the English language of their European host community). The fact that the Indo-European contact language here is a Germanic idiom spoken in west-central Europe rather than Italian or Greek conveys that the Levantine forebears of Ashkenazi Jews would still have spoken their native Hebrew language as they spread from southern Europe to other parts of the continent. (For more information, see Yes, Ashkenazi Jews (Including Gal Gadot) Are People of Color.)
STRUCTURE and phylogenetic analysis of Jewish populations. Ashkenazi Jews cluster with Mizrahi Jews from Syria and other Jewish groups. These Jewish populations carry both Levantine/Arabian (pink component) and European admixture elements (blue component). Ashkenazi individuals trace just over half of their ancestry to the Levantine/Arabian component (~60%), intermediate between Palestinians and southern European populations. This is consistent with the observation that the Semitic male ancestors of Ashkenazi and Sephardic Jews interbred with native women in southern Europe (Atzmon et al. (2010)). (*N.B. For more details, see Are Ashkenazi Jews and Sephardic Jews really of the same ancestral origin as Mizrahi Jews, Palestinians, Lebanese, peninsular Arabs, Assyrians, Mahra and other Semites?)
Structural genome analysis indicates that individuals belonging to the Pre-Pottery Neolithic culture of the Levant (Levant Neolithic) interbred significantly with incoming Southern European-related groups, who bore the Anatolian Neolithic ancestral component (Schuenemann et al. (2017)). Consequently, Levantine samples from this Neolithic period acquired substantial “Mediterranean” ancestry that is not pronounced among their Mesolithic Natufian ancestors. Bronze Age migrations of peoples from the Caucasus/Iranian plateau would subsequently introduce the paternal haplogroup J and the Caucasus Hunter-Gatherer/Iran Neolithic component to the ancestors of the Semitic-speaking natives of the Levant, Arabian peninsula and Mesopotamia. This would have the effect of diluting the Semites’ original Natufian ancestry as well as their Anatolian Neolithic admixture, thereby bringing them genetically closer than before to populations in the Caucasus and Iranian plateau. Modern exceptions are groups like the Mehri of southern Arabia, Yemenis of the Marib governorate, and Bedouin Arabs of the Negev Desert, who, due to their geographical isolation, have managed to retain more of the Semites’ original Natufian ancestry.
Principal Component Analysis of modern and ancient individuals. The contemporary Semitic speakers cluster with the Bronze Age inhabitants of the Levant (represented by the autochthones of Sidon; they bore the Y-DNA haplogroup E, as well as the J clade from admixture with newcomers bearing Caucasus Hunter-Gatherer/Iran Neolithic ancestry), followed by the Neolithic Levantines (Pre-Pottery Neolithic makers; they bore the Y-DNA haplogroup E and the older haplogroup CT, as well as the T and H clades from admixture with new settlers carrying Anatolian Neolithic ancestry) and the Mesolithic Levantines (Natufians; they bore the Y-DNA haplogroup E, as well as the older haplogroup CT). Ergo, despite experiencing various foreign admixtures, the modern Semites have remained genetically close to their ancient Levantine forebears (Lazaridis et al. (2016).
Genome analysis of Semitic-speaking Yemeni individuals. The Mahra/Mehri samples generally have the most Arabian ancestry (teal); four of the examined Mahra individuals almost entirely belong to that Natufian-related ancestral component. This supports the long-held belief that the Mahra represent the Semites in their “purest” form. Conversely, Yemenis from the other coastal governates of Hadramout and Shabwah have the most foreign admixture. The latter primarily consists of Sub-Saharan African elements derived from Niger-Congo-speaking West African (light green) and Nilo-Saharan-speaking East African (dark green) sources, as well as Caucasus-related admixture (peach). Additionally, the Hadramis have the most European-related admixture (lilac). This is in agreement with the theory that the ancient Himyarites of Hadramout and the adjacent southwestern provinces bore substantial Mediterranean or Sardinian-like ancestry, whereas the ancient Sabaeans in the northern territories had more pronounced Caucasus influences (see Punt: an ancient civilization rediscovered). Correspondingly, modern Yemenis from the highlands and northwest have the most Caucasus-related admixture, except for individuals in the Marib and Al Jawb governates, who have retained more of the Semites’ original Natufian ancestry (Vyas (2017).
Reconstruction of a 7500 year old Early European Farmer woman excavated in Gibraltar. Ancient DNA analysis of the specimen (named “Calpeia”) has found that she was defined by the Anatolian Neolithic ancestral component. Craniometric analysis further indicates that she and other bearers of the Anatolian Neolithic component would have had a similar “Mediterranean Caucasoid” or Sardinian-like physiognomy. However, their skin tone would likely have approached a light brown since they only possessed the derived SLC24A5 allele, the primary contributor to lighter skin pigmentation. Besides this mutation, populations in Europe now have several additional, minor light skin-associated variants, which cumulatively are responsible for the white complexion of modern Europeans (Gibraltar National Museum).
Analysis of ancient DNA samples relative to modern European, Caucasus and Near Eastern populations indicates that the Anatolian Neolithic component (green) is today most frequent among Sardinians, Basques and other southern & western European groups. The Western Hunter-Gatherer/WHG component (blue) is most common among Russians and other eastern European groups. The Caucasus Hunter-Gatherer/CHG component (yellow) peaks among Georgians and other populations inhabiting the Caucasus. The Natufian component (red) climaxes among Saudis, Yemeni Jews, Palestinians and other Semitic peoples. Ashkenazi and Sephardic Jews derive much of their ancestry from the Near East and Caucasus, with a significant admixture from southern Europe (Sarno et al. (2017), Supplementary Fig. S3). Haplogroup analysis indicates that this is because these Jewish groups’ Semitic male ancestors intermarried with native women in southern Europe when they first arrived from their Levant homeland (Nebel et al. (2000), Costa et al. (2013)).
Genome analysis of Daza Toubou individuals using the Vahaduo Admixture JS program indicates that they are most similar to the Kababish “Arabs” of Sudan. They share the same assortment of predominant Eurasian ancestries, which consist of majority West Eurasian elements (ancient Egyptian, European Steppe, and Levantine Natufian components) and a minority East Eurasian element (East Asian component). However, the Daza have a greater Sub-Saharan admixture than the Kababish (45.4% vs. 34.9%), as well as a more elevated Iberomaurusian admixture (14.5% vs. 0.8%).
Genetic distance analysis of modern Daza Toubou, Baggara “Arabs” and Kababish “Arabs” using the Vahaduo Admixture JS program. Almost all of these individuals show a clear preference for the medieval Kulubnarti Nubian specimens from Sudan. This confirms that the Toubou, Baggara “Arabs” and Kababish Sudanese “Arabs” share a common origin, tracing most of their proximal or recent ancestry to earlier Cushitic-related peoples.
Genome analysis of Shuwa “Arab” individuals and other modern populations from Africa. The examined Shuwa “Arabs” (labeled here “Cameroon Arabe”) derive most of their Eurasian ancestry from a Cushitic-related source (pink component), followed by a Tunisian Maghrebi-related source (light blue component). This same composite autosomal DNA signature appears to also be present at lower frequencies among the Sahelian Kanuri and Kotoko populations (who are thought to have expanded westward from the Sudan), as well as among Chadic-speaking populations of northern Cameroon (who mostly bear the Eurasian R1b-V88 paternal haplogroup). Altogether, this suggests that the Shuwa “Arabs” may originally have been Chadic speakers, who inhabited the eastern Sahara and Nile Valley beside Nilo-Saharan and Kordofanian speakers. There, through population contact, they seem to have picked up the Nilo-Saharan purple component. The Shuwa appear to have later spread to the Cameroon/Nigeria vicinity, where they settled amongst and further intermixed with Niger-Congo speakers. Such a population movement is also consistent with Basil Davidson (1968)‘s proposition that the iron-making tradition of the Nok culture in western Africa was established “perhaps by diffusion from the metal-traders of Meroe on the Middle Nile or the North African Berber states” (Bird et al. (2023)).
MSMC analysis inferring the prehistoric population size of the Toubou, Afro-Asiatic-speaking groups, and other global samples. According to the scientists, “Eurasian populations had a distinctive bottleneck at the time of their exodus from Africa ~60,000 ya. Compared to other Africans, admixed Africans (from a Eurasian gene flow), such as Egyptians, Ethiopians, and the Toubou, also showed a decline in population size during the same period” (Haber et al. (2016)).
In further support of this connection between Afro-Asiatic speakers, the ancient Sahara, and the Anatolian Neolithic component, Haber et al. (2016) indicate that the Toubou inhabitants of the Sahel carry haplogroup T at appreciable frequencies (31%). They also report that the oldest instances of the T paternal lineage have been observed among Early European Farmers from the Neolithic Linearbandkeramik (LBK) culture. Moreover, the researchers note that their sampled Toubou individuals, although admixed with neighboring Nilo-Saharan-speaking communities, are genomically more similar to the Afro-Asiatic-speaking populations in the Horn region than to local Nilo-Saharan speakers. Additionally, Haber et al. conducted f3 statistical analysis with the aim of determining the most plausible source of the Eurasian ancestry shared by these populations. They found that, among modern populations, the Afro-Asiatic speakers of the Horn and the Toubou showed greatest genetic affinity with Sardinians, and, among ancient populations, they showed greatest genetic affinity with the early Neolithic LBK farmers of Europe — both of which are characterized by the Anatolian Neolithic ancestral component. Vicente et al. (2019) also observed similar genome ties between their Toubou sample (Daza/Gorane or southern Toubou) and their Nubian and Afro-Asiatic-speaking samples from Northeast Africa. Furthermore, Haber et al. conducted an MSMC analysis to infer the prehistoric population size of their Toubou and Afro-Asiatic-speaking samples. They concluded that, like Eurasian populations that are thought to have undergone a bottleneck during the putative Out-of-Africa exodus (ca. 60,000 ybp), “compared to other Africans, admixed Africans (from a Eurasian gene flow), such as Egyptians, Ethiopians, and the Toubou, also showed a decline in population size during the same period.” They also found that “PCA of worldwide populations shows that Near Easterners and East Africans are intermediate to Eurasians and sub-Saharan Africans on PC1”, and that the Toubou individuals clustered toward the Ethiopian samples. In short, this suggests that both the Toubou and Nubians (specifically, descendants of the “red” Nubians), who now speak Nilo-Saharan languages, originally spoke languages belonging to the Afro-Asiatic family. The above securely places the earliest arrival of Minoan/LBK/Sardinian-related ancestry in Africa to the Neolithic period, thus giving a wider berth to Molinaro et al.’s thesis.
Admixture analysis of Yemeni individuals. At K=3, the Yemeni samples from the Marib governorate have the most indigenous Near Eastern (Natufian) ancestry of all the examined groups, with little-to-no extraneous influences (Haber et al. (2019)).
It is in agreement with the skin pigmentation alleles that are presently found in the Horn. Researchers have observed that around 60% of the modern Afro-Asiatic speakers in the region carry the derived variant of the SLC24A5 gene, which is associated with lighter skin pigmentation (cf. ALFRED; et al. (2015)). This mutation has been detected at fixation rates (100%) among ancient Cushitic pastoralists in East Africa (Wang et al. (2020)), as well as among Minoan specimens from Crete and Kelif el-Boroud individuals. However, Natufian specimens were found to bear the ancestral SLC24A5 allele linked with darker pigmentation. The Natufians, therefore, were not the ancient population from whom the early Cushitic settlers inherited the derived SLC24A5 allele. Likewise, the Natufians do not appear to have been directly ancestral to the predynastic Egyptians. Coon (1939) indicates that many exhumed Badarian specimens were well-preserved, and their “skin was apparently brunet white, while the hair was black or dark brown in color, thick, of fine texture, and usually wavy in form.” Could perhaps, then, the derived SLC24A5 mutation have been introduced to the Cushites and Egyptians by autochthonous hunter-gatherers? It certainly was not by Ari/South Omotic peoples because the Mota specimen did not have any light skin-associated alleles. Moreover, we know from archaeogenetic analysis that during the Iron Age, Khoisan-related foragers were present as far north as coastal Kenya, where they were in contact with some Cushitic “Azanian” settlers (see Who were the ancient Azanians?). However, here too we can dismiss the Khoisan as a source of the derived rs1426654 variant at the SLC24A5 gene because a) as Lin et al. (2018) note, this “most common derived haplotype is identical among European, eastern African, and KhoeSan individuals”, b) the oldest Khoisan-related specimens analyzed (c. 4000 years old) did not carry that polymorphism (Schlebusch et al. (2017) indicate: “the SLC24A5 G allele is near fixation in African populations and all individuals with enough data exhibit the alternative G variant for the SLC24A5 gene SNP rs1426654, which codes for darker skin color”; cf. Supplementary Materials), and c) Lin et al. (2018) found evidence that the mutation was actually introduced quite recently into the Khoisan gene pool, around 2000 years ago, and likely by migrating ancient Cushitic pastoralists. Thus, it is most probable that an Anatolian Neolithic-related population (e.g. Sea Peoples, Minoan, Linearbandkeramik) and/or a Neolithic Levant-related population (Pre-Pottery Neolithic) ancestral to the early Cushites is responsible for the presence of the derived SLC24A5 variant in the Horn.
South Arabian men of the Janaba and Yal Wahiba tribes. Besides the fact that the Levantine Natufians did not carry any mutations associated with lighter coloration, the idea that these ancient individuals were deeply pigmented is also suggested by the dark skin complexion of the modern Semitic peoples who carry the most Natufian ancestry. Among these “pure” Semites are the Janaba and Yal Wahiba, who have many members that are quite literally black-skinned.
Reconstruction of a Mesolithic Natufian man, based on the Jericho Skull excavated in Israel (British Museum). Ancient DNA analysis of Natufian specimens has revealed that they were ancestral to the Levantine Neolithic population, forebears of the contemporary Semitic-speaking groups of the Middle East (Lazaridis et al. (2016)).
Craniometric analysis of ancient pastoralists excavated at Wadi Takarkori Acacus, southwestern Libya. The Takarkori sample is most similar to Early Holocene (Kiffian culture) and Mid-Holocene (Tenerian culture) specimens from the Gobero site in the Sahara. Although chronologically younger than the Takarkori individuals, the Fezzan specimens (Fewet, Wadi-el-Ajal, Tahala) have retained a more archaic morphology and cluster instead with the late Middle Pleistocene Herto skull (Homo sapiens idaltu) (Vincenzo et al. (2015)).
Correspondingly, ancient DNA analysis of early pastoralists from Takarkori, Libya, which date from this “Green Sahara” period, were found to belong to the N maternal haplogroup (Vai et al. (2019)). Vincenzo et al. (2015) conducted cranial analysis on the Takarkori skeletal remains and observed that these individuals were most similar to Early Holocene (Kiffian culture) and Mid-Holocene (Tenerian culture) specimens from the Gobero site in the Sahara. Earlier morphological analyses report that these Gobero specimens had Iberomaurusian & Capsian and Mediterranean affinities, respectively (cf. Sereno et al. (2008); Wilford (2008)). Vincenzo et al. also found that the Takarkori individuals were physically distinct from the Fewet, Wadi-el-Ajal and Tahala specimens from the Fezzan. Although much more recent than the Takarkori sample, the Fezzan cohort had a more “archaic” morphology and clustered instead with the late Middle Pleistocene Herto skull (Homo sapiens idaltu):
The Neighbour Joining tree shows that the small sample from Takarkori takes relationships with the populations from Gobero in Niger, either coeval (Gobero B) or more ancient (Gobero A), thus with humans from sub-Saharan regions which are characterized by a wide morphological variation. Populations from the Fezzan such as Fewet, Wadi-el-Ajal and Tahala, much younger chronologically than the Takarkori sample, are separated from it (according to the length of the branches that are proportional to the Euclidean distances between the metrical variables) and are, by contrast, more in relationship with one of the earliest representatives of our species such as the cranium from Herto (Ethiopia, late Middle Pleistocene). Paradoxically, therefore, although more recent than the two women from Takarkori, samples from this time period appear more “archaic” and closer to the root of the tree.
Moreover, Haak et al. (2005) discovered that the N1a sublineage (which is common today among the Afro-Asiatic-speaking populations in Northeast Africa) was prevalent among the Early European Farmers of Neolithic Europe. This is consistent with Marcus et al.’s proposal, and also coheres well with Molinaro et al.’s suggested later in-migration of Sea Peoples.
Craniometric analysis of various global populations. The Ethiopia sample clusters in between the Paleo Europe and modern Egypt samples (Froment (1998)).
That said, it is almost certain that there were also older population movements from and/or to the Levant. Iberomaurusian samples and early Neolithic specimens from the Ifri n’Amr or Moussa site in Morocco were found to belong to a Near East-affiliated ancestral component, which they shared with the Mesolithic Natufians and the Pre-Pottery Neolithic culture bearers of the Levant. Furthermore, these ancient specimens had no apparent Sub-Saharan affinities: Loosdrecht et. al (2018) initially posited that Taforalt had some Sub-Saharan admixture related to modern populations in West Africa. However, Lazaridis et al. (2018) subsequently found that “West Africans (represented by Yoruba) had 12.5±1.1% ancestry from a Taforalt-related group rather than Taforalt having ancestry from an unknown Sub-Saharan African source.” Lazaridis et al. (2016) also assert that “no affinity of Natufians to sub-Saharan Africans is evident in our genome-wide analysis, as present-day sub-Saharan Africans do not share more alleles with Natufians than with other ancient Eurasians.”
Genome analysis identifying a substantial North African Iberomaurusian component (orange element) in the ancestral composition of ancient Levantine individuals from the Pre-Pottery Neolithic culture of c. 8000-6000 BCE (D’Atanasio et al. (2023)).
Craniometrically, Terrazas Mata and Benavente (2013) report that no firm conclusions can be drawn on the population affinities of the Iberomaurusians. Their Taforalt sample almost exclusively grouped with other ancient individuals dating from the Middle Pleistocene, Upper Paleolithic and Early Holocene, and therefore in their judgement “these populations are still too far away from the Neolithic and Historic samples, so they could not represent direct ancestors of any present human population.” However, their modern Horn of Africa and Dynastic Egypt samples clustered together. The scientists indicate that this is because “these groups seem to represent migrations of groups from the Middle East, who spoke languages of the Afro-Asiatic family”. (*N.B. Terrazas Mata and Benavente state that “only two recent populations, both from Western Africa, have been integrated into the third branch, being near to the Paleolithic sample from Taforalt, Algeria”. However, the scientists do not explain the reason behind this apparent affinity. It may be that Terrazas Mata and Benavente’s Taforalt sample contains some admixed individuals since Brace et al. (2006) observed that their Taforalt/Afalou cohort clustered with Upper Paleolithic European crania rather than with any of their Sub-Saharan samples. Another possibility is that this is a manifestation of gene flow in the other direction, from the Iberomaurusians into the Proto-Niger-Congo population. As suggested above (under Final observations and recommendations), the Iberomaurusians may have introduced the paternal haplogroup E to adjacent Sub-Saharan African groups, a clade of which they and the related Natufians are now known to be the oldest carriers. This admixture event(s) probably also transmitted some West Eurasian genes, which Lazaridis and other researchers have detected in their Yoruba and other modern West African samples. Alternatively, a population(s) with considerable Iberomaurusian ancestry (e.g. the ancestral Fulani) may have been responsible for spreading the E lineage to the Niger-Congo-speaking peoples. The Niger-Congo speakers may instead have originally borne the archaic A and B haplogroups given: 1) the young TMRCA of the E1b1a or E-M2 subclade, which most Niger-Congo-speaking individuals today carry (estimated at just 6175-6588 years before present; cf. Ansari Pour et al. (2013)), 2) the genomic ties between Niger-Congo speakers and the primary carriers of the A and B lineages (viz. Khoisan, Pygmies, Nilo-Saharan speakers, Hadza and other East African hunter-gatherers), 3) the disconnect between the autosomal DNA of the Niger-Congo-speaking populations and those of the other haplogroup CT-descended global populations, 4) the fact that, with the exception of the Mota sample (which has trace levels of Eurasian admixture — see Erratum above), all of the pre-Bantu Migration ancient specimens in Sub-Saharan Africa that have thus far been genetically analysed belong to the archaic A and B paternal haplogroups (Lipson et al. (2022), Extended Data Table 1), and 5) the earliest and exclusive presence of the CT and DE parent clades among ancient specimens in Asia.)
Ultimately, the Iberomaurusians appear to have originated from outside Africa; most probably somewhere near Europe given their craniometric and mtDNA ties with Upper Paleolithic Europeans. Accordingly, the oldest instance of U6*, a signature maternal haplogroup borne by Iberomaurusian specimens, has been observed in Peştera Muierii 1 (PM1), a 35,000 year old individual excavated in Romania (Hervella et al. (2016)):
After the dispersal of modern humans (Homo sapiens) Out of Africa, hominins with a similar morphology to that of present-day humans initiated the gradual demographic expansion into Eurasia. The mitogenome (33-fold coverage) of the Peştera Muierii 1 individual (PM1) from Romania (35 ky cal BP) we present in this article corresponds fully to Homo sapiens, whilst exhibiting a mosaic of morphological features related to both modern humans and Neandertals. We have identified the PM1 mitogenome as a basal haplogroup U6*, not previously found in any ancient or present-day humans. The derived U6 haplotypes are predominantly found in present-day North-Western African populations. Concomitantly, those found in Europe have been attributed to recent gene-flow from North Africa. The presence of the basal haplogroup U6* in South East Europe (Romania) at 35 ky BP confirms a Eurasian origin of the U6 mitochondrial lineage. Consequently, we propose that the PM1 lineage is an offshoot to South East Europe that can be traced to the Early Upper Paleolithic back migration from Western Asia to North Africa, during which the U6 lineage diversified, until the emergence of the present-day U6 African lineages.
Furthermore, this part of Europe, the Balkans, remains a hub of V13, the most prevalent Europe-centered derivative of the E1b1b paternal haplogroup. The Iberomaurusians carry E1b1b’s M78 sublineage, which is the parent clade of V13. There are, therefore, also Y-DNA ties. In short, since the ancient West African-related population that the Iberomaurusians are hypothesized to have had contact with was indigenous to Africa, it follows that the Iberomaurusians must have lacked any such admixture when they first settled there. Future testing of older Iberomaurusian specimens is thus likely to yield examples of relatively unadmixed individuals. (For further details, see Did the Capsians of North Africa descend from the Iberomaurusians?, What are the genetic affinities of the Aterians?, and Have the Kiffians and Tenerians of the Gobero been genetically analysed yet?.)
Moreover, the Iberomaurusians of Taforalt, the Ifri n’Amr or Moussa specimens, the Pre-Pottery Neolithic makers and the Natufians all bore the E1b1b paternal clade. This makes them, along with a 12,000-8,000 BCE Mesolithic specimen from the Belt Cave in Iran (Narasimhan et al. (2019)), the oldest known carriers of the haplogroup E. Some Natufian individuals were also found to carry the Y-DNA haplogroup CT, which is ancestral to the DE (parent clade of haplogroups D and E) and CF (parent clade of haplogroups C and F) lineages. Additionally, multiple other instances of both CT and DE have been observed among ancient specimens in Asia, whereas old examples of these lineages have yet to be observed anywhere in Africa (cf. Narasimhan et al. (2019), Table S1-S5). Since these are the earliest and only instances of these clades in the archaeogenetic record, this strongly bolsters the argument that Asia rather than Africa is the true point of origin and dispersal of the CT and DE haplogroups. These early peoples likewise all harbored maternal lineages derived from the M and N mtDNA macroclades. Altogether, this suggests that, as in the case of the paternal haplogroup T, the contemporary Afro-Asiatic speakers in the Horn of Africa and North Africa inherited their E1b1b sublineages from ancestors with a Eurasian genetic background (Fregel et al. (2018); Loosdrecht et al. (2018); Lazaridis et al. (2016)).
African admixture component
When the Mota man was excavated in Ethiopia, some scholars wondered if the population to which he belonged might represent the native hunter-gatherers whom the Afro-Asiatic-speaking agropastoralists encountered and possibly mixed with when the latter first settled in the Horn. More extensive DNA analysis has, however, shown that gene flow between the two communities, although present, was minimal (Lipson et al. (2020) places Mota-related admixture in the Cushitic-speaking Agaw at 8%; no other hunter-gatherer-related admixture has been detected in the Horn). Due to this apparent lack of a suitable ancient proxy sample, researchers have formulated alternative admixture scenarios using various modern populations. In qpAdm and other genomic programs, these contemporary reference groups have served as stand-ins for the elusive ancient African contact population(s). The Dinka, Nuer and other northern Nilotic communities have often been the go-to populations utilized as the African proxies. The problem is that all of these groups have proven to be even less reliable surrogates than Mota.
Let us examine why that is:
By contrast, in various autosomal DNA studies, the northern Nilotes have appeared to be almost completely of African ancestry. For example, in the African Genome Variation Project’s analysis, the Dinka sample showed no extraneous influence at K=2 (cf. Gurdasani et al. (2015)). This apparent lack of non-African affinities is why the Dinka, Nuer and other northern Nilotes have often been used by researchers as proxies for the African component. In 2017, Skoglund et al. compared the same AGVP Dinka sample to that of an ancient South Cushitic pastoralist (Luxmanda), the first such specimen to be genetically analyzed. In their admixture analysis, the Dinka individuals now all of a sudden showed almost 30% non-African ancestry at the K=2 level. Judging by the existing uniparental marker data, it’s pretty clear why that is: there was non-African ancestry buried within the northern Nilote gene pool, and that ancestry was specifically derived from earlier Cushitic peoples such as Luxmanda.This, in a nutshell, explains why the Dinka, Nuer and other northern Nilotes often fit well as proxies for the African component in admixture testing models. However, the problem with using such admixed groups in these analyses is that doing so leads to highly distorted estimates of ancestry proportions. For example, let’s say a researcher is testing admixture models on qpAdm and finds that his modern Cushitic or Ethiosemitic-speaking population is best modeled as 50% Neolithic Levantine + 50% Dinka. What that program is really telling him is that his sample is best modeled as 50% Neolithic Levantine + X% ancient Nilotic + X% ancient Cushitic. This again stems from the fact that the Dinka are not a purely African population, but rather a Nilote-descended community with significant Cushitic admixture (whence was derived their non-African ancestry). When factoring in the ~30% of non-African ancestry that Skoglund et al. (2017) detected in their Dinka reference sample, the estimated whole genome ancestry apportionment then actually becomes 50% Neolithic Levantine + 35% African + 15% unclassified Eurasian. This necessary adjustment therefore brings the non-African total to around 65%. The same corrective adjustment would have to be made if the African reference sample were a southern Nilotic group, such as the Maasai Nilotes (e.g. when the ~30% of West Eurasian admixture in Ali et al. (2020)‘s East African proxy sample (Maasai) is taken into account, the estimated non-African ancestry for their northeastern Somali sample rises to 70%; this more accurate total is close to Hodgson et al.’s 66% average for their general ethnic Somali sample). That’s also before correcting for linkage disequilibrium bias, which, as Hodgson et al. (2014) observed and Choudhury et al. (2020) also demonstrated, would increase the Cushitic speakers’ Eurasian total even further. Put simply, there is considerable extra non-African ancestry in the genome of the Afro-Asiatic-speaking sample which is not being counted. This results in an inaccurate overall estimation of ancestry proportions. It also conflicts with scientific data gathered through other means (viz. craniometric analysis, anthropometric analysis, linguistics).
MtDNA (maternal) haplogroup affinities of various populations in Africa and Asia. The Afro-Asiatic-speaking Somali, Oromo, Afar, Tigrai and Gurage samples cluster with Yemeni individuals of Hadramout, as well as the Dawoodi Bohra and other Muslim groups of South Asia that originated from Arabia. The Dinka, Nuer and Datog Nilote samples cluster instead with other Nilotic, Bantu and hunter-gatherer populations of East Africa (Non (2010)). (*N.B. Mohamoud (2006) observed similar affinities in his analysis of HLA antigens, noting that “the result of HLA class I and class II antigen frequencies show that the Somali population appear more similar to Arab or Caucasoid than to African populations”.)
The maternal haplogroups distributed today among the northern Cushitic and Ethiosemitic-speaking populations in the Horn of Africa do not support the claim that there was significant admixture between their Afro-Asiatic-speaking male ancestors and Nilotic females. This idea first gained prominence after the publication of Watson et al. (1996) and Watson et al. (1997), two early mtDNA studies which reported that the analyzed Somali (sampled in Kenya) and Tuareg Berber (sampled in Niger) individuals primarily belonged to the L3 mtDNA haplogroup. Comas et al. (1999) similarly indicated that only 5%-27% of Ethiopians bore non-African mtDNA lineages. However, it turns out that these results were inaccurate since the L3 macroclade’s phylogeny had not yet been fully resolved. Specifically, many of the samples that were at the time presumed to represent L3 carriers actually belonged to derivatives of the N haplogroup. Martin Richards, who served as a co-author on Watson et al. (1997), explains the situation thusly:
The analysis of Watson et al. (1997) was based largely on control-region sequence data, which fails to resolve many mtDNA haplogroups. By targeting newly identified coding-region variants, Quintana-Murci et al. (1999) distinguished two major clades in non-Africans, within haplogroup L3. One of these had already been identified as the Asian super-haplogroup M (Torroni et al. 1994a); Quintana-Murci et al. (1999) showed that all other non-African L3 lineages fell into a second major clade, later named haplogroup N[…] haplogroup M was present at high frequency in Ethiopia and Somalia
Non (2010) later retested Watson et al.’s Somali sample, in addition to Oromo, Afar, Amhara and Tigray samples from Ethiopia. She found, given the now better understood mtDNA phylogeny, that around 60% of the examined individuals actually bore the Eurasian M and N haplogroups. Ottoni et al. (2009) likewise reanalysed Tuareg individuals and noted that the most common mitochondrial haplogroup among them was, in fact, the H1 lineage, which is associated with Europe. Subsequent mtDNA studies by Kivisild et al. (2004), Holden (2005) and Mikkelsen et al. (2012) observed comparable haplogroup frequencies for these Afro-Asiatic-speaking populations, including a very low occurrence of the L5 clade (<1% on average). This is of particular importance since the L5a subclade is the most common maternal lineage among the Dinka Nilotes in the Sudan area; 31% of Dinka individuals there belong to this subhaplogroup (cf. Hassan et al. (2009)). Thus, instead of showing appreciable Nilotic affinities, the mtDNA profiles of the northern Cushitic and Ethiosemitic-speaking populations in the Horn region parallel those of certain other Afro-Asiatic-speaking communities in the Arabian peninsula and North Africa (e.g. Non (2010) observed that her Somali sample clustered with a Yemeni sample from Hadramout, Kujanova ́ et al. (2009) note that 68.6% of Egyptians from El-Hayez oasis carry M and N lineages, Bekada et al. (2015) report a similar frequency of ~63% M and N haplogroups among their Algerian cohort from Algiers (see S5 Table), Coudray et al. (2009) likewise reveal that 55.3% of Figuig Berbers from northern Morocco are M and N carriers, and Kivisild et al. (2004) indicate that 56.4% of their Yemeni sample bears M and N derivatives.) These findings are why Shriner et al. (2016) propose that African introgression may have primarily occurred through the paternal line via the E1b1b haplogroup rather than through the “Arabian” maternal line. However, this argument, too, is no longer tenable because ancient DNA analysis has demonstrated that the haplogroup E was first affiliated with Epipaleolithic & Neolithic North African and Middle Eastern groups rather than Nilo-Saharan-speaking populations. The early Nilo-Saharans instead appear to have originally belonged to the Y-DNA haplogroup A (see Ancient DNA from Sudan).
Population tree estimating Fst genetic distance between ethnic groups in Africa. The Cushitic Beja samples (Beni Amer & Hadendoa) cluster with the Mozabite Berber sample. Despite some recent Arabian and Sub-Saharan contacts, this suggests that the Beja have retained much of their Cushitic ancestry (as also observed by Dobon et al. (2015) and Gopalan et al. (2019)). These continued ties ultimately reflect the northern origin of the early Cushitic settlers of East Africa (Scheinfeldt et al. (2019)).
Having identified a Cushitic-like substratum in Modern South Arabian, Militarev (1984, 18-19; cf. also Belova 2003) proposes that Cushites originally lived throughout the Arabian Peninsula; thus they would be the original southern neighbors of the Semites, who then assimilated those Cushites who did not move into Ethiopia. This hypothesis is supported by Anati (1968, 180-84), who analyzed the rock art of Central Arabia. He connected the pictures of the ‘oval-headed’ people depicted with shields with the Arabian ‘Cushites’ from the Old Testament [Genesis 10.6-12; Isaiah 45.14] described also with specific shields [Jeremiah 46.9; Ezekiel 38.5]. The spread of Cushites in Africa is connected with the Rift Valley. In the coastal area of Eritrea and Djibuti, where the Rift enters into the African mainland, three archaic representatives of the North, Central (= Agaw) and Eastern branches of Cushitic appear: Beja, Bilin and Afar-Saho respectively. In this place the disintegration of Cushitic probably began.
Vyas (2017) also genetically analyzed Mehri (Mahra) individuals from Yemen’s Mahra governorate, who natively speak Mahra, a Modern South Arabian language. He found that, in contrast to other Arabian and Levantine populations, his Mehri sample experienced minimal exogenous admixture:
While admixture and gene flow generally characterize the Levantine and southern Arabian populations in my studies, some groups have been found to be minimally admixed. When I studied the ancestral components present in Levantine and southern Arabian populations, I found that many of the Yemeni from the Mahra governate were largely non-admixed and (almost) all of their ancestry derived from the Arabian IAC. All the sample donors from the Mahra governate reported Mehri (i.e., a Modern South Arabian language spoken in parts of Yemen and Oman) as their primary language; the people of Mahra are thus linguistically isolated from the neighboring Arabic-speaking groups (Rubin 2008; Rubin 2010).
Had there been a pronounced Sub-Saharan influence among the Proto-Cushites, it likely would also have been present in the affiliated Modern South Arabian populations. Instead, the opposite is true: Mehri, as a representative MSA group, are the least admixed Semitic-speaking population. Vyas’ finding therefore supports the view that the Proto-Cushites were not significantly admixed with any Sub-Saharan groups. (*N.B. A few Mehri also settled in the Horn, where their descendants are today represented by the Carab Salaax. This tiny group numbers a few hundred individuals, who dwell in the northeastern Puntland region of Somalia.)
The Makalia skull, which belonged to an early Cushitic settler of the Pastoral Neolithic. Although since lost, Louis Leakey (in the cranium’s original form) and G. P. Rightmire (in its cast form) observed that the specimen shared closest morphological ties with “Mediterranean Caucasoids.”
Only around 1,300 years after the earliest radiocarbon-dated Pastoral Neolithic sample does the first probable evidence of Nilotic influence, in the form of the Y-DNA haplogroup A and the mtDNA clade L4, appear among specimens excavated at the Naishi Rockshelter and Keringet Cave. As the years pass, we can see the Afro-Asiatic-associated mtDNA lineages further giving way to hunter-gatherer and Nilotic-affiliated clades. The Great Lakes would, by the Iron Age, eventually become dominated by its current primary occupants, the Bantus (descended from the Iron Age/IA culture makers) and the Nilotes (descended from the Pastoral Iron Age/PIA people). This gradual population replacement of that region’s early Cushitic pastoralists is also reflected in the osteological record (see G. P. Rightmire’s analysis on The Elongated African fallacy).
Wang et al. (2020) analysed additional genomes belonging to the ancient Cushitic pastoralists of East Africa. Like Prendergast et al., the researchers found that the earliest of these Cushitic settlers largely bore North African-related ancestry. They also formulated a three-way admixture model using Levantine Chalcolithic, Dinka and Mota specimens as hypothetical ancestral populations to the ancient Cushitic herders. However, thanks to Wang et al.’s improved and more broad-based dataset (which includes several new Pastoral Neolithic samples from Kenya), we are now able to confirm that the first Cushitic arrivals in the region indeed were not significantly admixed with Nilo-Saharan speakers or other Sub-Saharan groups. Admixture analysis instead shows that the early Cushitic pastoralists had almost identical ancestry proportions as modern Afro-Asiatic-speaking North African populations, deriving around 80% of their genome from a West Eurasian ancestral population (red component). Furthermore, according to both Wang et al.’s admixture analysis and three-way admixture modeling, these earliest Cushitic settlers had just 5%-10% of supposed admixture from Nilo-Saharan peoples (purple). This completely debunks the claim that the Proto-Cushitic speakers, who evolved in the Egypt/Libya/northern Sudan area and subsequently migrated southwards, carried significant ancestry related to Nilotes. The later group of Cushitic Pastoral Neolithic herders were very similar to the earlier Cushitic pastoralists in that they derived ~70% of their ancestry from a North African/Levantine-related ancestral population. At the time of their first settlement in the region, these ancient Cushitic peoples would have borne even higher non-African ancestry (~90%). We know this because they were also found to carry some East Africa-specific forager admixture (blue component), which they must have picked up locally after their arrival (cf. Supplementary Material; Tables S1 to S10).
Moreover, Wang et al. examined two of the ancient Cushitic settlers for phenotype alleles (a 2300 year old sample from Hyrax Hill and a 1500 year old sample from Molo Cave, both in Kenya). The scientists reported that both of the specimens bore the derived allele of the SLC24A5 gene, which is associated with lighter skin pigmentation. This suggests that the individuals had a light brown complexion (cf. Table S7).
(*N.B. Wang et al. also compared the ancient Cushitic pastoralists’ genomes with those of various modern samples, including individuals from the Cushitic-speaking Afar, Agaw, Somali and Oromo groups (cf. Supplementary Material). The researchers did not include any Abyssinian/Habesha/Ethiosemitic-speaking samples in their admixture analysis, presumably because Abyssinians are not the best representatives of the early Cushites since they interbred with Semitic and Omotic peoples (Wang et al. genomically model the Agaw sample as having 5.6% more Levant Chalcolithic ancestry than the Amhara test group due to the Amhara cohort’s higher Omotic/Mota-related admixture; see Tables S1 to S10). In the admixture analysis, the Afar and Agaw samples had virtually the same estimated ancestry proportions as the later Pastoral Neolithic samples. However, the Oromo and Somali samples had slightly less of the North African-related component. This is probably because the Ajuran and Garre Somali clans in Kenya have intermarried with Borana Oromos (cf. Helland (1980)), and Borana Oromos have some Omotic admixture (the Somali sample in Wang et al. (2020) was taken from Prendergast et al. (2018), who used the Simons Genome Diversity Project’s Somali sample; this SGDP sample was collected in Kenya by George Ayodo). Future analyses should be sure to incorporate Oromo samples from central Ethiopia, which are less affected by Omotic admixture. They should also include ethnic Somali samples from the overlooked Puntland and Somaliland regions in northern Somalia.)
Young ethnic Somali men in the southern Jubaland region of Somalia (top), young ethnic Somali women in the northeastern Puntland region of Somalia (center top), young ethnic Somali women in the northwestern Somaliland region of Somalia (center bottom), and young Bilen/northern Agaw women in Eritrea (bottom). Genetic analysis has shown that around 60% of Cushitic, Ethiosemitic and North Omotic-speaking individuals in the Horn of Africa carry the derived allele of the SLC24A5 gene. This lighter skin conferring variant has been measured to have an effect on pigmentation of 7.6–11.4 average melanin units (cf. (Beleza et al. (2013)).
The general morphology of the skeletal remains, expressed in classical racial terms, showed a somewhat different picture. The results of the physical anthropological study pointed to the prevalence of the Europoid type (84.5%) in the burial complexes CI-III as well as in the N cemetery, with a small share of mixed (11.4%) and Negroid (4.1%) types. On the other hand, people buried in the A cemetery were mostly of an intermediate morphology (57.1%) with more Europoids (33.3%) than Negroids (9.6%).
Of the few “Negroid” individuals living amongst the mostly “Europoid” Pan-Grave culture bearers, research has found that the former showed archaic characteristics, possessing robust bones with strong muscular attachments as well as distinctly “Negroid” features and woolly hair (Strouhal and Jungwirth (1984); Säve-Söderbergh (1989); Friedman (2001); Brunton (1937)). By contrast, individuals belonging to the Afro-Asiatic-speaking Kerma, C-Group and ancient Egyptian cultures generally had lithe builds, with “Caucasoid” features and non-woolly hair (for specifics on the physiognomy of the Kerma and C-Group culture bearers, see The Elongated African fallacy). Furthermore, Strouhal and Jungwirth (1979) compared the discrete/non-metric osteological traits of the mainly “Europoid” individuals from the three CI-III Pan-Grave cemeteries at Sayala against those of the Pan-Grave specimens from the N cemetery and A cemetery, as well as against those of various ancient Egyptian series (Badari, early Naqada, late Naqada, Hierakonpolis, Abydos, Tarkhan, Sedment, Qurna, Qau, Giza and Hawara), a C-Group Nubian sample (thought to represent early Berber-speaking inhabitants of the Nile Valley), a modern East African “Negroes” sample (Rwanda and Burundi), and a modern South African “Negroes” sample (Zulu, Xosa, Sotho and Venda). The researchers observed that the CI-III specimens were morphologically distinct from all of these comparative samples, a fact which they suggest indicates that the Pan-Grave people of the CI-III burial complex likely represent the Blemmyes of the Eastern Desert. According to Winstedt (1909), an early Coptic inscription refers to the Blemmyes as the Bougaioi. During the later Aksumite era, this name is rendered as Beya in an Adulis text — a clear allusion to the Cushitic-speaking Beja people. This morphological dichotomy between the Pan-Grave culture’s “Europoid” Cushitic majority and its assimilated “Negroid” Nilotic minority is also reflected in ancient artwork. To this end, a 12th Dynasty Nile Valley mural depicts a “Caucasoid” Cushitic Beja figure, whereas a painted ox skull from the same epoch portrays a “Negroid” Nilotic figure.
A Somali man, retaining the fine features, facial orthognathism and soft-textured hair of his Cushitic ancestors.
The alveolar index (gnathic index) among global populations, which measures levels of prognathism (facial projection). The Somali (94), ancient Kerma (96.7), and early and modern North African samples have a similarly low alveolar/gnathic index as the ancient and modern European samples (Hanihara (2000)).
Interbreeding between European individuals and Nilotic or Bantu individuals will not produce any of the three main phenotypes that are found today among the northern Cushitic and Ethiosemitic-speaking populations in the Horn of Africa (see discussion below on the mariin, cad and yusuur phenotypes). This obvious fact is frequently overlooked by geneticists when constructing their statistical admixture models, probably because they often are not trained in physical anthropology.
Rather than anthropometrically clustering in between Europeans and Nilotic/Bantu individuals like actual biracial persons do, the Afro-Asiatic speakers in the Horn region group with other Afro-Asiatic speakers in North Africa and the Arabian peninsula (cf. Billy (1988); Leguebe (1981)). Instead of having nasal indices between those of Europeans and Nilotes/Bantus like actual biracial persons have, the nasal indices of the northern Cushitic and Ethiosemitic-speaking populations also fit comfortably within the “Caucasoid” range (Olapido et al. (2011)). Likewise, Gedda (1961) indicates that “the nasal breadth of the American Negro (40.9) is much less than in African Negroes (42.4-45.5), that for White Americans being 34.9 and for Somali only 34.7.” Skeletal analysis of ancient Cushitic specimens by Louis Leakey observed similarly leptorrhine (narrow) nasal indices (see The Elongated African fallacy). This was, therefore, the original nasal form of the Cushitic peoples.
Oromo women, possessing the “Caucasoid” physiognomy and soft-textured hair characteristic of “pure” Cushitic people.
— Somali men with the two most common hair forms distributed among ethnic Somali males (over 80% in non-Sab clans). These consist of wavy strands and frizzy curls, both of soft-texture (jileec) (Coon (1939); Puccioni (1937)).
Hair form provides a similar indication, for Coon (1939) notes that approximately 86% of the Somalis he examined had soft-textured curly, wavy or straight hair (also see Charpin and Georget (1977) chart above). The autonym jileec (“soft-haired”), which ethnic Somalis traditionally use to describe themselves, suggests that this was the original hair form of the Cushitic peoples (Bernus (2000)). Indeed, the medieval Portuguese missionairies João dos Santos and Jerónimo Lobo state that the Maracatos (Katwa or Garre Somali) — notorious slave traders who castrated their slaves (cf. Steiner (1999)) — that they encountered in southern Somalia were tall, swarthy people with fine-features and lank hair (cabello corredio). We also know from forensic analysis that the early Cushitic speakers of Kerma in Sudan almost unanimously possessed non-kinky hair texture (Lepsius (1915)). Contrarily, many if not most European-Nilotic/Bantu biracial individuals have kinky hair texture like their Nilotic/Bantu parent (this often does not apply to New World biracial persons, who, due to the extra European admixture and Native American admixture that is already present in their African American/Afro-Caribbean parent, may have soft-textured hair).
Ancient Egyptian engravings of an early Cushitic individual (left) and an early Nilotic individual (right). The contrast in physiognomy between the two is evident, pointing to different ancestral origins. The Cushitic individual is fine-featured and lean, and has soft-textured wavy hair, medium-sized and non-everted lips, and an orthognathous profile. Conversely, the Nilotic individual has broad and flat facial features, plaited kinky hair, large everted lips, and is markedly prognathous.
A Hadendoa Beja man preserving in all essential aspects the physiognomy of the engraved ancient Cushitic figure above.
Furthermore, instead of sharing at least some of the prognathism (facial projection) that characterizes Nilo-Saharan/Niger-Congo groups, the Afro-Asiatic speakers in Northeast Africa are among the most orthognathous populations in the world. Hanihara (2000) thus observed that the alveolar index (gnathic index), which measures prognathism levels, is particularly low among his Somali sample (94); the same goes for the ancient and modern North African samples, including the Kerma cohort (96.7). Leigh (1934) similarly indicates that “the ancient Egyptians have a low gnathic index, my average for sixty-nine males being 95.2, and for seventy-one females, 95.6.” By contrast, Wellcome Tropical Research Laboratories (1908) report that the Nuer and Bari Nilotes of the Sudan area have a “very considerable” gnathic index of 106.6 and 109, respectively. This is an even greater level of prognathism than that measured among the Niger-Congo-speaking samples. In fact, some of the highest gnathic values ever have been recorded among the Nilotic peoples. Lockyer (1910) reports on the skull of a Nilotic woman (dated to c. 2nd-4th centuries CE), which had an extreme gnathic index of 123.3, almost 19 points higher than the 104.4 prognathous average of African “Negroes.” (*N.B. With regard to the alveolar index or gnathic index, craniofacial profiles that are orthognathous=below 98, mesognathous=98.1-103, and prognathous=above 103.)
Structural genome analysis of Afro-Asiatic-speaking and Nubian populations in Northeast Africa. At K=7, Cushitic ancestry (yellow component) is maximized among the Somali North sample (Ethiopian Somali). The Somali South sample (Sab Somali) is most similar to the Somali North sample, but differs from it by having greater Sub-Saharan admixture (Nilo-Saharan/orange & Niger-Congo/purple components) (Vicente et al. (2019)). This is consistent with anthropometric (Billy (1988)) and craniometric (Rightmire (1976)) analyses of these groups. (*N.B. Vicente et al. (2019) indicate in their Table S1 that their “Somali North” sample was taken from Gurdasani et al. (2015). As explained above, Gurdasani et al. stipulate that “although these samples were collected from Ethiopia, many of these individuals are from Somalia (Mogadishu).” Hence, Vicente et al. (2019)’s “Somali North” sample is in actuality primarily a Mogadishu Somali or South-Central Somali sample (i.e., Hawiye Somali). Furthermore, Vicente et al. state in their Table S1 that their “Somali South” sample was borrowed from Triska et al. (2015). Triska et al., in turn, note in their own Table S1 that their Somali sample consists of 11 individuals who speak an Afro-Asiatic language, practice a sedentarian subsistence system, and originate from the geographical coordinates of 42.63 longitude and 2.00 latitude. GPS-coordinates.org specifies that these geographical coordinates correspond to the town of Diinsoor, located in the southwestern Bay region of Somalia. This is an area almost exclusively inhabited by the Rahanweyn or Sab Somali agropastoralists. I. M. Lewis writes that “the Sab are a conglomerate people, an amalgam of many different Somali groups with Galla and negroid elements,” thus explaining the excess Sub-Saharan African admixture in this “Somali South”/Sab cohort.)
The anthropometric data measured by Puccioni (1931) and Coon (1939) suggests that the populations responsible for the African admixture component in the Horn of Africa were generally of short stature like the Mota, Gogoshiis Qabe and Buur Heybe hunter-gatherers rather than of tall stature like Nilotes. This is indicated by the fact that the most African-admixed groups among the Afro-Asiatic speakers in both Ethiopia and Somalia also happen to be the shortest on average (viz. Sidama and Sab), although they are still taller than the local foragers (viz. Chabo, Sabue, Eyle, Boni, Watta and Ribi; Puccioni reports an average height of 165.15 cm/1651.5 mm for the Ribi). Conversely, Puccioni observed that there was a positive correlation between high stature, lighter skin pigmentation and non-woolly hair texture in his cross-analysis. Patenostre (1922) likewise found that his Somali sample had a crural index or tibio-femoral index (which estimates leg limb proportions) of 83.2, similar to the 83.4 average of the Berber and Arabic-speaking populations of North Africa. Both Afro-Asiatic-speaking groups were also distant from the “Negro” sample, which had a crural index of 86.8. Moreover, genome analysis of the Early Pastoral Neolithic specimens of Kenya, who are the oldest Cushitic settlers in eastern Africa to be genetically analysed, indicates that they bore little Nilo-Saharan-related ancestry (8.5% on average). Since the rest of their Sub-Saharan admixture consists of a minor East African hunter-gatherer element (~14.7%), which is native to the region, this implies that these first Cushitic arrivals in eastern Africa would have borne almost exclusively non-African ancestry at the time of their first settlement in the area (~91.5%). Taken together, this supports Carleton Coon’s theory that the moderately tall height of modern Cushitic speakers was directly inherited from their “Caucasoid” Cushitic ancestors as opposed to acquired by mixing with Nilotes (as Coon phrased it, “there can be no doubt that the tall stature of the Gallas, Somalis, and Agaus is an old Hamitic trait[…] The bodily build of the East African Hamites is typically Mediterranean in the ratio of arms, legs, and trunk”). (See Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa for details.)
Crural index (tibiofemoral index) of various populations in Africa and Europe (Patenostre (1922)).
Tuareg Berber nobles of the Kel Tamasheq confederation.
Among the northern Cushitic groups in the Horn region, three main phenotypes are distinguished: mariin (“red-brown”), cad (“light”) and yusuur (“ebony”). The mariin phenotype is the most widespread, and is characterized by tall stature, leptorrhine facial features, copper-brown skin complexion, soft-textured curly, wavy or straight hair, medium-sized lips with no eversion, dolichocephaly, orthognathism and oftentimes slightly oblique eye folds (cf. Puccioni (1931); Gallo (1979)).
Other examples:
Muna Khalif (Tuni, Rahanweyn Somali). Although cad individuals in Somalia are concentrated in the northern Puntland and Somaliland regions, a number of persons with this phenotype, such as fashion designer Muna Khalif, can also be found in southern areas.
The cad phenotype is essentially the same as the mariin one, with the exception that cad individuals have a distinctive olive to light brown skin tone. This physiognomy can be found throughout the Horn, but is largely concentrated in northern Somalia, eastern Ethiopia, and parts of Djibouti and Eritrea. Cushitic individuals with the mariin and cad phenotypes, the “pure” Cushitic physiognomies, have traditionally been hailed for their good looks. They were often described in the literature as beautiful, handsome or prepossessing (e.g. the explorer Thomas Heazle Parke writes: “The Somali’s type of feature is Asiatic[…] These Somalis are also very agile in their movements–compared with the Negroes[…] Their skin is of a coppery tint; the natural expression of the face is strikingly bright–with quick, dark, and very mobile eyes–and, altogether, their appearance is prepossessing”). This custom dates back to Herodotus, who asserted that the ancient Macrobians (thought to be an early Cushitic people) were the “handsomest race of men in the world.”
The yusuur phenotype also resembles the mariin one, except that individuals with this physiognomy frequently show traces of admixture with the aboriginal hunter-gatherers of the Horn in that they have a blackish-brown skin tone and sometimes shorter height and frizzier hair. This phenotype occurs across the region, but is most common in southern Somalia, particularly among the Sab (Puccioni (1931); Coon (1939)).
According to Coon, the mariin and cad phenotypes found especially among Somalis closely resemble the amrani and Ihaggaren (beidan) phenotypes, respectively, which are characteristic of the Tuareg Berber nobles in the southern Maghreb.
Serological analysis of the Tuareg Berbers, the Ikelan (the slave class within Tuareg society), and other populations of the Sahara and Sahel. The Tuareg individuals cluster with the Arabic-speaking Maghrebi samples. The Ikelan group instead with the Niger-Congo-speaking Serer, Wolof and Toucouleur (Haalpulaar), and more remotely with the Fulani (Peul) of Senegal and the Haratin (Froment (1999)).
Note that the amrani/mariin and beidan/cad phenotypes are not characteristic of the Tuareg’s slave class, known as Ikelan, whose members instead have a “Negroid” physiognomy (see Coon’s discussion below). Genetic analysis has confirmed that the Ikelan are of West African origin since they carry the E1b1a paternal haplogroup, which today is the modal clade among Niger-Congo speakers. The Tuareg nobles, on the other hand, are of Berber origin. They thus primarily belong to the E1b1b lineage, which is typical of the Afro-Asiatic-speaking populations in North Africa and the Horn of Africa (cf. Ottoni et al. (2011)):
The Tuareg nobles are tall men, with mean statures running tribally from 172 cm. to 178 cm.; about 174 cm. would be their total mean. They are lean, long-armed, and long-legged, with narrow shoulders, narrow hips, and chests which are narrow in an antero-posterior direction; their hands and feet are long and very narrow, their fingers long and thin. The very fine wrists and ankles which we have observed among the Somalis are also present here. The addition of Negro blood to this Tuareg bodily type broadens the shoulders, shortens the legs, and makes the hands and feet wider and larger. The Tuareg relative sitting height mean of 49, indicating that the sitting height is less than half the stature, serves to illustrate the extremely linear constitutional type of this people. The heads of the Tuareg are dolichocephalic and large; tribal means in the cephalic index vary between 72 and 75, but 73 is the central point of the whole. No brachycephals are found among the white nobles, although they occasionally appear among negroids of other classes. […]
The Tuareg in their pure form belong to a specialized Mediterranean sub-type, the creation of which is partly a matter of isolation and selection under extreme environmental stimuli. They resemble the East African Hamites very closely, and especially the whiter element among the Somali, but in their extreme stature and great head size they seem closer than most other living Mediterraneans to the pre-Neolithic East African men.[…]
the unexposed skin of the non-negroid nobles and Imrad is seen to be a brunet-white, without brownish tinge; the mixed bloods, however, who are predominantly Tuareg and only in a minor degree negroid, assume a constant and characteristic dark brown color, known in North Africa as amrani, and comparable to the characteristic hue of the Somalis.
The persistence of the mariin and cad phenotypes today in the Horn of Africa further supports the idea that the earliest Cushitic settlers in the region were not significantly admixed with Nilotic populations, for, if they had been, their modern descendants would not continue to resemble the amrani and beidan Tuareg Berber nobles so closely.
Somali model Jawahir Ahmed (Darod clan), center, and North African contestants at a beauty pageant. Cushitic individuals with an cad phenotype have a similar complexion as North African individuals with a beidan phenotype, though they are often taller like the Tuareg Berbers.
Distribution of oblique eye folds and other forms of palpebral fissure (eye slits) among individuals in southern Somalia (Gallo (1979)).
Intermarrying with adjacent Arab or Persian groups is also clearly not responsible for these phenotypes. Gallo (1979) reports that most individuals in southern Somalia have either oblique eye folds (29.9%), slight Mongolian eye folds (19.9%), or external eye folds (19.9%), with the remainder possessing normal palpebral fissures/eye slits (36.6%). However, the southern Arabs generally do not have oblique eye folds nor are they particularly tall, and most are instead brachycephalic (broad-skulled). While Persians are frequently dolichocephalic (long-skulled; the cephalic/cranial index being a key difference between them and the southern Arabs), they too do not tend towards high stature or almond-shaped eyes (cf. Coon (1939)). (For more details, see Are you sure that the oblique eye folds, which are common among Afro-Asiatic speakers and Nubians in the Horn and North Africa, are not due to absorption of Khoisan peoples? What makes you ascribe this trait to Central Asian or East Asian admixture? and Is there any other evidence of an ancient Central Asian or East Asian-related presence in the Nile Valley?.)
Ethnic Somali women in northern Somalia. Almond-shaped eyes, which frequently characterize Berber individuals with the beidan phenotype, are also rather common among Cushitic individuals with the “pure” cad and mariin phenotypes. Ancient DNA analysis has confirmed that this similarity, like the resemblance between the mariin and amrani phenotypes, originates from the fact that the ancient Cushites were a population of North African ancestral stock.
Somali man with oblique eye folds. This form of palpebral fissure or eye slits likely evolved as an environmental adaptation among early Afro-Asiatic-speaking populations in the Sahara. Such eye folds are still rather frequent among Cushitic and especially Berber speakers; particularly individuals with the cad/beidan and mariin/amrani phenotypes.
Cephalic indices of populations in Northeast Africa, the Arabian Peninsula and Near East. The Cushitic and Ethiosemitic speakers and the North Arabs (Bedouins) are dolichocephalic (CI ≤74.9), whereas the South Arabs and Armenians are brachycephalic (CI ≥80).
Moreover, recent Arab and Persian admixture among Cushitic speakers in the Horn is primarily confined to the Benadiri Somalis (Reer Xamar or Xamar Cad-Cad), the latter of whom dwell in the coastal Benadir region in southern Somalia rather than in the northern parts of the country (Puntland and Somaliland territories) and other areas where cad individuals are concentrated. The early Benadiri intermarried with Arabian and Persian merchants and established the Somali-Arab-Persian ruling dynasties of the Mogadishu Sultanate. In this regard, the 1st century CE Periplus of the Erythraean Sea indicates that the ancient Azanians — the Cushitic ancestors of the Benadiri Somalis and the Bravanese, a subgroup of the Benadiri based in the port town of Brava — interbred with Arab traders on the coast of southern Somalia. This is why, to this day, especially tall Benadiri/Bravanese individuals can still be found (i.e., the great height was inherited from the Benadiri/Bravanese’s “giant” Cushitic Azanian progenitors, the latter of whom the Periplus describes as being “very great in stature”). Ibn Battuta also informs us that at the time of his visit in 1331, the Sultan of Mogadishu was one Abū Bakr ibn ‘Umar, a “Berber” originally from northern Somalia (Upshur et al. (2011); Freeman-Grenville (1963)). This first dynasty of the Mogadishu Sultanate, known as the Fakr ad-Din dynasty, was succeeded after 1500 by al-Muẓaffar dynasty (Freeman-Grenville (1963)). Correspondingly, Immel and Kleiber (2009) observed that most Benadiri individuals today carry the E1b1b and T paternal haplogroups like other ethnic Somalis (~52%; 35.48% E1b1b and 16.13% T), albeit with notable frequencies of the Arab-mediated J clade (~29%) and the Persian-affiliated R lineage (~6%). Genome analysis of modern Benadiris similarly indicates that these individuals mostly bear Cushitic-related ancestry (averaging ~50%), with significant western Asian admixture derived from peninsular Arabs (~25%) (see details on Who were the ancient Azanians?). Benadiri (as Arabized Cushites) are thus the Horn of Africa’s equivalent to North Africa’s Arabic-speaking populations (Arabized Berbers).
Admixture analysis of a Bajuni individual of Swahili type from Somalia. The person (labeled S25) was found to mainly harbor Bantu-related ancestry (light green component) with minor West Eurasian admixture, like other Swahili speakers from the Comoros and East Africa. However, unlike the Comorian Swahilis, the West Eurasian admixture borne by this Bajuni individual and the other East Africa Swahili samples is primarily associated with Cushitic-speaking populations of the Horn region (dark grey component) rather than Arabs (red component) or Iranians (light pink component) (Brucato et al. (2019)).
Benadiri/Bravanese are also ancestrally distinct from the Swahili. Although the Bravanese recently adopted the Swahili language during the Zanzibar Sultanate’s presence in Benadir, other Benadiri still speak their native Cushitic tongue (Somali). By contrast, the Swahili are people of Bantu origin and language, who have some Cushitic, Arab and Persian admixture; particularly among the Bajuni Swahili in southern Somalia. Brucato et al. (2018) observed that Swahili Bantu individuals of coastal East Africa (Mombasa, Lamu and Kilifi in Kenya) and Swahili Bantu individuals of the Comoros Islands predominantly bear Bantu-related autosomal DNA (~75%), with some minor Eurasian admixture derived from absorbed Cushitic peoples and Austronesian, Arab & Iranian peoples, respectively. Raaum et al. (2017) likewise report that the Swahili carry diverse Y-DNA haplogroups (including the Arabian-mediated J clade and the Cushitic-affiliated E1b1b and T lineages), though their most frequently occurring paternal haplogroup is the Bantu-associated E1b1a clade. The mtDNA of both the early Swahili (74%-89%) and modern Swahili (93%) almost entirely consists of African derivatives of the macrohaplogroup L, which are shared with neighboring Bantu and Nilotic populations (also see Msaidie et al. (2011)). Lastly, ancient DNA analysis of Swahili individuals at various Middle Ages coastal sites in Kenya and Tanzania indicates that they sustained considerably more gene flow from western Asia than their comparatively less admixed modern Swahili Bantu successors (50% Near Eastern-related admixture for the former vs. under 25% for the latter). The west Asian admixture that these medieval Swahili specimens carry is also primarily associated with Persians, whereas contemporary Benadiri/Bravanese Somalis to the north mostly derive their west Asian admixture from peninsular Arabs (cf. Brielle et al. (2022), Proctor et al. (2015), and Sloan (2010) for more ancient DNA of the early Swahili Bantus.)
Additional evidence that the mariin and cad phenotypes were not introduced by Arabs or Persians comes from historical texts. For example, in the 16th century Andalusian explorer Leo Africanus informs us that “the people of Adel are of the colour of an olive.” This is an allusion to the denizens of the Adal Sultanate, which he describes as “a very large kindome, and extendeth from the mouth of the Arabian gulfe to the cape of Guardafu.” The Portuguese missionary Francisco Alvares, a contemporary of Leo Africanus, confirms that indeed “the kingdom of Adel (as they say) is a big kingdom, and it extends over the Cape of Guardafuy, and in that part another rules subject to it.” Correspondingly, this realm is the exact general area in northern Somalia (Puntland and Somaliland), Djibouti and eastern Ethiopia where Cushitic individuals with the cad phenotype are still concentrated. According to the chronicler Shihāb al-Dīn’s coeval treatise Futuh Al-Habasa (“Conquest of Abyssinia”), the Adalites largely consisted of ethnic Somali clans, among whom were: the Gorgorāh Dir; Harti Darod (Warsangali, Majerteen, Dhulbahante), whom he calls the “people of Mait” after the northwestern port city of Mait; Marraiḥān Darod; Girri Ogaden Darod; Yabarre Ogaden Darod; Hawiya; and Habr Maqdi Isaaq.
(*N.B. Leo Africanus asserts that the Adalites of the Horn were of Arabian origin. In connection with this claim, modern genome analysis of ethnic Somali individuals born in the northeastern Puntland region of Somalia, who mostly belong to the Majerteen subclan of the Darod, has detected a predominant non-African ancestry. This Eurasian ancestry is comparable in frequency to that carried by North African individuals. Following Clark (1954), we suggest that this is because these northern Somali individuals are of relatively pure Cushitic stock rather than of recent peninsular Arab origin. For details, refer to Which Afro-Asiatic-speaking population of the Horn region has the most non-African ancestry?. Also see I have heard that the V12 and V32 subclades of the paternal haplogroup E1b1b, which are today common among Afro-Asiatic speakers in Northeast Africa, are old Arabian lineages. Is this true?.)
Oral testimonies transcribed by colonial officials in British East Africa during the early 20th century are in a similar vein. One such recorded eyewitness account from a Karimojong Nilote elder describes a cattle raid in the late 1800s that his tribe had made against one of the last remaining Cushitic groups in Kenya (which has since been assimilated into a hunter-gatherer community called the Oropom). He avers that this pastoralist population was “very light skinned and had pronounced slant eyes[…] The men wore their hair long in a pigtail, like Indian women” (cf. Wilson (1970)). Evidently, this is a description of the “white Galla,” whom J. C. Prichard and other early European adventurers reported having encountered in the area i.e., Cushitic individuals with the cad phenotype.
![Berber dancer and actress Sofia Boutella. Note the close likeness to Jawahir Ahmed. This calls to mind Coon (1939)'s assertion that "the Tuareg in their pure form belong to a specialized Mediterranean sub-type[...] They resemble the East African Hamites very closely, and especially the whiter element among the Somali."](https://landofpunt.files.wordpress.com/2015/11/berber-beidan.jpg?w=209)
11 Saturday Mar 2023
Tags
A. Batrawi, A. E. Mourant, A. M. El Hassan, Ala111Thr, Archaeology, Bantu, Bertram Thomas, C-Group, Capsian, Carleton Coon, Caucasoid, Charles Gabriel Seligmann, Christopher Ehret, Daniel Stiles, East Africa, Eburran, Elizabeth W. Ikin, Elmenteitan, Elongated African, G. Billy, G. P. Rightmire, George P. Murdock, Gunter Bräuer, Hamitic, Harold C. Fleming, Hubert Jules Deschamps, J. D. Fage, J. E. G. Sutton, Jean Hiernaux, John Hanning Speke, Joseph Greenberg, K. L. G. Goldsmith, Karl Richard Lepsius, Kerma, Khoisan, Louis Leakey, Marianne Bechaus-Gerst, Negroid, Nella Puccioni, Oric Bates, Pastoral Neolithic, Peter Behrens, Philipp Paulitschke, R. Protsch, Rodolfo Fattovich, Savanna Pastoral Neolithic, Sonia Cole, Steven H. Ambrose, Stone Bowl Culture, W. W. Howells
The Elongated African theory is an evolutionary hypothesis devised by the late Belgian anthropologist Jean Hiernaux. It was introduced in his 1974 book The People of Africa, a work which seeks to explain the existing physical and genetic diversity in Sub-Saharan Africa through various developmental processes. Touting itself as using a then-new non-racial approach, the narrative places an emphasis on environmental adaptation as one of the primary driving forces behind human biological variation.
Hiernaux suggests therein that the swarthy “Caucasoid” (“Hamitic”) phenotypes which typify the Afro-Asiatic-speaking populations in the Horn of Africa and the Moors in the Sahara evolved through interbreeding between, on the one hand, Arabs or Berbers, and on the other, naturally narrow-featured, hypothetical inhabitants of Sub-Saharan Africa that he dubs “Elongated Africans.” To prove his theory, Hiernaux employs a series of logical fallacies, contradictions and factual inaccuracies, many of which were apparent even at the time of publication. For this reason, his Elongated African hypothesis never really took off. It was instead criticized by some of his own colleagues, eventually abandoned by Hiernaux himself, corrected a few years later by more comprehensive anthropometric, craniometric and serological studies, and debunked altogether through ancient DNA analysis. Although now obsolete, the theory remains an instructive example of how excessive post-colonial guilt can easily lapse into faulty science and guesswork.
Development of the theory
Map showing the spread of the Niger-Congo-speaking ancestors of the Tutsi-Hima Bantus and Hutu Bantus from their suggested Original Bantu Homeland on the Nigeria-Cameroon border in West Africa (Jaja et al. (2011)).
Hiernaux was a physician and anthropologist by training. Beginning in the 1950s, he alongside the historian and archaeologist Emma Maquet carried out some of the first excavations in present-day Rwanda, Burundi and the Democratic Republic of the Congo in eastern Central Africa. These Great Lakes territories were at the time under Belgian colonial rule.
Phylogeny of the Niger-Congo language family. Kinyarwanda, the mother tongue of the Tutsi Bantus and their brethren the Hutu Bantus, belongs to the Narrow Bantu branch of the Niger-Congo phylum (Bostoen (2004)).
While stationed in the area, Hiernaux also conducted a series of anthropometric studies on the local Tutsi pastoralist, Hutu agriculturalist and Twa hunter-gatherer populations. Although all three groups speak the same Kinyarwanda language (a member of the Bantu branch of the Niger-Congo family), he argued that they each had distinct origins and suggested that these differing ancestral backgrounds could be discerned anthropologically. To this end, in several of his earlier works, Hiernaux insists that a “Hamitic” influence is evident among the Tutsi despite their linguistic, cultural and physical commonalities with the “Bantu” Hutus and “Pygmy” Twa.
This assertion was in part drawn from the Tutsis’ own longstanding oral traditions, as documented by the explorer John Hanning Speke, among others. These local accounts held that the founders of the various Great Lakes kingdoms were “Caucasoid” peoples, who had arrived in the region several centuries prior from either the Horn or North Africa. The Hamitic migrants are then said to have ruled over the local “Negroid” inhabitants, gradually adopted the latter’s Bantu languages, and eventually amalgamated with the autochthones through intermarriage. These stories are embodied in the person of Kintu, a figurative representation of the early Cushitic settlers in the lacustrine region. A. B. Fisher (1904), writing on the territory north of Buganda, states:
The oldest inhabitants of the country related to me how a white man and woman many years ago landed on the right bank of Lake Albert, and settled in Bunyoro. On being questioned as to whether they referred to Sir Samuel and Lady Baker, they replied that they remembered them well, and described them to me. But, said they, these other white people arrived a long, long time ago, and founded their first dynasty. The story is that this man and his wife came down from the Nile, and caused a great sensation, the people all flocking round them, exclaiming, “Kintu ke?” (What is it?). So they named the man Kintu.
As explained in part two of this essay, there is some truth to the above narrative, for there exists today a minor Cushitic paternal influence among the Tutsi Bantus.
Portrait of Kintu and his wife Nambi. Note the couple’s “Caucasoid” appearance. According to A.J. Mounteney Jephson (1893), this is because Kintu and his bride are the “personification of the influential immigrants from Galla countries” in Northeast Africa, who moved south and settled in the Great Lakes region.
Hiernaux later distanced himself from his initial writings, apparently in response to intensified tensions between the Tutsis and Hutus. In his 1968 work, while ironically decrying what he termed “classificatory mania”, he still maintained that the Tutsi were biologically distinct from the other Bantu-speaking Great Lakes aborigines, the Hutu and Twa. It was around this period when Hiernaux developed what he would eventually call his “Elongated African” hypothesis.
In the first iteration of his theory, Hiernaux argued that the Tutsis were largely of “Ethiopid” origin (traditionally a synonym for “Eastern Hamite”). This ancestral stock, he wrote, was neither Europoid nor Negroid nor a mixture thereof, but rather a race unto itself. By 1972, Hiernaux would assert that there was no significant exotic component in the Tutsi, suggesting instead that they evolved their physique through genetic adaptation to hot and dry conditions. He would also propose that the Afro-Asiatic-speaking populations of the Horn and the Moors of the Sahara were ancestrally formed through a mixture of Arabs and a mysterious, environmentally-molded “African” stock similar to the Tutsi. Hiernaux thus essentially changed the direction of the gene flow, arguing now that it was the Afro-Asiatic speakers that experienced a Tutsi-like influence rather than vice versa.
This second incarnation of his theory was primarily inspired by the work of Hubert Jules Deschamps, a French historian and colonial administrator. Based on anthropometric means and blood work, Deschamps had asserted in 1970 that Somalis, Ethiopians, Moors and certain other groups on the southern rim of North Africa all appeared to have considerable exotic affinities (“Arab” in his nomenclature). He suggested that this ancestral affiliation manifested itself most conspicuously through traits that were not found in combination elsewhere below the Sahara, such as non-kinky hair texture, keen facial features, and oftentimes lighter skin pigmentation. Deschamps, however, conceded that it was impossible to quantify this ancestry without knowing first just what exactly was the nature of the African element in these populations. He also spoke of a Moors-Somali Warsingali “constellation”, and noted that these Afro-Asiatic-speaking groups lived in similar “biotopes” or habitats.
Phylogeny of human stocks in Africa. The Ethiopian “Hamites” and “Mediterranean” peoples of the Horn of Africa and North Africa descend from a common “Proto-Caucasoid” stock (Cole (1954)).
A few years later, the pioneering anthropologist and archaeologist Sonia Cole would invite Hiernaux to write for her Peoples of the World book series published by Charles Scribner’s Sons. Cole had released The Prehistory of East Africa in 1954, a seminal work that provides strong evidence for an early Caucasoid presence in the region as well as in North Africa. She had most recently authored the British Museum of Natural History guide Races of Man (1963), which draws on her many years of field work in Africa and again emphasizes an ancient Caucasoid presence on the continent.
Distribution map of the peoples of Africa, including the “Caucasoid” Hamites and Semites. The Tutsi are among the Bantu-speaking Group of “Negroid” peoples (Cole (1963)).
Against this backdrop, Hiernaux, then serving as the Director of Research at the National Centre for Scientific Research in Paris, would pen his The People of Africa in 1974.
Ancient skeletons in East Africa
Hiernaux starts off by arguing that an ancestral “Elongated African” population can be found in the makers of the Upper Kenya Capsian (Eburran), a Mesolithic hunter-gatherer culture centered in East Africa. He is aware of the Upper Kenya Capsian people’s proposed Caucasoid metrical affinities, measurements which he does not dispute. He also acknowledges that that lithic industry has ties with the coeval Capsian culture of North Africa, whose makers he indicates were gracile “Mediterraneans”:
The makers of the Capsian are less well known physically than the makers of the Ibero-Maurusian. They are Mediterraneans, whose lighter build contrasts with the robustness of the Mechta people. Probably the Berbers are their descendants, with a possible admixture of the Mechta element in some places.
The oldest remains of Homo sapiens sapiens found in East Africa were associated with an industry having similarities with the Capsian. It has been called the Upper Kenya Capsian, although its derivation from the North African Capsian is far from certain. At Gamble’s Cave in Kenya, five human skeletons were associated with a late phase of the industry, Upper Kenya Capsian C, which contains pottery. A similar association is presumed for a skeleton found at Olduvai, which resembles those from Gamble’s Cave[…] The skeletons are of very tall people. They had long, narrow heads, and relatively long, narrow faces. The nose was medium width; and prognathism, when present, was restricted to the alveolar, or tooth-bearing, region. Many authors regard these people as physically akin to the Mediterraneans, hence the label of ‘Caucasoids’ (or European-like) generally attached to them.
Where Hiernaux differs is in the actual origin of those Caucasoid osteological affinities. He suggests that those traits developed in situ in East Africa, independently from the Capsian Mediterraneans in North Africa. The reason why Hiernaux asserts this is because he believes that all of these features can be found in the modern Tutsi Bantus and Maasai Nilotes of the Great Lakes, two populations that he indicates (wrongly, as it turns out) have little-to-no exotic biological influences.
Hiernaux’s first error is in assuming that the Gamble’s Cave skeletal remains of his day were in the same condition as when Louis Leakey first excavated and described them in 1928/29. These fossils were from the start fragmentary, and only two of the five could later be reconstructed. The specimens incurred further damage when the Royal College of Surgeons, where they had subsequently been stored, was bombed during WWII. For this reason, the anthropologist G. Philip Rightmire — who, unlike Hiernaux, had the opportunity to study the remains in the 1970s — correctly notes that no firm conclusions on the fossils’ affinities could by then be ascertained:
Deposits belonging to this “Gamble’s Cave Shoreline” complex have now been dated to between 8000 and 10,000 B.P. Of the five Gamble’s Cave skeletons, only two could be reconstructed, and this job was carried out in England after the material had been sent there from East Africa. Results were certainly far from perfect, owing to warping and crushing of the original bone, and further insult was to follow. The Royal College of Surgeons in London and the skeletal collections housed there received heavy bomb damage during World War II. So by the time that the skulls were transferred to the British Museum (Natural History) in 1948, they were scarcely in mint condition. Skull number 4 is the less well preserved of the two, and all of the base as well as a substantial portion of the facial skeleton are present only in plaster. Distortion renders this specimen quite unfit for measurement. Number 5 also lacks much of the skull base, and the missing parts have been heavily reconstructed. Although these skulls have been called non-Negro in morphology, the evidence is certainly far from clear cut, and any such diagnosis is questionable by virtue of the state of the material alone.
The Cro-Magnon (left) and Gamble’s Cave Number 4 (right) skulls. Note especially the distorted eye orbit shape of the latter, heavily reconstructed cranium (Oschinsky (1963)).
Furthermore, Hiernaux mistakenly assumes continuity in the skeletal record from the makers of “the Upper Kenya Capsian of Gamble’s Cave, Naivasha and Olduvai, who may date to about 4,000 BC; [to] the makers of the ensuing mesolithic Elmenteitan culture of Bromhead’s Site; [to] the remains associated with the neolithic stone bowl culture at Hyrax Hill and Njoro River Cave (dated by carbon 14 to 960 BC), and with the more recent stone bowl culture at Willey’s Kopje, Makalia and Nakuru, which almost certainly date from the Iron Age.”
As we saw, the actual affinities of the fragmentary Upper Kenya Capsian remains are uncertain. Because the Gamble’s Cave Number 4 skull was heavily reconstructed, its morphological status is somewhat conjectural. Oschinsky (1963), for instance, remarks that the “Gambles Cave II, No. 4 skull shows lateral compression which has distorted the shape of orbits, the left zygomatic arch is twice as long as the right one, the basic occipital region is displaced to the left, the cranio-facial juncture has been crushed toward the rear of the neuro-cranium on the right side causing the short right zygomatic arch. The same pressure from the front of the skull has caused the alveolar region to be flattened and the palate to be deepened.”
According to the anthropologist Steven H. Ambrose, the industries at the Njoro River Cave, Makalia and several other Rift Valley sites that Hiernaux indicates were part of the Stone Bowl Culture were also actually later expressions of the Elmenteitan culture:
Extraction of the central incisors may prove significant for confirming correlations between modern and prehistoric cultures. Although this is not exclusively a Nilotic practice, it is most common among modern Nilotic populations in Tanzania and Sudan (Kilma, 1970: 8; Murdock, 1959: 173). The central incisors were removed from all 79 crania recovered from Njoro River Cave, an Elmenteitan cremation burial cave (Leakey and Leakey, 1950: 76). They may therefore have been of Nilotic origin. This practice is also evident in an early Iron Age context at Wiley’s Kopje and the Makalia Burial Site, both located on the western side of the Rift Valley (L. S. B. Leakey, 1935: 95, 107-108). The latter site may represent an Iron Age expression of the Elmenteitan Industry (Chittick et al., in press).
I. M. Lewis excavating a cairn at Gaan Libah in northern Somalia (Lewis (1957)).
The Stone Bowl Culture, also known as the Savanna Pastoral Neolithic, is generally associated with early Southern Cushitic settlers. However, just who exactly were the makers of the Elmenteitan culture is a question that, until recently, was disputed. Louis Leakey and Mary Leakey first analyzed human remains at the Elmenteitan-associated Njoro River site in Kenya. They noted that these specimens had a “non-negro” morphology and were markedly different from the present-day Bantu and Nilotic inhabitants, including the groups with some Cushitic admixture. The Leakeys observed that the Njoro River skeletons had a hyper-leptorrhine (very narrow) nasal index of 47.88. By comparison, Leakey and Leakey (1950)’s lowest scoring Bantu/Nilotic sample, the Tanganyika natives, had a platyrrhine (broad) nasal index of 55.4. Hiernaux (1960)’s Iron Age Rwanda sample was also platyrrhine, with a nasal index of 58.6.
Reconstructed necklaces from the Njoro River Cave, an Elmenteitan site of the Pastoral Neolithic. Similar beads have been found in predynastic Egyptian burials (Cole (1954)).
In 1957, I. M. Lewis excavated a series of cairns at Gaan Libah in northern Somalia, which were radiocarbon-dated to a maximum of 250 years before present. The skeletons buried within the graves were subjected to an anthropometric analysis. In stark contrast to the Bantu/Nilotic samples, the Gaan Libah specimens turned out to be hyper-leptorrhine, with a virtually identical nasal index as the Njoro River individuals (viz. 47.8). This only further validated the Leakeys’ theory on the identity of the original Elmenteitan culture bearers. (Note: the nasal indices above were calculated based on the skulls of the deceased; for the nasal indices of living individuals, see below under Anthropometry). Likewise, after examining 16 beads that were recovered from the Njoro cave, H. Beck opined that “some of them show great resemblance to the pre-dynastic Egyptian work” (Leakey and Leakey (1950)).
Analysis of human crania from ancient burial sites in the Rift Valley and those of various Bantu, Khoisan and Egyptian groups. The ancient Cushitic Makalia and Baharini samples cluster with the male and female Egyptian samples. The Rwanda male and female samples cluster instead with the Sotho Bantus and other Niger-Congo-speaking populations (Rightmire (1975)).
In 1975, Rightmire compared human crania from ancient burial sites in the Rift Valley with those of various Bantu (including Rwanda), Khoisan (Bushman, Hottentot), and Egyptian (Egyptian E/Gizeh) groups. He observed that several of the analyzed Rift Valley skulls shared greatest affinities with the Egyptian samples; namely, the Baharini, Makalia I, and (albeit tentatively) Elmenteitan F1 specimens. As Rightmire put it, “there is little doubt about Baharini as Egyptian female.” The rest of the skeletons either clustered with the “negro” crania or were in the process of being assimilated into these Bantu and Khoisan groupings. This reflects the gradual replacement of the early Egyptian-related Cushitic settlers in the Great Lakes area with the incoming Bantu/Nilotic populations; the latter peoples now constitute the region’s predominant inhabitants.
Based on these skeletal affinities as well as population dynamics and historiolinguistics, Ambrose correlates the Caucasoid skulls with the first Cushitic pastoralists and the Negroid crania with those of the ancestral Nilotic populations:
Craniometric studies undertaken by Rightmire (1975b, this volume) suggest that among the crania from Neolithic burial sites in the Rift Valley were representatives of Nilotic Negroid populations, as well as those whose closest correlates are found among prehistoric Egyptian populations. Rightmire suggests that many of the Rift Valley crania may represent speakers of a Nilotic language (Rightmire, 1974; 1975a, b). One cannot at this point correlate each individual cranium with an individual industry, due to small sample sizes and lack of precise information on artifactual associations. However, we may be dealing with as many as three distinct physical types: a Nilotic Negroid, a Cushitic “Caucasoid,” and an indigenous Negroid hunting population (Gramly and Rightmire, 1973). Cushites and Nilotes have a history of several thousand years of contact along a major geographic and ecological boundary that follows the Ethiopia/Sudan and northern Kenya/Uganda border regions (Ehret, 1974b). Archaeological and linguistic evidence demonstrates that the region of contact continued south along the Rift Valley, Kenya, to Lake Eyasi, Tanzania. The Eyasi region is a cul-de-sac where representatives of the four major language phyla of Africa — Khoisan (Hadza and Sandawe), Afro-Asiatic (Southern Cushitic), Nilo-Saharan (Eastern and Southern Nilotic), and Niger-Congo (Bantu) — are spoken today. Racial differences have undoubtedly been minimized by intermarriage during the long period of contact. Thus, although there is a strong skeletal evidence for two different Neolithic populations, with Sudanic and Ethiopian origins, respectively, it may never be possible unerringly to correlate skeletal types with industries.
With regard to the affinities of the makers of the Kenya Capsian/Eburran culture, the historian and linguist Christopher Ehret indicates that the “Mediterranean Caucasoid” skulls that have often been associated with this industry likewise appear to have actually belonged to the Cushitic makers of the Savanna Pastoral Neolithic/Stone Bowl Culture. He suggests that the makers of the Eburran culture were instead likely of Khoisan ancestral stock:
The poorly understood Eburran Industry has misled other people on the basis of a spurious, misinterpreted association of “Mediterranean Caucasoid” skeletons (Leakey 1931; Protsch 1975, 1978), which are more likely to be of Highland Savanna PN origin. This association has led some people to postulate an early center of Cushitic speech in central Kenya (Fleming 1964, 1969, 1976: 265; Murdock 1959: 197). Since there are no skeletons that can actually be attributed to pre-Neolithic industries, it is more likely that the makers of this industry originated from a local population with great antiquity. Given evidence for the formerly widespread distribution of Khoisan languages throughout East Africa, it is more likely to have been spoken by Eburran hunter-gatherers and other pre-Neolithic East African populations than an Afroasiatic language.
Correlation of the Savanna PN “Stone Bowl Culture” with Southern Cushitic groups has been the orthodox interpretation for many years (Sutton 1966, 1971, 1973; Odner 1972; Phillipson 1977a). Southern Cushitic speakers are theorized to have been the earliest food producers in East Africa, possessing cattle, sheep, and goats and probably cultivating grain. They would correlate with the Lowland Savanna PN, the first pastoralists, who made stone bowls and Nderit Ware in the Turkana region between 5,000 to 3,000 b.p. and later, at 3,300 b.p., spread to the highlands of Kenya and northern Tanzania.
This interpretation is not controverted by the skeletal evidence amassed by Rightmire (1975), which shows that many of the Neolithic peoples of the Rift Valley have their closest affinities with Egyptian populations. The skeletal evidence, however, also demonstrates the presence of peoples whose closest affinities were with modern Negroid populations and who were not “Mediterranean Caucasoids,” as Leakey proposed (1935). Thus two groups of people, of different racial and geographic origins, were present in the Rift Valley.
This is in keeping with Ambrose’s assertion that there was a third early physical type present in the region alongside the Cushitic Caucasoids and Nilotic Negroids. Leakey (1936) likewise notes “Bushman affinities” for the Later Stone Age (40,000 ybp-2,000 ybp) inhabitants of the Lake Victoria area. In contrast, he emphasizes that the Gumban/Savanna Pastoral Neolithic makers were of “a physical type which is almost European”. The anthropologist Gunter Bräuer reaches a similar conclusion in his large study on the morphological differentiation of anatomically modern humans in Africa:
although the East African highlands probably cannot be regarded as the centre of differentiation of modern man in Africa, this part of the continent does represent an important region which was inhabited by Europid, Negrid and Khoisanid populations in prehistoric times
Distribution of ancestral populations in Africa circa 9,000 BCE (Bräuer (1978)).
Ambrose further confirms the above when he observes that the chronological date proposed by R. Protsch for the cairn burials at Gamble’s Cave is grossly inaccurate. To this end, Ambrose notes that conventional charcoal dates for the older (and thus deeper) Phase 3 layer at the site range from 8,000 to 8,500 years before present. Protsch, however, had mistakenly suggested that the cairns — which were buried in a deposit above the Eburran’s final/most recent Phase 5A layer; Phase 5A was, in turn, situated around four meters above the Phase 3 layer — dated to a similar 8,020 ybp, give or take a few years. Thus, the cairns are in fact chronologically more recent than even the last Eburran cultural phase, and by extension, so are the skeletons within them. The specimens’ “Mediterranean Caucasoid” morphology therefore indeed likely does not represent the general physical type of the makers of the Eburran culture. It instead appears to have arrived in the region later with the Cushitic makers of the Savanna Pastoral Neolithic/Stone Bowl Culture, as Ehret had correctly deduced. Ambrose writes:
Dates sites comprising Phase 5A of the Eburran listed in Table ie show that this phase began before 2900 B.P. and ended well after 2000 B.P.[…] The same reservations apply to Protsch’s date of 8020±260 (Protsch, 1978: 103) for the cairn burial overlying the Eburran 5A horizon at Gamble’s Cave, as this date is inconsistent with conventional charcoal dates ranging from 8500 to 8000 B.P. on Phase 3 in this site from 4 meters below this horizon. The Gamble’s Cave burials actually overlay the Phase 5A horizon (L. S. B. Leakey, 1931: 117), and are thus later than and unrelated to this horizon, and may not represent the Eburran physical type[…] Therefore, conventional dating evidence indicates that the Mediterranean Caucasoid physical type belongs to the Neolithic era.
J. E. G. Sutton makes the same observation, remarking that Leakey himself indicates in his excavation notes that the burial cairns containing the “Mediterranean Caucasoid” skeletons belong to a separate, much later population than the makers of the Kenya Capsian/Eburran:
The ‘Kenya Capsian’ is a blade-and-burin industry, which in North Africa or Europe would be classified as ‘Advanced Palaeolithic’. In fact, Leakey originally called it ‘Kenya Aurignacian’ by comparing it with French materials, later allowing its redesignation as ‘Kenya Capsian’ on account of Maghrebian comparisons. The type-site (indeed the only site with a collection of reasonable size, satisfactorily stratified and at least cursorily described in print) is Gamble’s Cave (the lowest ‘occupation level’), which Leakey excavated in the late 1920s.[…] The base of these deposits has now been radiocarbon-dated (following a test cutting by Glynn Isaac and Ron Clark in 1964) to the seventh millennium B.C. It was here that Leakey had collected, beside the stone tools and waste in quantity, both fish-bones and a broken harpoon, though this later find was not recognized till thirty years after the excavation![…]
Repeatedly in the literature the makers of the ‘Kenya Capsian’ are described as a ‘tall Caucasoid’ or ‘Afro-Mediterranean’ people, a deduction based on examination of burials which Leakey found while digging Gamble’s Cave. Whether this racial attribution is roughly correct or not is irrelevant here. For, as is plain in Leakey’s ‘diagrammatic section’ and notes of his excavation, these burials were placed in a layer well above that containing the true ‘Kenya Capsian’ materials with the fish-bones, harpoon and ‘dotted wavy-line’ potsherd. The skeletons probably belong to a different population several thousand years later. There is therefore no direct evidence of the physical type of the makers of the ‘Kenya Capsian’.
In the early 1980s, the anthropologist and archaeologist Daniel Stiles set out to assess the affinities of ancient human fossils buried in several Savanna Pastoral Neolithic/Stone Bowl Culture sites as well as in Azanian cairns in the Chalbi Desert. Stiles had helped establish the Department of Archaeology at the University of Nairobi during the 1970s, so he was well qualified for the task. He later published three papers on the excavations, Stone Cairn Burials at Kokurmatakore, Northern Kenya (1981), The Azanian Civilization and the Megalithic Cushites Revisited (1984), and The Azanian Civilisation Revisted (2004).
Daniel Stiles unearthing the 3,500 year old (~1,500 BCE) Savanna Pastoral Neolithic mound cairn, which he indicates is “the oldest dated stone structure in East Africa”.
Many of the skeletons that Stiles excavated were complete and in good enough condition for him to measure their standing height. They were generally quite tall individuals of Caucasoid physical type, consistent with oral and written tradition. Stiles was also able to accurately date the cairns that the specimens were interred in. He found three different types of cairns, each belonging to separate epochs. The oldest cairns were of mound type; they dated to around 3,000 years ago, and were evidently made by early Southern Cushites of the Stone Bowl Culture. The next oldest cairns were of platform type; they dated to about 1,000 years ago and thus were probably made by early Eastern Cushites. The third group of cairns were of ring type and dated to around 600 years ago. Unlike the more ancient specimens in the mound and platform cairns, the skeletons within the ring cairns showed some evidence of Nilotic cultural influence since all of them had their lower incisors removed. Stiles therefore suggests that these more recent cairns probably contained the Cushitic ancestors of the Rendille, who are known to have intermarried and exchanged customs over the years with the adjacent Samburu Nilotes, including the practice of incisor extraction.
On one of the Savanna Pastoral Neolithic-associated mound cairns that Stiles excavated, he writes:
The ~3,500 year old (~1,500 B.C.) mound cairn was almost certainly made by Southern Cushitic speaking ‘Stone Bowl’ pastoralists. A stone bowl fragment was found buried at the base of the cairn, and obsidian tools and goat bones were found inside the cairn. The person buried inside measured 190cm tall (about 6’4”), an incredible height for someone of that antiquity.
In short, Hiernaux was completely mistaken about the affinities of the various ancient skeletons in East Africa. He erroneously assumed that the Kenya Capsian/Eburran, Elmenteitan and Savanna Pastoral Neolithic/Stone Bowl cultures were all made by a single hypothetical population, his “Elongated Africans”. In reality, the Kenya Capsian/Eburran was likely the work of early hunter-gatherer peoples, who, based on their lithic industry, probably had some cultural (if not demic) contacts with the Capsians of North Africa. Ancient DNA analysis has, moreover, proven that the original makers of both the Savanna Pastoral Neolithic and Elmenteitan cultures were early Cushitic pastoralists (see below under Exotic influences). Additionally, archaeogenetics has revealed that the ancient Nilotes were instead responsible for the Pastoral Iron Age sites. The “Mediterranean Caucasoid” morphology is also associated with only one of those groups, the Cushitic settlers of the Pastoral Neolithic.
Three-dimensional stereogram comparing various modern populations with some early specimens from the Great Lakes, Southern Africa and the Nile Valley (Bräuer (1980a)).
That said, just how do these ancient Caucasoid specimens of the Great Lakes relate to the present-day Afro-Asiatic-speaking populations to the north, in the Horn region? Are they, as Hiernaux postulates, ancestral to Cushitic speakers in general? That is, are they the immediate forebears of the Eastern Cushites (like the Somalis, Afar, Saho, Sidamo and Galla/Oromo), Northern Cushites (Beja), Central Cushites (Agaw), and the largely extinct Southern Cushites? Or are they instead early Southern Cushites alone, as Stiles asserts? Hiernaux does not offer a comparative analysis, but thankfully Bräuer does. Bräuer (1980a) finds that his modern Galla-Somali lumped sample is more closely related to ancient Afro-Asiatic groups from the Nile Valley, particularly the predynastic Egyptians of Naqada and the C-Group peoples of Lower Nubia/Northern Sudan. This is true in all three principal coordinate dimensions; notably, in the first axis, which contains most of the variation between the examined populations (26.2%).
Modern haplogroup and lactase persistence allele analyses provide a similar indication, for they establish that the early Afro-Asiatic-speaking presence in the Great Lakes region was mainly represented by the Southern Cushites (see below under physiognomy and exotic influences). These ancient Caucasoids south of the Horn would eventually be absorbed by the aboriginal hunter-gatherer populations and the incoming Nilotic and Bantu groups.
In part two of the Elongated African fallacy, we demonstrate how we know for a fact that the Caucasoid morphology did not develop in situ in East Africa as Hiernaux had proposed. We also show how the physical anthropology and ecology underpinning his theory are likewise flawed, contradictory and generally inaccurate.
In part one of the Elongated African fallacy, we saw how Jean Hiernaux was initially one of the prime exponents of Hamitic scholarship in African historiography. Under post-colonial duress, he gradually distanced himself from his earlier writings and drew inspiration from the work of Hubert Jules Deschamps. We also saw in detail how and why Hiernaux was mistaken about the affinities of the various ancient human fossils in East Africa, specimens which he erroneously assumed were all of the same physical type. In this second and final part of the Elongated African fallacy, we further demonstrate the inherent flaws and contradictions in his hypothesis by deconstructing the actual physical, serological and environmental evidence that he invokes to support it.
Exotic influences
In his book The People of Africa, Hiernaux includes a brief chapter on biological, cultural and linguistic classifications, where he asserts that his “Elongated African” populations are “much too diverse to form a taxon”. He then devotes an entire chapter to his Elongated African theory, where he also discusses the Nilotes. Hiernaux starts off by defining what exactly he believes physically constitutes an “Elongated African”, as well as which four populations today best represent this putative morphology in its unaltered state:
As already discussed in Chapters 5 and 6, a number of African populations have an elongated body build, with narrow head, face and nose. Their skin is dark (in varying degree), their hair is spiralled, and they have thick but not everted lips. In many of these people, such as the Tutsi of Rwanda and Burundi and related Hima of Uganda, the Masai of the East African steppes and the Ful communities of the Western Sudanic savanna, there is no evidence of an exotic (Arabic or North African) element in their gene pool. Their physical features can best be explained in terms of genetic adaptation to dry heat. Apparently they represent the result of a peculiar evolution in the semi-arid crescent which caps sub-Saharan Africa to the north and north-east.
A Nilotic Maasai man. The Maasai are sometimes termed “Hamiticized Negroes” due to the fact that their Nilote ancestors in the Great Lakes region assimilated some earlier Southern Cushites.
Right off the bat, Hiernaux makes a fundamental factual error that all but invalidates his theory. He mistakenly assumes that the Tutsi-Hima Bantus, Maasai Nilotes and Ful West Africans — his “pure” Elongated Africans — do not have any extraneous physical influences that could account for their more Caucasoid-leaning morphology than other Bantus, Nilotes and West Africans, respectively. In reality, all of these populations have low-to-moderate levels of Afro-Asiatic admixture, which their ancestors acquired through interbreeding with early Cushitic and Berber groups. For this reason, such populations were often referred to in the anthropological literature as “Hamiticized Negroes”, or, if the Hamitic influence was believed to be a bit more salient, as “Negro-Hamites” or “Half-Hamites”. Sonia Cole in her masterwork The Prehistory of East Africa describes the situation thusly:
The Upper Kenya Capsian people were very tall, over 5 feet 10 inches, and had long, narrow skulls with prominent chins and noses. They are apparently of Caucasoid or Mediterranean type, and may be termed ‘Proto-Hamites’ to emphasize their resemblance to the present inhabitants of North Africa and the Horn (the term ‘Hamite’, often used to describe North and North-East African people with marked racial characteristics, should strictly speaking apply to a language group only). Possibly a mixing of this proto-Hamitic type with incoming Negroes (of which there is no sign in Africa during the Upper Paleolithic) gave rise to some of the modern semi-Hamitic negroid peoples, such as the Masai.
Hima Bantus (sitting) and Bairu Bantus (standing). Like their Tutsi Bantu relatives further west in the Great Lakes region, the Hima of Uganda share a close physical and genetic resemblance with the Bantu peoples dwelling near them, the Bairu.
Tutsi Bantu men styling their natural afro-textured hair in the traditional amasunzu coiffure.
Hiernaux, on some level, appears to be aware of this since he mentions various Afro-Asiatic-speaking populations on the continent as “possible sources of ‘Hamiticization’.” He also remarks that the Maasai and Tutsi “have some characteristically Cushitic food habits.” Additionally, in reference to the linguist Joseph Greenberg and the anthropologist George P. Murdock, Hiernaux notes that “the pastoral Masai[…] their language belongs to the Eastern Sudanic class in Greenberg’s classification. However, both their language and culture show a strong Cushitic influence, which makes Murdock describe them as ‘Kushitized Nilotes’.”
DNA
In the case of the Tutsi and Maasai, a notable Cushitic male influence is indeed evident in their uniparental lineages. Trombetta et al. (2015) observed that around 78% of Tutsis in Burundi and 76% of Maasai in Kenya carry typical Bantu and Nilotic clades. However, a significant minority are M293 bearers (Tutsi=22%; Maasai=24%). M293 is a subclade of E1b1b (E3b), a paternal haplogroup that is most common among Afro-Asiatic speakers in the Horn and North Africa. The M293 sublineage is specifically associated with early Southern Cushites, for it peaks among remnant Southern Cushitic speakers in the Great Lakes and its modern geographical distribution also closely mirrors the historical distribution of the Southern Cushitic languages. More importantly, the earliest occurrences of M293 have been identified in ancient skeletons belonging to the Pastoral Neolithic, a Cushitic-affiliated cultural complex (see below). Thus, we now have genetic confirmation that the subhaplogroup was indeed introduced to the lacustrine region by the Southern Cushites.
Although the South Cushitic-linked E1b1b-M293 clade has been detected at low-to-moderate frequencies among Tutsis and Maasai, more extensive Y-DNA analysis has unveiled that the majority of individuals from these communities actually belong to the E1b1a or E3a haplogroup. E1b1a is today the most common paternal lineage among Bantu speakers. Luis et al. (2004) and Luis et al. (2007) report that almost all Tutsis in Rwanda bear this clade (over 75%). This suggests that the Cushitic admixture existing among the Tutsi Bantus is more pronounced in Burundi than in Rwanda. Furthermore, Wood et al. (2005) note a high frequency (50%) of the typical Afro-Asiatic-affiliated haplogroup E1b1b in a sample of Maasai individuals in Kenya. Trombetta et al. (2015), however, observed a lower percentage (37.8%) of all E1b1b subclades in their Maasai sample from the same territory (cf. Supplementary Table 7). Hirbo (2011) reports an even lower E1b1b or E3b percentage of just 16.7% for his Kenyan Maasai cohort. Likewise, Mwema (2011) found that the most common patrilineal STR markers carried by Maasai in Tanzania (51 individuals, the largest Y-DNA sampling of Maasai) are instead assigned to the E1b1a clade. Hence, the Cushitic admixture present among the Maasai Nilotes appears to be focalized in only select areas in Kenya; the “purer” Maasai population in Tanzania and elsewhere in the Great Lakes region shows a considerably lower Afro-Asiatic influence (as also indicated by autosomal DNA analysis; see below).
Y-DNA haplogroups of the Tutsi Bantus, Hutu Bantus and other Sub-Saharan African populations. Most Tutsi individuals (80%) and Hutu individuals (73%) carry derivatives of the E3a or E1b1a lineage like other Niger-Congo-speaking groups, which mainly consist of the basal E3a* clade and the E3a7 subclade (Hirbo (2011)).
Similarly, Hassan et al. 2008 found that around 54% of Ful/Fulbe/Fulani migrants in Sudan are haplogroup R1 carriers. This clade is today the most common paternal haplogroup among males in Europe. The presence in Africa of its R1b-V88 sublineage (which now is mainly borne by Fulani and Chadic speakers) is believed to represent late-glacial period diffusion from the Franco-Cantabrian area of southwestern Europe. This is supported by genome analysis of modern Fulani individuals, who carry a bit of Anatolian Neolithic-related ancestry like that characteristic of Iberian peoples (see Vahaduo Multi analysis here; the Fulani individuals on average bear ~7.6% Anatolian Neolithic admixture, as exemplified by the TUR_Marmara_Barcin_N sample). A further 35% of Ful belong to M78, a subclade of haplogroup E1b1b/E3b that is thought to have originated in Egypt. Hence, contrary to what Hiernaux claims, there is not one but at least two separate sources of exotic influence among the Ful: an ancient Iberian one and a Berber one.
Unlike their Y-DNA/paternal DNA, the mtDNA/maternal DNA of the Tutsi, Maasai and Ful appears to show little exotic influences (cf. Cerný et al. (2006), Castri et al. (2008)). This suggests that the Afro-Asiatic admixture in these populations was instead mainly acquired through the past assimilation of Cushitic and Berber males. In regards to the Tutsi, Göbel et al. (2019) found that almost all individuals in their Rwanda sample belonged 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:
The Rwandan sample[‘s] […] broader haplogroup pattern, mirrored also by higher MNPD (Table 1), is[…] highly similar to that reported from Kenya.[…] Most of the non-L low-frequency lineages found in the novel datasets are highly informative about human history. Lineages M1(a) and U6(a) are explained as signals of ancient backflow into North Africa from the Mediterranean area in the Early Upper Paleolithic. N1a1a is a low-frequency lineage with a relict distribution likely indicating a Pleistocene dispersal from Arabia. All are reported also in other East and North African populations. The single haplotype of K1a, a lineage found across West Eurasia according to EMPOP, might be attributable to more recent migration to Rwanda and is, intriguingly, shared with the dataset from Somalia.
This stands in sharp distinction to the neighboring Iraqw of Tanzania, a relict South Cushitic-speaking population that has managed to retain significant frequencies of Afro-Asiatic-associated uniparental markers (both Y-DNA and mtDNA) despite being completely surrounded by hunter-gatherer/Bantu/Nilotic communities. In this respect, Hirbo (2011) reports that 55.3% of his Iraqw sample bore the E1b1b paternal clade (of which 51% was assigned to the M293 or E3b6 subclade), with 12.8% carrying the T or K2 lineage. The examined Iraqw individuals also had a moderate incidence of West Eurasian maternal haplogroups (~22%), much of which consisted of the M1a clade (8.7%).
Analysis of HLA antigens among populations in Africa, the Middle East and Europe. Ethiopians, Beja and Sudanese “Arabs” cluster with other Afro-Asiatic-speaking groups inhabiting North Africa and the Middle East, as do Nubians. By contrast, Nilo-Saharan and Niger-Congo-speaking populations cluster together, separately from the Afro-Asiatic speakers (Aamer (2016)).
With regard to HLA antigens, Tang et al. (2000) write:
Cumulative anthropologic and genetic evidence indicates that centuries of intermarriage have largely dissolved the ethnic distinctions originally separating tribes (e.g. Hutu and Tutsi subgroups).
As such, the scientists observed that their Rwandan female sample’s HLA profile was generally unique to it, though broad commonalities could be discerned with certain other populations in Africa. Of these, the Rwandan cohort’s DRB1 and DQB1 haplotypes were again most similar to those of the Kenyan samples, but rather different from those of the West and Central African samples. This finding concurs with the Y-DNA haplogroup data on the Tutsis, for it suggests that they and the Hutus are most closely related to other Bantu populations in the Great Lakes region (e.g. the Kikuyu Bantus, who experienced comparable hunter-gatherer, Nilotic, and Cushitic admixtures as the Tutsi Bantus). Ali et al. (2020) likewise indicate that the Rwanda population’s HLA profile is analogous to those from other territories in eastern central Africa (viz. Kongo Kinshasa, Central African Republic, Uganda). By contrast, Lulli et al. (2009) report on the Fulani/Fulbe that: “In particular, the DRB1*04 allele is absent or rare in all Sub-Saharan African populations, except in the Fulani and in Amhara-Oromo from Ethiopia, where it reaches a frequency close to that of Europeans. These observations are in agreement with the hypothesis that the Fulani’s genetic make-up includes an appreciable Caucasoid component of possible East-African origin, which has been suggested on the basis of their physical features and cultural traditions.” This is also supported by Mohamoud (2006), who notes that “the result of HLA class I and class II antigen frequencies show that the Somali population appear more similar to Arab or Caucasoid than to African populations.”
DNA analysis of the long-horned “Ankole” cattle kept by Tutsi-Hima herders indicates that they are a variety of “Sanga,” the most common cattle breed owned today by Niger-Congo and Nilo-Saharan-speaking pastoralists in Africa. Sanga cattle is a crossbreed of the “Hamitic Longhorn” or taurine cattle of ancient Egypt (Bos taurus) and the zebu cattle of the Indian peninsula (Bos indicus). The Hamitic Longhorn, now almost extinct, has long since been replaced by the zebu and the camel among Afro-Asiatic speakers in Northeast Africa and by different varieties of the hardier Sanga cattle among Niger-Congo/Nilo-Saharan speakers elsewhere. The lone exception is among Fulani pastoralists of the Sahel, who keep herds of N’Dama cattle. N’Dama is the only bovine variety in Africa today that still has predominant taurine ancestry, consistent with the Fulani’s suggested links to the ancient Nile Valley (Kwon et al. (2022); Tijjani et al. (2022)).
Further evidence of exotic influence among the Maasai and Ful can be seen in the lactase persistence alleles that they carry. Tishkoff et al. (2007) observed that up to 58% of the Maasai bear the C-14010 variant. According to Breton et al. (2014), this lactose tolerance allele originated among Afro-Asiatic-speaking pastoralists in East Africa (likely Southern Cushites), who then spread it in the surrounding area all the way to the Khoe-inhabited parts of Southern Africa. This postulated diffusion is now supported by ancient DNA analysis (see below), which has identified the C-14010 variant in an early Cushitic specimen associated with the Pastoral Neolithic. This points to a rapid expansion of lactose tolerance in East Africa, just as has been observed in Europe. Similarly, Lokki et al. (2011) found that around 37% of Fulani in Mali carry the T-13910 mutation, the most common lactase persistence allele among Europeans. DNA analysis of the long-horned cattle raised by Fulani pastoralists, moreover, indicates that the “N’Dama” breed they keep is largely the same “Hamitic Longhorn” or taurine variety (Bos taurus) of ancient Egypt (cf. Kwon et al. (2022); Tijjani et al. (2022)). Altogether, this is a reflection of the aforementioned gene flow from the Franco-Cantabrian area of southwestern Europe, which impacted the ancestral Ful.
Worldwide allele frequencies of the TAS2R38 gene, which is associated with bitter taste perception. Most of the Somali and Ethiopian Jew individuals as well as the other Afro-Asiatic-speaking samples from the Middle East carry the global PA variant, followed by the Europe-centered AV allele. By contrast, most of the Maasai Nilotes bear the global PA variant, followed by the Africa-centered AA allele (Mourali-Chebil et al. (2022)).
As regards bitter taste perception, which is genetically inherited, analytical data indicates that Maasai individuals share similar allelic variants as other Nilo-Saharan and Niger-Congo speakers. Mourali-Chebil et al. (2022) observed that most of the Maasai Nilotes they examined bore the global PA variant of the TAS2R38 gene, followed by the Africa-specific AA allele. By contrast, after the generic PA variant, most of the Somalis, Ethiopian Jews and other Afro-Asiatic-speaking samples from the Middle East were found to carry the Europe-centered AV allele, with few individuals bearing the AA variant.
Global frequency distribution of the various isoforms of the APOE gene. After the globally-distributed APOE3 allele (ε3), most Tutsi Bantus and other native populations of central Africa bear the Africa-centered APOE4 or ε4 variant (nearly 40%). In contrast, Afro-Asiatic speakers from the Horn of Africa and Arabian peninsula that have been examined almost all carry the APOE3 isoform, which climaxes among Amerindian groups as well as Basques and Sardinians in southern Europe (Abondio (2021)).
Concerning variability in the apolipoprotein E (APOE) gene, which influences longevity in humans, Abondio (2021) reports that, after the APOE3 isoform (ε3) which prevails in all contemporary populations, her Tutsi Bantu cohort, Aka Pygmies and other native samples from central Africa carried among the highest worldwide frequencies (close to 40%) of the Africa-centered APOE4 isoform (ε4). By contrast, the Afro-Asiatic-speaking individuals from Ethiopia and the Arabian peninsula whom she studied almost exclusively bore the APOE3 variant (>85%), which peaks among Sardinians and Basques of southern Europe as well as among Amerindian populations. Abondio indicates:
ε3 is currently the most frequent isoform in all modern populations, peaking at 94% in the Hutterite people of Canada, 90% in Mayas, 88% in the Basque and Sardinian populations, and 86% in Han Chinese people. Isoform ε4 shows the maximum percentage in a range of indigenous populations of Central Africa (40% in Aka Pygmies, 38% in Tutsis), Oceania (49% in the Hui population of New Guinea) and Central America (27% for the Huychol people of Mexico). Moreover, a decreasing North-South gradient (Figure 3.5.1 and Appendix A1) characterizes this isoform in Europe (5% to 10% in the Mediterranean basin; 16% in Central and Western Europe; 23% in the Scandinavian peninsula, with peaks of 31% in Finland) and China (Zekraoui et al. 1997; Corbo and Scacchi, 1999; Singh et al. 2006; Hu et al. 2011; Ojeda-Granados et al. 2017). Isoform ε2 has an inconsistent distribution, with peaks in Southeast Asia, Australia, Africa (19%) but is absent Native American groups (Singh et al. 2006).
Density map of sickle cell in Africa and the Middle East (Ridley (2003)). Most modern Niger-Congo-speaking populations carry sickle cell alleles at variable frequencies, including the Tutsi Bantus and their Hutu brethren in the Great Lakes region (estimated at 1%-5% and 5%-15%, respectively; cf. Bain (2006)). Although they today inhabit the malarial zone, the Afro-Asiatic-speaking populations in Northeast Africa generally do not bear any sickle cell variants. Sickle cell mutations are, however, present among some individuals in Northwest Africa and the Middle East, primarily in the form of the Arab-Indian haplotype (cf. Al-Ali et al. (2020)).
With respect to sickle cell anemia, Bain (2006) reports that 1%-5% of Rwandan Tutsi, 5%-15% of Rwandan Hutu, and 1.5%-26% of Burundi inhabitants have the associated Haemoglobin S (HbS). This trait is generally not found among the Cushitic-speaking populations of the Horn of Africa (Bain notes a ~0% HbS frequency in Djibouti and Somalia), but it does occur among a significant minority of modern Bantus and other Niger-Congo-speaking groups. Prendergast et al. (2018) further indicate that their early Bantu sample from the Deloraine Farm in Kenya’s Rift Valley (dated to c. 1170-970 years before present) lacks any sickle cell alleles (cf. Table S12). This suggests that the Tutsi Bantus and Hutu Bantus did not derive their sickle cell variants directly from the Iron Age Bantus, but instead from the same, more recent source as other contemporary Bantu speakers in the Great Lakes region.
In terms of the GM/KM immunoglobulin allotype system, Weber et al. (2000) analysed Tutsi, Hutu and Twa individuals from Rwanda/Burundi and found that “the Tutsi and Bantu show close genetic associations.” Coudray et al. (2004) likewise observed that their Fulani sample from Senegal clustered with other Niger-Congo-speaking populations. On the other hand, their Cushitic-speaking (Issa Somalis of Djibouti) and Ethiosemitic-speaking (Amhara and Tigray of Ethiopia) samples from the Horn of Africa showed affinities with their Berber and Arabic-speaking samples from North Africa. Coudray et al. suggest that this is because these Afro-Asiatic-speaking populations share European GM haplotypes:
L’axe 1 de cette deuxième AFC permet de mettre en évidence 2 grands groupes de populations. Un groupe comprenant les populations nord et est africaines : Berbères du Maroc, d’Algérie, de Tunisie ; Touaregs d’Algérie et populations est africaines d’Ethiopie et de Djibouti. Cet ensemble se divise en trois sous-groupes selon l’axe 2 : Touaregs d’Algérie, Berbères et Amhara Tigrai et Issas d’Afrique de l’Est. L’autre grand groupe rassemble les populations sub-sahariennes (Mali, Sénégal, Côte d’Ivoire, Nigeria et République Centrafrique). La position de ces groupes aux extrémités de l’axe 1 révèle l’importante distance génétique les séparant. Cette distance se traduit par une différenciation entre les haplotypes Gm « européens » (Gm21,28;1,17;.., Gm21,28;1,2,17;.., Gm5*;3;23 et Gm5*;3;..) pour le groupe nord et est africain et les haplotypes Gm « africains » (Gm5*,28;1,17;.., Gm5*;1,17, Gm5,6,11,24;1,17;.., Gm15;1,17;.., Gm5,6,10,11,14,28;1,17;.. et Gm5,6,10,11,14;1,17;..) pour les populations sub-sahariennes. Nous pouvons aussi remarquer que cette distinction de 2 ensembles de populations africaines se superpose à une distinction géographique de part et d’autre du Sahara. Le désert ne représente pas pour autant une barrière aux gènes car on trouve chez les Berbères nord-africains environ 20 % d’haplotypes « sub-sahariens ».
Axis 1 of this second AFC makes it possible to highlight 2 large groups of populations. A group comprising the North and East African populations: Berbers from Morocco, Algeria, Tunisia; Tuaregs from Algeria and East African populations from Ethiopia and Djibouti. This set is divided into three sub-groups along axis 2: Tuaregs from Algeria, Berbers and Amhara Tigrai and Issas from East Africa. The other large group brings together the sub-Saharan populations (Mali, Senegal, Ivory Coast, Nigeria and the Central African Republic). The position of these groups at the ends of axis 1 reveals the great genetic distance between them. This distance results in a differentiation between the “European” Gm haplotypes (Gm21,28; 1,17; .., Gm21,28; 1,2,17; .., Gm5 *; 3; 23 and Gm5 *; 3 ; ..) for the North and East African group and the “African” Gm haplotypes (Gm5 *, 28; 1.17; .., Gm5 *; 1.17, Gm5,6,11,24; 1.17; .., Gm15; 1,17; .., Gm5,6,10,11,14,28; 1,17; .. and Gm5,6,10,11,14; 1,17; ..) for the sub-Saharan populations. We can also notice that this distinction of 2 sets of African populations is superimposed on a geographical distinction on either side of the Sahara. However, the desert does not represent a barrier to genes because we find among the North African Berbers about 20% of “sub-Saharan” haplotypes.
— Global autosomal STR affinities. The Tutsi Bantus again cluster with the Hutu Bantus and other Niger-Congo-speaking individuals. By contrast, the northern Sudanese (SUD) group with Afro-Asiatic-speaking Egyptian (EGY) and Omani (OMN) individuals (From top left, clockwise: Coudray et al. (2006); Simms et al. (2008); Shepard and Herrera (2006b); Shepard and Herrera (2006a)).
In terms of autosomal DNA — which is inherited from both parents (and thus often used to gauge overall ancestry), and studied by analyzing single nucleotide polymorphisms (SNPs) and autosomal short tandem repeats (autosomal STRs or microsatellites) — researchers have again noted a moderate Afro-Asiatic influence among the Maasai and Ful. Autosomal SNP data suggests that this genetic introgression was derived from early Cushitic and Berber peoples, who were absorbed by the Maasai’s Nilotic and the Ful’s West African ancestors, respectively (cf. Dobon et al. (2015); Henn et al. (2012)). Cherni et al. (2016) and Hodgson et al. (2014) estimate that the Maasai of Tanzania and Kenya (Kinyawa), respectively, have on average 12% and 21% of ancient Cushitic admixture, while the Fulani have around 28% of ancient Berber admixture (Table S6).
Genome analysis of contemporary Fulani individuals and various ancient and modern global samples. At K=6, the analysed Fulani individuals form two distinct genetic clusters: a FulaniA group that is distinguished by higher non-Sub-Saharan African admixture (consisting of an Iberomaurusian-related orange component, a Pre-Pottery Neolithic-related azure component, and a Balkan European-related red component), and a FulaniB group that mostly harbors Niger-Congo/Nilo-Saharan-related ancestry (yellow component) like the neighboring Wolof, Mande and other Niger-Congo/Nilo-Saharan speakers. The Berber samples appear to represent the source population that contributed most of the non-Sub-Saharan African ancestry borne by the Fulani, as these Afro-Asiatic speakers carry all of the aforementioned ancestral elements. Additionally, the Berber individuals harbor some extra Iran Neolithic-related admixture (dark blue component). They seem to have acquired this genome element after the contact period between their Berber ancestors and the ancestral Fulani because the contemporary Fulani samples do not bear this component. Furthermore, although the Afro-Asiatic speakers from the Horn of Africa share most of the same ancestry as the Berber samples, these Cushitic and Ethiosemitic-speaking individuals are differentiated by minor Late Stone Age (c. 3000-1000 BCE) Great Lakes hunter-gatherer admixture (light blue component); they also do not show any trace of the Balkan European red element. Overall, this supports the traditional interpretation that most of the Fulani’s non-Sub-Saharan African admixture was derived from a Berber-affiliated source population(s) (D’Atanasio et al. (2023)).
Genome analysis on the Vahaduo Admixture JS program’s Single function. Fulanis from Ziniare and peninsular Arab bedouins can be modeled as source populations for Mozabite Berbers and other Maghrebi individuals. The ancestral composition of the Mozabite Berbers here consists of around 75% Fulani-derived ancestry and 25% peninsular Arab Bedouin-derived ancestry. However, this estimate is inaccurate since the modern Fulani Ziniare are not a “pure” West African source population. The average Fulani Ziniare individual bears about 25% Eurasian ancestry, which was specifically acquired through admixture with Maghrebis (similar to the percentage of Cushitic-derived Eurasian admixture, which is now carried by the Maasai Nilotes in eastern Africa). What this Vahaduo Single analysis does establish is that, compared to other West African populations, there is considerable gene sharing between the Fulani Ziniare and Maghrebis. This is consistent with Maghrebi gene flow into the ancestral Fulani population.
Genome analysis of Fulani individuals from Cameroon and Sudan, as well as other modern samples from Africa (Bird et al. (2023)). Both examined Fulani groups almost exclusively derive their Eurasian ancestry from a Morocco Berber-related source (dark blue component). They completely lack the signal of ancient Chadic ancestry, which consists of a composite Eurasian ancestry of Cushitic and Tunisian Maghrebi origin (pink and light blue components). By contrast, this genetic signature of ancient Chadic heritage is found among contemporary Chadic speakers in northern Cameroon, among Sahelian groups like the Kanuri and Kotoko, and reaches a frequency peak among the Shuwa/Baggara “Arabs” of Cameroon (who likely originally spoke an Afro-Asiatic language of the Chadic branch). It is interesting to note that, despite inhabiting western Africa, the Shuwa “Arabs” and Cameroon Sahelian samples also bear Nilo-Saharan admixture (purple component). This supports the suggestion that the Shuwa migrated to their present location from the eastern Sahara and Nile Valley, where their Afro-Asiatic-speaking ancestors would have interacted with local Nilo-Saharan speakers before spreading westward. All in all, the study concurs with our Vahaduo genome analysis above, which detected a clear affinity between the Fulani samples from Ziniare, Burkina Faso and Berber individuals. We can now confirm that this association is indeed primarily attributable to ancient Berber gene flow into the Fulani population.
Published autosomal STR analysis on the Tutsi indicates that they are biologically related to the Hutus and other Bantu populations, with little exogenous affiliations (Simms et al. (2008); Simms et al. (2010); Shepard and Herrera (2006a); Shepard and Herrera (2006b); Coudray et al. (2006)). Although no such peer-reviewed autosomal SNP work on the Tutsi-Hima has yet been published, available data on their Hema congeners from the Democratic Republic of the Congo points to a similar Cushitic admixture as found among the Maasai Nilotes (~20%; cf. Xing et al. (2010)). Similarly, autosomal SNP analysis conducted by 23andme, AncestryDNA and other genetic testing companies has traced most of these Tutsi individuals’ ancestry to Niger-Congo/Nilo-Saharan-related forebears (~70% on average); the remainder consists of Cushitic admixture (~30%), which was derived from earlier Afro-Asiatic speakers from Northeast Africa whom the Tutsis’ Bantu/Nilotic ancestors assimilated. These observed predominant Niger-Congo/Nilo-Saharan affinities are in line with a DNA test taken by the Rwandan leader Paul Kagame. According to the Financial Times, although various different admixture strains were detected, the Tutsi politician ultimately shared ancestral links with the neighboring Hutu, consistent with both groups’ spoken Bantu language:
Mr Kagame has taken a DNA test that, he says, reveals him to be of particularly complex genetic mix. The implication, he says, is that he, the ultimate symbol of Tutsi authority, has some Hutu in his genetic make-up.
This is in stark contrast to the Cushitic and Ethiosemitic speakers to the north in the Horn of Africa, who instead share close autosomal SNP (Hodgson et al. (2014), Supplementary Text S1; Dobon et al. (2015); Almarri et al. (2021), Table S4) and autosomal STR ties with other Afro-Asiatic-speaking communities in North Africa and the Middle East (cf. Steele et al. (2014), the largest global autosomal STR analysis).
(*N.B. Using the mAUTO formula, which Stefflova et al. (2009) developed to infer autosomal DNA from known mtDNA and Y-DNA haplogroup frequencies, we may surmise that the Tutsi Bantus of Rwanda bear under 5% total Eurasian ancestry. Since this estimate is around 15 percentage points lower than what is usually projected for them (see estimates below), it would appear that the uniparental lineage frequencies of the Rwandan Tutsi have been shaped by a founder effect. For further details on the mAUTO formula, see Can the Y-DNA and mtDNA of a population be used to infer its autosomal DNA?.)
Genome analysis of Northern Somalis, Oromos, Ethiopian Jews and other global populations, including Maasai. Most of the Northern Somali individuals, who were born in the northeastern Puntland region of Somalia (hailing from the Majerteen Darod clan), cluster in a position parallel to but separate from the North African/Arabian cluster. A few of the examined Northern Somalis are outliers, grouping instead with either the Ethiopian Jew or Oromo samples or shifting toward samples from South Asia. All of the Sub-Saharan African samples are distinct from the Afro-Asiatic-speaking populations from the Horn, North Africa and the Middle East. However, the Maasai Nilotes and Kikuyu Bantus do pull toward the Afro-Asiatic speakers, due to past contact between their Bantu/Nilotic ancestors and neighboring Cushitic groups (Osman and Jonasson (2022)).
Genetic analysis of the ancient Cushitic settlers of the Pastoral Neolithic indicates that “PN individuals, including Elmenteitan and those within the heterogenous SPN category (whom we refer to as “other PN”), mostly form a tight cluster near present-day Afro-Asiatic speakers” (Prendergast et al. (2018)). More detailed genetic modeling of these early Cushites of the Pastoral Neolithic using Global25 further indicates that they bore a predominant non-African ancestry averaging 70% (see Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa).
In 2018, Prendergast et al. analyzed the DNA of various ancient skeletons in East Africa. The researchers found that the Maasai and other Nilotes in their sample set genetically clustered with the Pastoral Iron Age (PIA) specimens, whereas the Afro-Asiatic-speaking populations grouped with the Pastoral Neolithic samples (PN; comprising the Savanna Pastoral Neolithic and Elmenteitan cultures). They further observed that the Pastoral Iron Age individuals were characterized by ancestry related to modern Nilotic speakers in Sudan, while the earliest Pastoral Neolithic specimens derived most of their ancestry from northeastern Africa/Levant. Additionally, the scientists noted some gene flow from the PN specimens into the later PIA and Maasai samples, consistent with the documented Cushitic influence on the language and culture of the Maasai Nilotes. The researchers therefore suggest that the Pastoral Iron Age individuals were ancestral to the Maasai and other present-day Nilo-Saharan populations in the Great Lakes, and that the Pastoral Neolithic specimens instead represent the forebears of the Afro-Asiatic speakers:
PN individuals, including Elmenteitan and those within the heterogenous SPN category (whom we refer to as “other PN”), mostly form a tight cluster near present-day Afro-Asiatic speakers, with a small number of modest outliers, including the two individuals buried at Prettejohn’s Gully, whose ear-lier date (~4000 BP) coincides with the initial limited spread of herding into the area. Finally, five Iron Age individuals are shifted to the left in the PCA: four PIA individuals toward Nilotic speakers, and an IA child from Deloraine Farm (I8802)—the earliest agricultural site in Kenya’s Rift Valley (32)—toward western Africans and Bantu speakers.[…]
four PIA individuals spanning an ~800-year period show greater affinity to present-day Nilotic speakers and are associated with an influx of Sudan (Dinka)-related ancestry. Similarities between archaeologically and ethnographically documented material culture suggest that PIA sites may be associated with ancestors of present-day Kenyan Nilotic speakers such as the Kalenjin or Maasai (32, 47). Both the PIA individuals and present-day Maasai retain substantial components of PN-related ancestry, showing that the ancestry composition of PIA and more recent pastoralists reflects mixture with previously established herder groups in eastern Africa.
Wang et al. (2020) later reanalysed the ancient Cushitic pastoralists. Thanks to their improved and more broad-based dataset, including several new Pastoral Neolithic samples from Kenya, the researchers were able to more accurately determine the settlers’ ancestral composition. Like Prendergast et al., they formulated a three-way admixture model using Levantine Chalcolithic, Dinka and Mota specimens as hypothetical ancestral populations to the ancient Cushitic herders. However, admixture analysis confirmed instead that the early Cushitic pastoralists had almost identical ancestry proportions as modern Afro-Asiatic-speaking North African groups: they derived around 80% of their genome from a West Eurasian ancestral population, and had just 5%-10% of supposed admixture from Nilo-Saharan peoples. Thus, the ancient Cushitic pastoralists appear to have belonged to a transplanted North African population(s), which gradually moved southwards and settled in East Africa (cf. Supplementary Material).
Wang et al. (2022) reported that a Kerma period individual dating from the Middle Kingdom (c. 4000 year old), who was excavated at the Kadruka site in Upper Nubia (northern Sudan), has a genome that “is genetically indistinguishable from that of early Neolithic eastern African pastoralists located 2500 kms away.” The scientists note further that “our findings are consistent with established models for the southward dispersal of Middle Nile Valley pastoral populations to the Rift Valley of eastern Africa, and provide a possible genetic source population for this dispersal.” Hence, it now has been confirmed that the Cushites of the Pastoral Neolithic indeed belonged to a transplanted ancient North African population, and one with a predominant West Eurasian ancestry at that. (For more details, see Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa.)
To find out for ourselves what exactly are the autosomal DNA affinities of the Tutsi-Hima and their Hutu kinsmen, we conducted our own genome analysis of Tutsi, Hima and Hutu individuals using the Vahaduo Admixture JS program. Our Target samples mostly consisted of 23andme and AncestryDNA customers, who posted online their converted and scaled Global25 coordinates. For our Source proxies, we used all of the ancient individuals listed on Eurogenes‘ official Global25_PCA_scaled datasheet. Our results showed that, in terms of proximal ancestry (i.e. recent ancestry), the Tutsi and Hima largely descend from the Cushitic-admixed Nilotes of the Pastoral Iron Age or PIA. Prendergast et al. (2018) analysed the remains of PIA specimens and note that these ancient “PIA individuals spanning an ~800-year period show greater affinity to present-day Nilotic speakers and are associated with an influx of Sudan (Dinka)-related ancestry[…] PIA sites may be associated with ancestors of present-day Kenyan Nilotic speakers such as the Kalenjin or Maasai.” Here are the results for a typical Tutsi individual from Rwanda (see here for the full proximal analysis results for all of the Tutsi-Hima and Hutu individuals in the datasheet):
To determine the overall ancestral composition of the Tutsi, Hima and Hutu populations, we then performed another Vahaduo Single analysis, this time using as our Source proxies the “purest” Sub-Saharan African samples (i.e. least Eurasian-admixed) and the “purest” Eurasian samples (i.e. least Sub-Saharan African-admixed) available on the Global25_PCA_scaled datasheet (the methodology followed is described here). Our results showed that, in terms of distal ancestry (i.e. remote ancestry), the Tutsi, Hima and Hutu each traced most of their ancestry to the Kakapel_300BP specimen, which is the “purest” known ancient Nilo-Saharan sample. Here are the results for a typical Tutsi individual from Rwanda (see here for the full distal analysis results for all of the Tutsi-Hima and Hutu individuals in the datasheet):
(*N.B. The Vahaduo Single analyses above of the Tutsi, Hima and Hutu individuals should be viewed with discretion since, unlike our Vahaduo Single analyses of the southern Somali individuals (which are based on unscaled or raw Global25 coordinates), these analyses are predicated upon scaled Global25 coordinates. Scaled coordinates are less accurate, filtered versions of unscaled/raw coordinates. As such, they capture less of the genetic variation that an individual carries. We have, nonetheless, made use of these scaled coordinates in our analyses because currently they are the only coordinates of Tutsi-Hima and Hutu individuals that are publicly available. If/when unscaled or raw Global25 coordinates are ever made accessible, we shall update our analysis to reflect this real, unfiltered output.)
The Vahaduo Single analyses above of Tutsi and Hima individuals are also supported by genetic distance analysis. When we carry out a Vahaduo Distance investigation with a focus on proximal or near ancestry, these Tutsi-Hima samples all have a closest genetic distance to the Cushitic-admixed Nilotes of the Pastoral Iron Age. This is in sharp contrast to the South Cushitic-speaking Iraqw, who, despite having sustained considerable hunter-gatherer/Nilo-Saharan/Niger-Congo admixture, are genetically nearest to the Cushites of the Pastoral Neolithic. This confirms our earlier observation that the Iraqw are Cushitic peoples who experienced significant hunter-gatherer/Nilo-Saharan/Niger-Congo admixture, whereas, oppositely, the Tutsi-Hima are Nilo-Saharan/Niger-Congo peoples who experienced some moderate Cushitic admixture.
Vahaduo Distance analysis of South Somali and Kenyan Somali individuals. All of these Cushitic speakers show closest genetic distance to other Afro-Asiatic-speaking individuals from the Horn vicinity (just 1%-3% genetic differentiation), including samples of other Cushitic (Oromo, Afar, Agaw, Rendille, Iraqw), Ethiosemitic (Amhara, Tigray, Ethiopian Jew), and North Omotic (Wolayta) speakers. Unlike the Iraqw, the Tutsi-Hima do not cluster with the Horn region’s Afro-Asiatic speakers, nor do the Maasai or any other Nilo-Saharan/Niger-Congo/Khoisan populations. This highlights the fact that the Tutsi-Hima are peoples of Bantu/Nilotic/hunter-gatherer ancestral origin who received moderate Cushitic admixture, whereas, oppositely, the Iraqw are people of Cushitic ancestral origin who received Bantu/Nilotic/hunter-gatherer admixture.
In order to better quantify and describe our findings, we ended by running an analysis using the Vahaduo program’s Multi function. Our examined Tutsi and Hima individuals all wound up bearing a predominant Sub-Saharan African ancestry averaging 72%, a majority of which belongs to the “pure” ancient Nilo-Saharan Kakapel_300BP component (~64%) and a minority of which is derived from indigenous East African hunter-gatherers, as represented by the “pure” forager cohort MWI_Chencherere (~8%). The rest of the Tutsi and Hima individuals’ ancestral makeup comprises Eurasian admixture (~22%, almost entirely consisting of Levantine Natufian-associated gene flow) and North African Iberomaurusian/Taforalt admixture (almost 6% on average), with no trace of ancient Niger-Congo-associated admixture (0% of the COG_Kindoki230BP component). By contrast, we detected around 31% of the COG_Kindoki230BP element in our Hutu samples but zero percent of the Taforalt component. This is in marked distinction to the Vahaduo Single proximal analysis above, which did identify a Bantu affiliation of about 10% in most of the examined Tutsi and Hima samples. Since Vahaduo Single analyses capture the exact ancestral makeup of an individual(s) and Vahaduo Multi analyses present that same data in a streamlined and simplified format, we may conclude that persons of Tutsi-Hima background do, in fact, carry some Bantu ancestry, albeit considerably less than what usually has been projected for them.
In brief, genome analysis indicates that the Bantu-speaking Tutsi, Hima and Hutu groups of the Great Lakes region are all closely related peoples. As such, the scholar Mahmood Mamdani (2020)‘s research — which asserts that the dichotomy between the Tutsi and Hutu is primarily one of social caste rather than of different ancestral origins — appears to be correct. On a broader scale, peer-reviewed autosomal STR studies as well as commercial genetic testing link the Tutsi-Hima and Hutu populations with other Niger-Congo-speaking communities inhabiting western Africa. However, our own independent genome analysis has found that, like their Nilo-Saharan-speaking Hema relatives in the Democratic Republic of the Congo, the Tutsi-Hima and Hutu actually descend from a common Nilo-Saharan-affiliated source population. In the archaeogenetic record, this ancestral community is best represented by the 300 BP sample from the Kakapel site in Kenya. The main difference between the Tutsi-Hima and the Hutu seems to be that the Tutsi-Hima had some later contact(s) with pastoralists from northern Africa (as evidenced by the presence among Tutsi-Hima individuals of both the Levantine Natufian and North African Iberomaurusian/Taforalt components) and received comparatively less gene flow from Bantu populations, whereas the Hutu had more intensified interactions with Niger-Congo-speaking groups (as evidenced by the greater frequency of the COG_Kindoki230BP component among Hutu individuals).
Our genome analysis above also seems to support the Kenyan historian Bethwell Ogot (1974)‘s assertion, in regards to the Tutsi-Hima, Maasai and other Bantu/Nilotic peoples that have some Cushitic admixture, that:
the lighter-skinned inhabitants of northern and north-eastern Africa should properly be classified ‘Caucasoid’, or, where necessary, ‘semi-Caucasoid’ or ‘Negroid-Caucasoid crosses’. But such descriptions would not extend to East African pastoralists such as Maasai, Karamojong, Turkana, Hima and Tusi[…] These peoples are decidedly Negroid, though some of them may have incorporated a small admixture of Caucasoid blood in their ancestry.
Physiognomy
With regard to skin pigmentation, Hiernaux contradicts himself again. He affirms above that the Tutsi are dark and specifies elsewhere in his book that they are “very dark skinned”. At one point, he goes as far as to suggest that:
In skin colour, the Tutsi are darker than the Hutu, in the reverse direction to that leading to the Caucasoids.
Yet, in his earlier Hamitic-centered work, Hiernaux maintains that the Tutsi have a lighter complexion than the Hutu. He further asserts that those inhabiting Rwanda are darker-skinned than their brethren in Burundi. This is quite telling since, per Luis et al. (2004), the “darker” Tutsi in Rwanda appear to have a significantly lower Cushitic male influence than Hiernaux’s “lighter” Tutsi in Burundi (they instead are almost all haplogroup E1b1a/E3a carriers, like the average Hutu and most other Bantus). As it is very unlikely that the Tutsi and Hutu incurred a diametric change in coloration within the span of a few years, it follows that Hiernaux’s sudden about-face was motivated by something other than the scientific data.
Global distribution of the derived Ala111Thr/rs1426654 allele, which is associated with lighter skin pigmentation. This SLC24A5 gene variant today has frequencies of around 60% or greater among the Afro-Asiatic-speaking populations in the Horn and North Africa.
Hiernaux makes another error here when he associates a priori dark skin pigmentation with non-Caucasoid ancestry. Ancient DNA analysis has, in fact, confirmed the opposite. That is, most Caucasoid populations — like all early humans — were until recently dark brown-skinned because the alleles linked with lighter coloration are only a few thousand years old. Various pre-Neolithic specimens from Europe with otherwise unexceptional genomes, including ancient individuals from Spain and Greece, were thus found to still have ancestral variants in several skin pigmentation genes. Consequently, even if the early Afro-Asiatic-speaking peoples that the forebears of the Tutsi Bantus, Maasai Nilotes and Ful West Africans encountered had been swarthy, that alone would not necessarily mean that they were not Caucasoids.
A Somali woman with grey eyes. One of the interesting discoveries of ancient DNA analysis on the Cushites of the Pastoral Neolithic is that some of them appear to have had light-colored eyes. This trait is still found today among a minority of their modern Cushitic-speaking descendants in Northeast Africa. (For more details, see I have read that some Cushitic peoples have blue eyes. Was this trait inherited from the ancient Cushites?.)
As it turns out, these ancient Afro-Asiatic speakers probably instead had light brown complexions. We know this from ancient DNA analysis of early Cushitic settlers in East Africa. Two Pastoral Neolithic individuals (a 2300 year old sample from Hyrax Hill and a 1500 year old sample from Molo Cave, both in Kenya) were tested for the SLC24A5 gene’s derived allele, Ala111Thr or rs1426654. This variant is associated with lighter skin pigmentation, and both specimens wound up carrying it (modern Europeans have this main allele and several additional minor ones; East Asians have their own separate variant). These ancient Cushitic herders also passed on the derived allele to their descendants in the Horn of Africa since the Afro-Asiatic speakers in that region today carry it at high frequencies (cf. Wang et al. (2020), Table S7). Furthermore, ancient DNA analysis of Cushitic individuals of the Pastoral Neolithic cultural complex has found that they bore the derived allele at the APBA2 locus (2 out of 2 examined specimens or 100%), which likewise confers lighter skin pigmentation. Some individuals also carried the derived rs1800404 variant at the OCA2/HERC2 locus, an allele associated with lighter eye color (2 out of 2 examined Pastoral Neolithic specimens from the Lukenya Hill site). These alleles are today still common among Cushitic-speaking groups. By contrast, the early Nilotes of the Pastoral Iron Age were found to bear the ancestral alleles at the SLC24A5, APBA2 and OCA2/HERC2 loci, which are associated with darker skin pigmentation and eye color (Wang et al. (2020), Table S7). (For further details, see What complexion were the ancient Cushitic settlers of eastern Africa? and I have read that some Cushitic peoples have blue eyes. Was this trait inherited from the ancient Cushites?.)
(*N.B. Lazaridis et al. (2022) conducted a comprehensive analysis of phenotypic traits borne by ancient individuals excavated in Europe and Asia. The scientists report that most of their Chalcolithic period sample from Israel (85.7%) had intermediate skin pigmentation, similar to their ancient Minoan cohort from Bronze Age Crete (80%) (cf. Supplementary Materials). According to Wang et al. (2020), this Chalcolithic Levant sample is the best-fitting proxy for the distal source of the West Eurasian ancestry which defines the Cushites of the Pastoral Neolithic. This suggests that these early Cushitic settlers indeed would have been relatively light-complected at the time of their initial arrival in eastern Africa.)
A Somali man (Hawiye clan), with the “Caucasoid” or European-like physiognomy typical of “pure” Cushitic peoples. Because of the close osteological affinities between skeletons belonging to the Pastoral Neolithic culture of eastern Africa and those of modern Afro-Asiatic speakers in the Horn region, anthropologists surmised that these peoples shared ancestral roots. This conjectured link has been confirmed by genomic analysis since contemporary Cushitic, Ethiosemitic and North Omotic-speaking individuals trace most of their proximal/recent ancestry to the Cushitic makers of the Pastoral Neolithic (see genetic analysis above).
In terms of hair form, Hiernaux is more consistent. He writes that “outside of the serological field, hair form appears to be the characteristic which shows the most sharply marked contrast between sub-Saharan Africa as a whole and surrounding areas[…] over most of the subcontinent, spiralled hair is the only category to be observed”. On this basis, he notes that the Tutsi and Maasai have spiralled/kinky hair.
Somali artist Hani UK (Isaaq clan), representing an individual of “pure” Cushitic stock. Note the marked difference in physiognomy between her and the “pure” Tutsi Ange Kagame (shown below), particularly in terms of hair form. This disparity is due to the fact that the Tutsi-Hima are ultimately of Bantu/Nilotic origin, with low-to-moderate Cushitic admixture.
By contrast, Hiernaux is well aware that this is not the characteristic hair form of the Afro-Asiatic-speaking populations in the Horn; especially the northern Cushitic groups like the Somali, Afar and Beja:
At the opposite end of the scale, the lowest frequencies of spiralled hair in sub-Saharan Africa have been observed in Ethiopia and Somalia, with a minimum in the Somali.
African-American singer Jaguar Wright, closely resembling average Tutsi-Hima Bantu women in terms of mesorrhine facial features, dark skin pigmentation, attenuated prognathism, and natural afro-textured hair. This likeness again underlines the fact that the Tutsi-Hima Bantus are of Niger-Congo/Nilo-Saharan origin like African Americans (~75%), with both groups also sharing a similar proportion of ancillary Eurasian admixture (~25%).
A Shuwa (Baggara) “Arab” woman. Genome analysis indicates that the Shuwa/Baggara of the Sahel carry a composite Eurasian ancestry, which consists of components of Cushitic and Berber origin. This composite Eurasian signature appears to be the signal of ancient Chadic ancestry since it is also borne at lower frequencies by Chadic speakers (cf. Bird et al. (2023)). As such, the Shuwa/Baggara seem to have originally spoken an Afro-Asiatic language of the Chadic branch. This explains why the Shuwa/Baggara are physically, genetically and culturally much closer to Afro-Asiatic speakers than are Sub-Saharan tribes like the Tutsi-Hima, despite the fact that the Tutsi-Hima’s Bantu ancestors received low-to-moderate gene flow from Cushitic peoples.
— Hutu and Tutsi Bantu individuals from Rwanda. To the layperson, the average Hutu and Tutsi are virtually indistinguishable, owing to their shared Bantu origins. They are, in fact, officially classified as the same Bantu ethnic group, known as Banyarwanda in Rwanda and Banyarundi in Burundi. However, some minor physical differences do exist between the two, mainly in terms of height and nasal index.
— Peoples of Nilotic origin. Top row, from left to right: Dinka man, Nuer man, Tutsi man, Maasai man. Bottom row, from left to right: Dinka woman, Nuer woman, Tutsi woman, Maasai woman. Genome analysis indicates that the Tutsi-Hima and the Maasai of the Great Lakes region trace most of their proximal or recent ancestry to the Nilotes of the Pastoral Iron Age. This shared Nilo-Saharan ancestry readily explains why the Tutsi-Hima, despite today speaking a Bantu language, most closely resemble the Maasai and other Nilotic peoples.
Because Hiernaux presumes that his ancient “Elongated African” stock — a monolithic entity which, as we saw in part one, simply does not exist in the archaeological record — had kinky hair, he believes that the non-kinky hair texture that is common today among the Afro-Asiatic-speaking populations in the Horn is an indicator of exotic genetic influence. He hints that it may have been introduced in antiquity by Arabs, specifically. In reality, this non-spiralled hair form was the characteristic texture of the ancestral Cushitic speakers themselves. It was thus simply passed down from them to their modern descendants. We know this because, within the Horn region itself, every hunter-gatherer, Bantu and Nilotic population that has had extensive contact with early Cushitic groups has at least some members that possess soft-textured hair as a direct result of that interaction. Puccioni (1931) thus reports that, among the Ribi foragers of Somalia (who have significant Cushitic admixture), 60% of the individuals he examined had frizzy hair while only 40% possessed woolly hair. We also know this because the ancient peoples of the Kerma civilization in Sudan, barring a few assimilated individuals of Nilotic origin, uniformly had non-kinky hair. According to linguistic analysis by Marianne Bechaus-Gerst (1989, 2000), these Kerma folks spoke languages from the Cushitic branch of the Afro-Asiatic family. The pioneering Prussian Egyptologist, linguist and archaeologist Karl Richard Lepsius had the opportunity to examine the Kerma remains, and notes the following:
Speaking not as a trained anatomist but as one who has had the characteristics of different Nile Valley races pointed out to him by Prof. Elliot Smith and has handled the material for years, I may venture to add that the bones are not those of Negroes. A few cases of prognathous skulls occur and even, in the case of a woman in grave Kerma, tightly curled black negro wool on the head; but most of the men, especially the principal skeletons, had fine heads with straight black hair.
(For more details, see Did the ancient Cushites have soft-textured hair?.)
Additionally, Hiernaux indicates that the ancient C-Group pastoralists of Lower Nubia entered the Nile Valley from the Sahara and also settled in the Horn. He bases this on similarities in material culture and rock art. For example, John Desmond Clark asserts that “cattle and caprids (meaning here sheep/goat) were introduced to the lowlands of the Horn and the fringes of the Ethiopian plateau by C-group pastoralists from the Nile Valley and the surrounding arid regions” (cf. Barnett (1999)). Arkell (1961), moreover, identified pottery and stone work in Agordat, Eritrea, akin to that produced by the C-Group peoples (Fage and Oliver (1970)). Reisner (1912) likewise states that the “black- and brown-polished wares generally with incised white-filled patterns, peculiar to the Early Predynastic, Early Dynastic and the C-group periods[…] this pottery may be made in imitation of ebony or gourd vessels. The C-group patterns are suggestive of modern Somali, Sudanese and Nubian basketwork patterns, especially in the coloured examples.” Similarly, Stuart Munro-Hay writes that “pottery studies have suggested that the pre-Aksumite culture might owe something to Nile valley influences, specifically to the C-group/Kerma cultures (see O’Connor, Ch. 34, this volume), and in rather later times to Meroe/Alodia, as well as to contacts across the Red Sea on the Yemeni and Saudi coasts” (cf. Andah et al. (2014)). Furthermore, rock art at Dhawaale in northwestern Somalia, Harurona in the Wolayta region of Ethiopia, Abka in Sudan, and other archaeological sites in Northeast Africa include cross-in-circle motifs (also known as the encircled cross or cross-filled circle) (Bachechi (2005)). Istituto per l’Oriente (1979) notes that these are common symbols in C-Group art, which still feature prominently in Egyptian Coptic magic.
Osteological affinities between the modern Afro-Asiatic-speaking groups of the Horn and the C-Group folks have also been established (see Bräuer (1980a) above; also Batrawi (1946)). According to Peter Behrens (1981) and Marianne Bechaus-Gerst (2000), the C-Group makers spoke languages from the Berber branch of the Afro-Asiatic family (this linguistic association is backed by the recent finding in Somalia of ancient Libyco-Berber/Tifinagh inscriptions, which were presumably left by early C-Group settlers). Like the coeval Kerma inhabitants, the C-Group folks were also found to have had non-kinky hair. In 1914, the Peabody Museum scholar Oric Bates led excavations in the Libyan/Western Desert, where he observed:
The so-called “C Group” cemeteries of Nubia, it was early recognized, were those of a distinctly non-Egyptian people. They extend in time from about the end of the Vlth Dynasty to the XVIIIth Dynasty, although the lower date is one to be stated with some reserve[…] Only by exception is the hair woolly or “peppercorn-like” ; as a rule it is straight or wavy.
With respect to lip form, Hiernaux asserts that the Tutsi Bantus actually tend to have thicker lips than the Hutu Bantus:
Lip thickness provides a similar case: on an average the lips of the Tutsi are thicker than those of the Hutu. In most cases, however, they are not everted as in many West Africans.
Finally, analysis of fingerprint distribution patterns around the world indicates that the Tutsi-Hima Bantu and Maasai Nilote inhabitants of the Great Lakes region share similar dermatoglyphic characteristics as other Niger-Congo, Nilo-Saharan and Khoisan-speaking groups. All of these populations have a low pattern index (PI) or pattern intensity index (PII) of under 12.00. Contrarily, the Afro-Asiatic-speaking populations of the Horn region have relatively high fingerprint pattern values of around 13.50 to 14.25, like other Afro-Asiatic speakers in North Africa and the Middle East (Rife (1953)). Of these groups, Yohannes and Bekele (2015) report that their Cushitic-speaking Oromo sample from Ethiopia has the highest pattern intensity index, valued at 15.01.
Global distribution of fingerprint (dermatoglyphic) pattern values. The Tutsi-Hima Bantus, Maasai Nilotes and other Niger-Congo, Nilo-Saharan and Khoisan-speaking populations are characterized by a low pattern index (PI) or pattern intensity index (PII) below 12.00. By contrast, the Afro-Asiatic-speaking populations in the Horn of Africa, North Africa and the Middle East have a relatively high pattern index of around 13.50 to 14.25 (Rife (1953)).
In summary, Hiernaux was gravely mistaken about the somatic traits of the ancient Caucasoids in East Africa. They were of a lighter complexion than he had envisioned, and they left behind a corresponding skin pigmentation allele to show for it. Since this mutation originated only a few thousand years ago in or near West Asia, these early Afro-Asiatic speakers could not have settled in East Africa earlier than the Neolithic. Likewise, genetic examination of associated human remains has established that these pastoralists did indeed ultimately arrive from North Africa and/or the Middle East. They also had non-kinky hair texture, quite unlike the spiraled hair that Hiernaux had envisaged for them. These vanished “Hamitic” peoples left a low-to-moderate imprint on various early Bantu and Nilotic groups that they encountered in the Great Lakes region. However, their actual Afro-Asiatic-speaking relatives can still be found to the north in the Horn.
Anthropometry
The core of Hiernaux’s theory is centered on a handful of external physical measurements on living subjects, which he argues establish an ancestral tie between his pure “Elongated Africans” (Tutsi-Hima, Maasai and Ful) and the Afro-Asiatic-speaking populations of the Horn (Beja, Galla/Oromo, Somali, etc.). Numbering 11 variables in total, these anthropometric means include stature, head length, head breadth, face height, face breadth, nose height, nose breadth, relative trunk length, cephalic index, facial index and nasal index.
Somali ruler Mohamoud Ali Shire, the 26th Sultan of the Warsingali Sultanate.
Hiernaux has two Somali samples in his dataset, neither of which he measured himself. The first is a small sample of southern Somali individuals, “mostly of Sab descent”. These Sab Somali measurements were collected by the Italian anthropologist Nella Puccioni during an officially commissioned study published in 1917. The second, “larger sample of northern Somali belong[s] to various groups, the best represented being the Warsingali”. Hiernaux credits Lawrence Oschinsky (1954) with having gathered the Warsingali Somali measurements, although Oschinsky himself actually indicates that he obtained the data from “The Mediterranean Race in East Africa”, a chapter in Carleton Coon’s influential 1939 work The Races of Europe.
Tutsi ruler Charles Mutara III Rudahigwa, the 27th Mwami of the Kingdom of Rwanda.
Hiernaux suggests that the two Somali samples manifest substantial anthropometric differences. Specifically, he contends that the southern Somali cohort bears a close similarity with his Tutsi sample from Rwanda (a sample drawn from an early 1954 study of his). Hiernaux also posits that the northern Somalis “are strongly Arabicized”, based mainly on their shorter stature and more narrow nose and face. In doing so, he makes several important errors.
First, Hiernaux somehow overlooks or ignores one major specification that Puccioni provides in his original description of the Sab collection. Puccioni explicitly states therein that the southern Sab cohort has greater Negroid influence than the other Somali groups.
As the historian I. M. Lewis notes:
Within the Ethiopic group, the Somali belong to the eastern division, and show very few Negroid characteristics as compared with the western division, which is quite notably Negroid. As a result of his very extensive examination of Somali physical types, Puccioni considers that the southern Sab confederacies show a higher degree of Negroid influence, corresponding to their part Negroid origin.
On the Sab, Lewis points out elsewhere that:
Linguistically the speech of the Sab differs from that of the northern pastoralists by about as much as French does from Italian. The gulf in language is thus much wider than that between any of the northern pastoral dialects. The distinction has also a strong historical component, for, as I shall presently show, the Sab are a conglomerate people, an amalgam of many different Somali groups with Galla and negroid elements. And where they differ from the northern pastoral clan-families their distinctive culture and social institutions reflect these mixed origins.
This would explain why the Sab are closer to the Tutsis than are the northern Somalis. That is, the higher Negroid element among the Sab is bringing them nearer to the Tutsis than otherwise would have been the case. The Tutsis are likewise drawn toward the Sab due to their greater Cushitic element than other Bantus.
Similarly, Hiernaux apparently ignores Coon’s detailed analysis of his northern Somali sample. This too is unfortunate because Coon stresses the group’s overall “Mediterranean Caucasoid” affinities:
There can be no doubt that the tall stature of the Gallas, Somalis, and Agaus is an old Hamitic trait, since both the negroid Sidamos and the Semites of Hadhramaut origin are much shorter. The tallness of this East African Mediterranean strain stands in contrast to the moderate stature of the Mediterranean Arabs across the Red Sea, and constitutes a characteristic difference between them. The bodily build of the East African Hamites is typically Mediterranean in the ratio of arms, legs, and trunk, but the special attenuation of the extremities among the Somalis is a strong local feature, which finds its closest parallels outside the white racial group, in southern India and in Australia.
Second, Hiernaux does not offer any comparative explanation to support his claims, much less multivariate analysis. He just takes it for granted that his 11 anthropometric measurements alone are similar enough to substantiate these assertions. But are they really?
Mean stature figures for the Sab (Dighil and Rahanuìn) and other Somali clans (Puccioni (1931)). *N.B. Puccioni’s stature average for the Dir includes Garre individuals. When measured alone, Boughey (1971) observed that the Dir proper are instead primarily tall, with a mean height of 172.9 cm/1729 mm.
A closer look at Hiernaux’s stature figures shows that the Sab, whom he indicates are the tallest Somalis, are three centimeters or a little over an inch shorter than the Tutsis (173 cm vs. 176 cm). The Warsingali Somalis and Galla are a full eight centimeters/three inches (168 cm vs. 176 cm) and five centimeters/two inches (171 cm vs. 176 cm), respectively, shorter than the Tutsi. Likewise, with respect to the facial index, the Sab Somalis are mesoproscopic or have a medium face type (88.5), whereas the Tutsi are leptoproscopic or have a long narrow face type (92.8). The Warsingali Somalis are, by contrast, almost hyper-leptoproscopic (94.1). As it is, these values don’t particularly lend strong support to Hiernaux’s contention that the Tutsi anthropometrically parallel the Sab. Worse, Hiernaux appears to outright misrepresent Puccioni’s original height data on the Sab (see Table 108 and chart above). Puccioni (1931) actually lists two mean stature figures for the Sab individuals that he examined: Digil (Dighil) males at 1677 mm/167.7 cm and Rahanweyn (Rahanuìn) males at 1698 mm/169.8 cm. These values make the Sab on average the shortest of the Somali clans, thereby directly undermining Hiernaux’s claims.
Franco et al. (2013) provide helpful visual guides on these anthropometric variables:
Moreover, although Hiernaux emphasizes that the Tutsi’s and the Maasai’s relatively low nasal index is a key diagnostic feature that brings them closer to Afro-Asiatic-speaking groups than other Bantu and Nilotic populations, he completely ignores why those values are low to begin with.
The nasal index is calculated by dividing the nasal width by the nasal height, and then multiplying that quotient by 100. From this equation, it is clear that a marked nasal height alone is sufficient to produce a low nasal index figure, even if the nasal width is not particularly low. This is, in fact, the situation with the Tutsi and Maasai. Unlike what Hiernaux insinuates, neither population has a nose breadth within the low range of the Afro-Asiatic-speaking populations (lowest for the Warsingali Somali at 34 mm). Both instead have the same moderate nasal width as the Ful sample from the northwestern region of the Central African Republic (39 mm). However, because of their unusually long noses, the Tutsi and Maasai still wind up with lower nasal indices than the Ful (69.5 for Tutsi, 72 for Maasai and 79.7 for Ful). Here too, though, their average values remain distant from that of the Warsingali Somali, who have a leptorrhine nasal index of 66 — squarely within the narrow Caucasoid range. Radlauer (1914) observed a similar leptorrhine nasal index of 65.7 in a general northern Somali (Hashiya) sample, so the Warsingali average is close to the modal range for northern Somalis as a whole.
Oladipo et al. (2011) explain this standard pattern:
Nasal index measurement can be utilized in the analysis and classification of fossil remains as well as the study of living populations (Alex et al., 1996). Studies have shown that the Negroid race mainly of African descent have the platyrrhine nose type (Carleton, 1989).
[…]Oladipo et al. (2007) conducted a study on the morphometric analysis of the nasal parameters of Igbo, Ijaw and Yoruba ethnic groups of Southern Nigeria. The results obtained showed that an average Igbo had a mean nasal index of 94.1±0.37, Yoruba 89.2±0.30 and Ijaw 96.37±1.06. Thus the Ijaws had a significant higher nasal index (p<0.05). Fawehinmi et al. (2008) reported a mean nasal index of 98.5±0.93 and 94.1±1.18 for male and females of Kalabari ethnic group of Nigeria. The Somalia people in East Africa have a nasal index similar to that of European Caucasoid of 69.90 or less, which is of leptorrhine nose type (Porter et al., 2003; Carleton, 1989). The nasal index of African-American women is 79.70 Bantu speaking negreos and the bushmen of Africa as well as the Australoids of Australia are platyrrhine having a broad nose and a nasal index of 85.00 or more (Mulchand, 2004).
Nasal indices of Maasai individuals in Kenya. Most of the Maasai cohort have high nasal indices in the mesorrhine-platyrrhine (moderately broad) range, like other Great Lakes Nilotes. However, the presence in the sample set of leptorrhine (fine-featured) individuals of recent Cushitic origin — such as the “Mukogodo Maasai” i.e., assimilated Yaaku individuals from the Mukogodo forest — lowers the overall group average to a mesorrhine (moderate) nasal index (Leys and Joyce (1913)).
David Blackwell (1984) went further than Hiernaux’s basic nasal index calculations and photogrammetrically measured all aspects of the Tutsi Bantu’s and the Hehe Bantus’ nasal morphology. He found that there were no significant differences with respect to these Niger-Congo-speaking populations’ other nose-related indices, nor any marked disparities between them in terms of their nasal prominence, nasal bridge distance, columella length, nasal height, nasal angle, nasal wing/septum relationship, or nasal region vertical relationship. By measuring the nasal breadth index and inter-occular nasal width index, Blackwell was also able to gauge just how wide the average Tutsi’s nose is relative to his/her facial breadth. He found that while the Tutsi individuals he examined had a lower nasal index than his Hehe cohort, the sampled Tutsis in fact had a considerably larger nasal breadth index and inter-occular nasal width index than their Hehe counterparts. According to the scientist:
These results suggest that although the Hehe’s noses are wider in absolute terms the Tutsi’s are wider relative to their facial widths. The nasal index, which is nasal height divided by nasal breadth, was larger in the Hehe and since there was no significant difference in nasal heights this again suggests the Tutsi’s noses are wider relative to face width than the Hehe’s.
Note that although the Hehe are Bantu speakers, these anthropometric variables are also compatible with the idea that the Tutsi are of Nilotic origin since the coupling of a narrow head/face and a proportionately broader nose is a distinguishing characteristic of Nilotes (see discussion and anthropometric table below under Ecology, rock art and genetics).
As we saw in part one, the tall height of the Tutsis, as well as their long heads and less broad facial features than other Bantus, is due to contact with the Southern Cushitic “giants” of yore, whom the Tutsis’ ancestors assimilated. These physical attributes were not obtained from a mythical “Elongated African” entity, as Hiernaux muses. After having been introduced, these traits were, per the French social geographer Dominique Franche, maintained via selective breeding (though diet may have also played a part). Through this process, the Tutsis’ Bantu forebears would choose mates according to how closely those individuals conformed to the new Cushitic-influenced beauty ideals. This is evident given the antiquity and low-to-moderate level of Cushitic gene flow into the Tutsi population, which on its own would not have been enough to substantially modify that community’s ancestral Bantu morphology. Prior to said Cushitic admixture, the Iron Age inhabitants of Rwanda all had a physiognomy like their Niger-Congo-speaking relatives in West Africa, and most still do (cf. Rightmire (1975); Hiernaux (1960)). With regard to the Maasai Nilotes, who it has been established also have some Southern Cushitic influence, their shorter height relative to the Tutsi is largely due to the fact that they more intensively interbred with diminutive hunter-gatherer groups (cf. Dobon et al. (2015)).
(For more details, see Were the ancient Cushites slender?.)
Later anthropometric and craniometric studies
After the publication of Hiernaux’s The People of Africa, a number of studies were independently released, which contradicted, invalidated or otherwise corrected his “Elongated African” theory. The earliest of these works was Hiernaux’s own “Afrique moyenne” published the following year, in 1975. It was part of the Afrika ein volume in Rassengeschichte der Menschheit, a series on human diversity overseen by the anthropologist Eugene Strouhal. Based on anthropometric means, Hiernaux calculated the biological distance between hundreds of populations from all major ethnolinguistic groups below the Sahara. He included the Moors and Beja in his multivariate analysis, two populations that he had mentioned in passing in his book but did not actually examine. Hiernaux found that, after the Mbuti Pygmies (who, on account of their diminutive height, were obviously very divergent), the Afro-Asiatic-speaking groups on the northern fringes were the most distant. Of these latter populations, the Warsingali Somali were the furthest at a distance of 981, followed closely by the Bisharin subgroup of the Beja at 977 and the Moors at an average of 931. The Warsingali Somalis and Moors also formed a “constellation” unto themselves. Of the Bantu and Nilotic populations, the Tutsis of Burundi and Rwanda at averages of 859 and 829, respectively, were the nearest to the Afro-Asiatic-speaking groups. This is to be expected given their Cushitic admixture. Hiernaux, however, went further and suggested that this demonstrated that the Tutsi were not of Bantu origin but were instead settlers from the Galla area in southern Ethiopia, who later mixed with Bantu agriculturalists in the Great Lakes region and adopted their language — basically, a return to his initial Hamitic origin theory.
In 1976, G. P. Rightmire set out to evaluate the strength of Hiernaux’s biodistance analysis. Using both cranial metric and non-metric multidimensional scaling on 61 populations in Sub-Saharan Africa, including the Maasai Nilotes, he found no evidence for an “Elongated African” ancestral stock. Rightmire instead confirmed that the Afro-Asiatic-speaking populations of Northeast Africa were biologically distinct from the Nilo-Saharan, Niger-Congo and Khoisan groups. Of all the samples examined, the scientist observed that the Warsingali Somali of northern Somalia were overall the most distant, with the Oromo/Galla of Ethiopia and the Sab Somali of southern Somalia following suit:
This departure from “typical” African frequencies has been confirmed for various marker genes among Ethiopians (Ikin and Mourant, ’62; Harrison et al. ’69), so that the present peripheral position of the Galla and related Cushitic speaking groups is not surprising.
Additionally, Rightmire noted that the Nilo-Saharan-speaking Kunama of Eritrea also fell into the Afro-Asiatic cluster, a situation ascribable to substantial gene flow from the neighboring Afro-Asiatic-speaking communities (for Y-DNA evidence of such genetic introgression, see Trombetta et al. (2015), Supplementary Table 7; 65% of Kunama individuals have been found to bear the Afro-Asiatic-affiliated E1b1b haplogroup). This affiliation with Afro-Asiatic speakers can, alternatively, be interpreted as supporting the Kunama’s claim that they are not of Nilotic origin, but instead Abyssinian descendants who adopted a Nilo-Saharan language (Elisée Reclus (1899) notes: “The Kunama, established in the country since time immemorial, claim to be immigrants of Abyssinian origin, and the Abyssinians themselves look upon them as descendants of the ancient Aksumites”). For their part, the Maasai Nilotes of the Great Lakes craniometrically grouped with the other Nilo-Saharan and Niger-Congo samples, consistent with their comparatively lower Cushitic admixture. However, the Maasai did evince some non-metric affinities with the Afro-Asiatic-speaking populations. This in itself does not mean much, though, since research has demonstrated that non-metric/discrete/epigenetic trait analysis, whether cranial or dental, is really only useful for intra-population or within-population studies. Wilson (2010) explicates:
The suite of nonmetric traits expressed on an individual’s skeleton and dentition has been assumed to be genetically inherited. In other words, it is assumed that the phenotype (observable characteristics) of an individual will provide direct information about his or her genotype (genetic constitution). This assumption has allowed many researchers to use nonmetric traits to assess genetic relatedness within and between populations in the archaeological record (Matsumura, 2007). Understanding these relationships in past populations (especially those without written histories) can provide information about migration patterns, residence patterns, population structures, and human origins and evolution (Hanihara, Ishida & Dodo, 2003; Hlusko, 2004; McLellan & Finnegan, 1990; Lane & Sublett, 1972; Turan-Ozdemir & Sendemir, 2006). The term “biodistance” is commonly used to describe genetic relatedness. Saunders and Rainey (2008) describe biodistance as a measure of the amount of divergence; less divergence is equal to a closer genetic relationship (Saunders & Rainey, 2008; Sherwood, Duren, Demerath, Czerwinski, Siervogel, & Towne 2008). It should be noted that research has shown nonmetric traits to be population specific and therefore only really useful for intrapopulation analyses (Cheverud & Buikstra, 1981; Kohn, 1991).
When utilized in an inter-population or between-population context, non-metric trait analysis instead often results in spurious biological groupings, which contradict data collected through other, more scientifically robust methods. Sarfo (2014) thus remarks:
Epigenetic traits of the human skeleton, particularly of the skull, have had a long history in the study of paleogenetics. The last century, in particular, witnessed both the rise and fall of the use of epigenetic nonmetric traits in paleopopulation research. Part of the fall was attributed to rise of molecular anthropology in the 1980s and 1990s, whereby actual DNA could be analyzed to determine genetic relationships. This was coupled with problems in the research designs of the traditional morphologically-based paleopopulation genetics and the misunderstanding of the expression of epigenetic traits in the development of morphological variation. The pioneering article by Berry and Berry (1967) posited that nonmetric epigenetic cranial traits were highly genetic, were independent of each other, were independent of age and sex, and could be easily scored and standardized. Critical evaluation of these assumptions in the latter decades of the 20th century resulted in many challenges, particularly in the development of proper trait lists (Ossenberg 1976). For example, of the original 30 traits overviewed by Berry and Berry, only 7 can be confidently used to study paleopopulation genetics (Molto personal communication 2014). Yet, the vast majority of researchers in the period between the 1970s and 1990s utilized the Berry and Berry trait list with conflicting results arising as a consequence. Researchers who examined the assumptions generally found that many nonmetric traits were not independent of age, sex and symmetry (Ossenberg 1969, Suchey 1975, Molto 1985), and above all were not easily scored and standardized (Molto 1983). The latter is a fundamental requirement of the scientific method.
Eckhardt (1989) summarizes the problem with non-metric analysis in inter-population/between population studies as follows:
As noted in preceding sections of this review, there are reasonable prospects that better data on specific modes of inheritance of various skeletal traits, or at least estimates of heritabilities and the numbers of genes contributing to multifactorial characteristics, can help us to resolve one current paradox in studies of population affinity: that sometimes nonmetric traits suggest different patterns of relationship from those based on other data, including metric skeletal traits (Corrucini 1974). According to Brothwell (1959) nonmetric traits showed Chinese and Peruvian crania to be more closely related than Anglo-Saxon and London crania; in the same study Chinese were grouped more closely to North American Indians than Anglo-Saxons were to recent Germans or Melanesians to Polynesians. In another investigation Berry & Berry (1967) got results that placed South American Indians closer to Burmese and even to Africans than to North American Indians; also, Egyptian crania were placed nearer to those from South America than to a population from Palestine. In the other direction, Finnegan (1972) could not differentiate Florida Indians from several groups of Northwest Coast Indians.
Given the caveats above, it is hardly surprising that non-metric cranial and dental analyses of the Horn region’s Afro-Asiatic-speaking populations have shown little consistency. The conclusions of these studies have instead proven dubious, being at odds with cranial/dental metric and genetic analyses conducted by other researchers. For instance, Rightmire (1976) noted “Mediterranean” cranial non-metric affinities for his Cushitic samples (Warsangali Somali, Saba Somali, Oromo/Galla), whereas Hanihara (2003) reported that his sample of Ogaden Somali individuals (i.e. Ethiopian Somali) showed Sub-Saharan cranial non-metric ties. We can be confident that this is just another example of non-metric analysis going awry, as Vicente et al. (2019) genomically observed that their Somali North sample (Ethiopian Somali) actually has less Sub-Saharan admixture than their Somali South sample (Sab Somali). Likewise, Haddow (2012) analysed dental non-metric traits and found that their contemporary Ethiopia sample had closest biodistances to their ancient and modern North Africa samples, while Irish et al. (2007)‘s Tigrayan sample from Ethiopia/Eritrea instead shared nearest dental non-metric similitude with the ancient Jebel Moya sample from Sudan. We can be reasonably certain that this, too, is incorrect since Mukherjee (1955) indicates that the Tigrayan individuals he examined craniometrically cluster with the ancient Afro-Asiatic-speaking samples from North Africa; his Jebel Moya cohort, in contrast, craniometrically groups with the Sub-Saharan samples.
Moreover, recent genome research has identified specific alleles that are associated with craniofacial development (e.g. the EDAR, DCHS2, RUNX2 and GLI3 genes). These genetic findings have substantiated the validity of cranial/dental metric analysis for both intra-population and inter-population studies (Roosenboom et al. (2016); Adel et al. (2021)).
![Cranial metric analysis of Afro-Asiatic, Nilo-Saharan, Niger-Congo and Khoisan-speaking populations in Sub-Saharan Africa. The Afro-Asiatic-speaking Warsingali Somali, Oromo and Sab Somali samples cluster separately from the Sub-Saharan samples (including the Cushitic-admixed Maasai Nilotes), owing to ancestral ties with "Mediterranean Caucasoids." As Rightmire notes, "the departure from typical "African" frequencies has been confirmed for various marker genes among Ethiopians[...] so that the present peripheral position of the Galla and related Cushitic speaking groups is not surprising." The Kunama of Eritrea also fall in the Afro-Asiatic cluster due to significant gene flow from neighboring Cushitic and Ethiosemitic-speaking populations. This is the standard clustering pattern for these samples in metric analysis (both cranial and dental), which research has established is genetically inherited (Rightmire (1976)).](https://landofpunt.files.wordpress.com/2015/09/rightmire1976-fig2.png?w=264)
All in all, Rightmire attributes the morphological distinctiveness of the Afro-Asiatic speakers of the Horn of Africa to shared ancestral ties with “Mediterranean Caucasoids.” On this point, the linguist and anthropologist Harold C. Fleming remarks that Hiernaux’s peculiar perspective was likely shaped by his specialization in Central Africa, where the presence of any exotic influences among the local Niger-Congo-speaking populations is uncommon. Among the Afro-Asiatic-speaking groups in the Horn region, on the other hand (Fleming’s own area of focus), such “Mediterranean” traits are the norm. Fleming writes:
The most recent study of physical classification in Africa, GP Rightmire (1976), corrected and expanded Jean Hiernaux’s massive attempt to classify sub-Saharans. Rightmire used sophisticated measures of genetic distance and multidimensional scaling on the problem, but still kept his focus on black Africa. He however found that his Nilotes (Nuer, Alur, Kakwa, Luo, Masai) were distant from his northeast Africans (Sab, Somali and Galla) and gently suggested that Garn’s East Africans really meant northeast Africans rather than Nilotes. Hiernaux had also lumped Nilotes with Ethiopians apparently because of the linearity common to both — sometimes. I will take it as established then that Rightmire’s work has corrected Garn and Hiernaux with respect to southern Sudanese differences from Ethiopians. It is also clear, as a metaphysical matter, that Africanists who do physical classifications look at Ethiopia from the vantage point of the Congo and rarely consider “Mediterranean admixture” in terms other than the dilution of a basically African population. This viewpoint can come as a great shock to an Ethiopianist, accustomed to the vista from Bablyon.
An Ababda Beja girl, retaining the fine-features, facial orthognathism and soft-textured hair of her Cushitic Blemmyes (Bougaioi) ancestors.
In 1979, the anthropologists Strouhal and Jungwirth conducted a study, which compared the cranial affinities of modern East African “Negroes” from Rwanda and Burundi against those of late Roman-early Byzantine period skeletons excavated at Sayala, Egyptian Nubia. The Sayala specimens all belonged to the Pan-Grave culture and were exhumed from five cemeteries at the site (three cemeteries labeled CI-III, the N cemetery, and the A cemetery). Strouhal and Jungwirth report that most of the individuals from the CI-III and N burial complexes were of “Europoid” physical type (84.5%), whereas the individuals interred at the A cemetery were primarily of mixed “Europoid-Negroid” physiognomy (57.1%). The scientists also cross-analysed the discrete (non-metric) cranial traits of the CI-III cemetery Pan-Grave specimens with those of their Rwanda and Burundi “Negro” samples, and concluded that there was no affinity. This finding once more underlines the non-Cushitic origins of the Tutsi-Hima Bantus since the “Europoid” Pan-Grave culture bearers of the CI-III cemetery — with whom the East African “Negroes” from Rwanda and Burundi share no morphological ties — are believed to have originally been of Cushitic ancestral stock. Specifically, Strouhal and Jungwirth assert that “the Sayala CI-III series would represent the Blemmyes, a population originating from the Eastern Desert.” An early Coptic inscription refers to the Blemmyes as the Bougaioi, an appellative that is rendered as Beya in a later text retrieved from Adulis. This is an obvious allusion to the Cushitic-speaking Beja people, who still inhabit the Eastern Desert like their Blemmyes ancestors at the CI-III cemetery in Sayala, Egyptian Nubia (cf. Winstedt (1909)). (*N.B. In due course, some Nilotic peoples would gradually have been assimilated into the Pan-Grave community. These outsiders appear to have contributed the alien “Negroid” and mixed “Europoid-Negroid” physical types that have also been observed at various Pan-Grave sites, opposite the original “Europoid” physical type of the Cushitic Blemmyes.)
Anthropometric analysis of various Afro-Asiatic, Niger-Congo, Nilo-Saharan and Khoisan-speaking samples. The unadmixed Cushitic speakers from Northeast Africa (Beni Amer Beja and Hadendoa/Amara Beja) are distinct from all of the examined Sub-Saharan African populations, sharing close affinities only with each other (1.09 biodistance). The mixed Cushitic sample from Southeast Africa (Iraqw) is nearest to the Datog Nilotes, due to documented intermarriages. The Tutsi Bantu individuals are not especially close to any population, though they do bear somewhat remote ties with the Datog Nilotes (3.06 biodistance). The Sandawe foragers of Tanzania are morphologically very similar to the adjacent Nyaturu Bantus. Of the hunter-gatherer/Nilo-Saharan/Niger-Congo populations, the Sandawe are also the least isolated from the unadmixed Cushitic samples (3.94 biodistance from the Beni Amer Beja). This is attributable to absorption of Cushitic pastoralists by the Tehla or herder division of the Sandawe (Ikeda and Hayama (1982)).
Ikeda and Hayama (1982) published an anthropometric analysis comparing the Tutsi Bantus, Hutu Bantus and other Niger-Congo-speaking populations with various Nilo-Saharan, Khoisan and Afro-Asiatic-speaking groups. The study is unique in that it included among its samples unadmixed Cushitic individuals from Northeast Africa (viz. Beni Amer Beja, Hadendoa Beja and Amara Beja), Cushitic individuals from Southeast Africa with significant Niger-Congo, Nilo-Saharan and hunter-gatherer admixtures (Iraqw), and Niger-Congo, Nilo-Saharan and hunter-gatherer individuals with significant Cushitic admixture (Tutsi, Datog and Sandawe, respectively). The unadmixed Cushitic groups were found to be distinct from all of the examined Sub-Saharan African populations, sharing a close relationship only with each other (1.09 biodistance between the Beni Amer and Hadendoa/Amara Beja samples). The mixed Iraqw from Tanzania were nearest to the Datog Nilotes, consistent with their documented intermarriages (0.77 biodistance). They also showed more remote ties with most of the other samples, which are in line with known population contacts. Interestingly, the Tutsi individuals were not particularly closely affiliated with any sample; their nearest association was with the Datog cohort, at a notable biodistance of 3.06. This was not the case for the Sandawe, who shared very close morphological affinities with the neighboring Nyaturu Bantus of northern Tanzania (0.41 biodistance). Of all the studied groups, the Sandawe were also the least isolated from the unadmixed Cushitic samples, at a biodistance of 3.94 from the Beni Amer Beja. This suggests that most of the Sandawe individuals in the dataset belonged to the Tehla, the herder division of the Sandawe forager community, which is known to have assimilated many Cushitic pastoralists (due to these past interactions, the Sandawe today have an average of ~30% Eurasian admixture; see Scheinfeldt et al. (2019)).
Anthropometric relations between ethnolinguistic groups in Africa and the Arabian peninsula. The Cushitic and Ethiosemitic-speaking populations in the Horn of Africa cluster with other Afro-Asiatic speakers in North Africa (northern Libyans and Egyptians) and the Arabian peninsula (Yemenis). Nubians also group with the Afro-Asiatic-speaking samples rather than with the Nilo-Saharan and Niger-Congo-speaking populations (Billy (1988)).
In 1988, G. Billy compared the morphological relationships of the Afro-Asiatic-speaking populations in the Horn with each other and with neighboring groups. He used the same northern Somali and southern Sab samples as Hiernaux did (taken from Coon and Puccioni, respectively). However, unlike Hiernaux, Billy did not restrict his samples to populations below the Sahara since he also examined other Afro-Asiatic speakers from North Africa and Southern Arabia. Running various multivariate analyses, which Hiernaux had likewise neglected to do in his book, Billy found that the Afro-Asiatic-speaking groups were all morphologically closest to each other. Furthermore, contrary to Hiernaux’s suggestion therein that the Somalis were anthropometrically intermediate between the Yemeni Arabs and the Tutsis and that the Galla were intermediate between the Somalis and the Tutsis, the Somali and Galla as well as the other northern Horn samples clustered nearest to all four Yemeni samples in the dataset in addition to the littoral Cyrenaican/northern Libyan and Egyptian samples, but not with any Bantu or Nilotic population. Unsurprisingly, given Puccioni’s caveat, the Sab sample grouped closest to populations inhabiting southern Ethiopia (mainly Omotic groups). This was due to what Billy described as a greater morphological tendency on their part toward broader noses and faces. However, the Sab were still closer overall to the other Afro-Asiatic-speaking populations.
Billy thus concluded that:
It appears that populations living at present in Egypt and Sudan are weakly influenced by the populations of Equatorial Africa but strongly related to coastal populations of Ethiopia and the arabian peninsula to a lesser extent.
The physician and anthropologist Alain Froment, the current Scientific Director of the anthropological collections at the Musée de l’Homme in Paris, would similarly highlight the morphological proximity between the Afro-Asiatic-speaking populations on either side of the Red Sea. Froment had worked closely with Hiernaux earlier in his career (they co-authored at least one publication in the 1970s), so he was quite familiar with his colleague’s theories. In a 1992 study, Froment, apparently unaware that Somalis and Gallas are Afro-Asiatic speakers with a different biohistory from Nilo-Saharan groups, lumped a Somali and Galla joint sample with a Nilotic Maasai one. Predictably, he found that his combined sample pulled towards Europe due to the Somali and Galla’s “Caucasoid” craniometric affinities, but also towards the Sub-Saharan African centroid on account of the presence of the Maasai Nilotes in the sample. By 1994, Froment had grown more familiar with the marked biological differences between the Afro-Asiatic-speaking populations in the Horn and Sahara versus their Bantu and Nilotic neighbors. In a retrospective paper on the ancient Egyptians published that year, he noted his joint Somali-Galla sample’s close craniometric ties with the Proto-Mediterraneans as well as with other Afro-Asiatic-speaking populations in North Africa and the Middle East. He also criticized Hiernaux’s consistent omission of North African samples from the latter’s various analyses. Froment quite rightly remarked that this oversight on the part of Hiernaux made it impossible to ascertain the actual morphological transitions between the Horn region and the Mediterranean Basin.
Additionally, during his The People of Africa phase, Hiernaux had argued that changes in morphology between populations inhabiting different regions below the Sahara were gradual and clinal. However, both Froment and W. W. Howells found evidence to the contrary in their multivariate analyses. They instead noted a sharp morphological discontinuity between the Afro-Asiatic-speaking populations in the Horn and the adjacent Bantu and Nilotic groups. I. M. Ribot writes:
Multivariate analyses of both Howells (1989) and Froment (1992a, b) agreed on the following points: in contrast to the observations of Hiernaux (1976), the variation within sub-Saharan Africa was not entirely continuous, as significant differences were observed between regions; and these marked geographical differences were well shown especially when using two cranial variables related to both bizygomatic breadth of face and nasal breadth.
[…]Furthermore, in agreement with Howells (1989) and Froment (1998) again, but in contrast to the observations of Hiernaux (1974), the range of variation in sub-Saharan Africa was not continuous or clinal, as significant differences were still observed between the different regions.
Craniometric relations between populations in Africa, the Middle East and Europe (Froment (1999)).
In 1999, Froment conducted another large-scale craniometric analysis for a paper on Sahelian populations. This time he included separate Somali-Galla and Sab samples, as well as Maasai, Tutsi, Ful (Peul), Berber, Nilotic and West African samples and Egyptian, Maghreban, Tuareg, Sub-Saharan, Middle Eastern and European centroids — 636 male and female global populations in total. Froment used six cranial measurements, including head length and head breadth, face length and face breadth, and nose length and nose breadth. He found that the Galla-Somali lumped sample was distant from the Maasai, Tutsi and Ful samples, and was instead nearest to the Middle Eastern and Bedouin/Lower Egyptian centroids. On the horizontal axis, where most of the variation between the samples was contained (77%), the Galla-Somali sample was again closest to the Bedouin/Lower Egyptian centroid in addition to the Sab Somalis, the Tuareg centroid, the Maghreban centroid, the Moors of the Western Sahara (the Sahrawi), and the Kel Kummer male Tuareg sample. The Sab Somali sample was, in turn, nearest to the Tuareg and Bedouin/Lower Egyptian centroids, while being closest to those samples on the key horizontal axis as well as to the Galla-Somali lumped sample, the Kel Kummer males and the Maghreb centroid.
Craniometric analysis of Afro-Asiatic, Niger-Congo, Nilo-Saharan and Khoisan populations. The Fulbe (Peul) and Tutsi samples are positioned on the margins of the Niger-Congo/Nilo-Saharan cluster, with a pull toward the separate Afro-Asiatic cluster. In the case of the Fulbe, this placement is primarily due to gene flow from ancient Berber and European populations. In the Tutsis’ case, this clustering pattern is attributable to both absorption of South Cushitic groups and sexual selection. Since the Tutsis experienced less overall gene flow from Afro-Asiatic-speaking populations than did the Fulbe, they would have had to have turned to careful mate selection in order to further alter their ancestral Bantu morphology (Froment (1998)).
In 2013, Terrazas Mata and Benavente craniometrically compared Tutsi, Hutu and other Bantu individuals from eastern Africa with various Niger-Congo, Nilo-Saharan, Khoisan and Afro-Asiatic-speaking populations. Their study is unique in that it includes a sample of an early Tutsi king, one Cyirima Rujugira (r. 1675-1708), borrowed from Ribot (2003). Hiernaux had previously analysed this ruler’s skeletal remains and reported that Cyirima shared affinities with the modern Tutsi (e.g. same average height of 176.539 cm) and to a lesser extent with the Hutu (cf. van Noten (1972)). Terrazas Mata and Benavente subsequently confirmed the close ties between the Tutsi king and Ribot’s other Bantu samples, lumping these related Niger-Congo-speaking individuals into a cohort labeled “Eastern Africa” (comprising King Cyirima and Rwandan Tutsi, general Burundi, Pare, Shambaa, Dschagga, Teita and Haya persons). The scientists found that this Eastern Africa Bantu sample clustered with two other Bantu-speaking groups from the Great Lakes region, the Hutu and Teita, “all known to be archaeologically and ethnographically similar and probably recently derived from a common ancestral population.” By contrast, their Horn of Africa lumped sample, consisting of Cushitic-speaking Somali, Beja and Oromo/Galla individuals, clustered instead with the Dynastic Egyptian sample. Terrazas Mata and Benevente indicate that this is because “these groups seem to represent migrations of groups from the Middle East, who spoke languages from the Afro-Asiatic family.”
Thus, contrary to Hiernaux’s suppositions, Terrazas Mata and Benavente, Froment, Billy and Rightmire had all empirically established that the Somali, Oromo/Galla and Beja are morphologically much closer to other Afro-Asiatic-speaking populations in North Africa and the Arabian Peninsula than to Nilo-Saharan and Niger-Congo-speaking populations, including groups with some Cushitic or Berber admixture like the Maasai, Tutsi and Ful.
Serology
Somali herdsmen (Paulitschke (1893)).
Besides anthropometry, Hiernaux in his book The People of Africa relies on serology, or blood work, to support his “Elongated African” hypothesis. He focuses on relative differences in frequencies in two serological systems: the M allele in the MN system, and the R0 (cDe) allele in the Rh system. With regard to M, Hiernaux indicates that the average frequency for the allele in Sub-Saharan Africa is around 49%. He writes that the mean percentage for the R0 allele in the same area is over 65%. Hiernaux regards any marked deviations from these average frequencies as potential indicators of exotic influence in a given population. To his credit, he then attempts to corroborate this with other lines of evidence.
Beja nomads (Seligmann (1913)).
Since Hiernaux already ruled out that the Neolithic pastoralists of East Africa were of Caucasoid stock (unjustifiably, as we saw in part one), preferring instead to assign to these first Afro-Asiatic-speaking settlers a speculative “Elongated African” origin, he is obliged to explain away the many exotic biological influences in the Horn as being recent imports from Southwest Asia. Hiernaux therefore prefaces his serological discussion by remarking that “history and culture show Ethiopia and the Horn of Africa to be strongly impregnated by Arabic influences”. Specifically, he alludes to four waves of settlement from Arabia: one represented by the Sabaeans, whom he suggests established the Axumite Kingdom in the Ethiopian and Eritrean highlands; another by the ancestors of the Gurage; a third wave consisting of Jewish proselytizers; and a final wave comprising the Muslim patriarchs of the various Somali and Afar clans. From this limited perspective, Hiernaux interprets any convergence or divergence of blood group frequencies from the Arabian mean as being equivalent to greater or lesser Caucasoid ancestry, respectively.
Qara Southern Arabian tribesmen (Thomas (1929)).
Thus, by Hiernaux’s reckoning, the Afar of Djibouti have considerable Arab influence since they have a relatively high frequency of 62% of the M allele, and the “MN blood groups are also useful for assessing a genetic influence from Arabia[…] the M allele has a high frequency in Arabia, above 65 per cent; in sub-Saharan Africa its frequency varies between 31 and 73 per cent”. He asserts that the Galla, Gurage, Amhara, Falasha and Tigre of Ethiopia and Eritrea, who are also Afro-Asiatic speakers, are in a similar position, for “all these populations have high M frequencies of between 63 and 68 per cent”. On the other hand, despite acknowledging strong anthropometric ties between the northern Somalis and Arabs, Hiernaux argues that the northern Somalis (whom he defines as all major Somali clans except for the Sab) have a comparatively weaker serological connection with Arabia because the M allele frequency is a lower 51% among both the Dir Somalis of northwestern Somalia and the Issa Somalis of Djibouti. This happens to be the standard percentage for ethnic Somalis as a whole (see Mourant’s serological Table 1 below). However, what Hiernaux overlooks or is perhaps unaware of is that moderate frequencies of around 50% for the M allele are actually the norm among many other Afro-Asiatic-speaking populations in the circum-Mediterranean area, including Egyptians. In fact, El Hassan et al. (1968) observed an even lower percentage of 48% among the Beja of Sudan. Arthur Ernest Mourant, who pioneered the use of serology and other genetic systems in anthropology, explains the situation thusly:
The frequency of the M gene is high, mostly well above 60 percent, in nearly every Indian population tested, whereas it is below 60 percent and mostly below 55 percent in the Mediterranean area, apart from Sardinia. Insofar as the scanty data available allow us to judge, there appears to be a sudden rise from low, typically European and Mediterranean M frequencies in Turkey (both in Turks and in Ed-Turks) and Lebanon and in the Armenians, to much higher values in Syria, Arabia, Iraq, and Iran. Egyptians show low, typically Mediterranean frequencies, but higher values occur in Sudan, Ethiopia, and Somalia, where they appear on the map as an extension, unique in Africa, of the high M area of Asia.
In other words, what Hiernaux mistakenly interprets as weaker serological Caucasoid affinities for northern Somalis compared to surrounding Afro-Asiatic-speaking groups are actually simply a different type of Caucasoid affiliation (i.e. more Egyptian than Arabian on this particular blood group trait).
As regards his pure “Elongated Africans”, the Tutsi-Hima, Maasai and Ful, Hiernaux is again inconsistent. He is silent on the M allele frequencies for the Maasai and the Ful. Besides the Nama Hottentots of Southern Africa (whom he admits have some ancient Cushitic admixture), he does, though, remark that “the only other sample to show an M frequency above 60 per cent is a small one of Tutsi from Rwanda and Burundi”. Nonetheless, Hiernaux does not propose this unusually high M percentage as evidence of Ethiopian or Arabian introgression into the Tutsi population. He instead maintains his untenable position that the Tutsi Bantus have virtually no exotic genetic influences.
With respect to the Rhesus (Rh) blood group system, Hiernaux writes that the R0 allele’s “frequency varies between 33 and 95 per cent in sub-Saharan Africa, while it never attains the lowest of these values in North Africa or Arabia, and in Europe it does not rise above a few per cent”. He thus asserts that “the R0 frequency is therefore useful for tracing an exotic influence”. On this basis, Hiernaux indicates that “R0 frequency has relatively low values also in northern and central Ethiopia, which has a long history of contacts with southern Arabia”. He notes that these R0 allele percentages are typically near or below 50% among said Afro-Asiatic-speaking populations; at around 48% for the Tigre and Falasha, 54% for the Galla, and 35% for the Gurage. Since the Afar of Djibouti have a similar R0 frequency of 51% whereas the Issa Somalis of the same territory have a much higher R0 frequency of 65%, Hiernaux argues again that Arabian influence is significantly greater among the Afar and the central and northern Afro-Asiatic-speaking Ethiopian groups than among the northern Somalis. But is this really justified?
Blood group frequencies of various Afro-Asiatic-speaking populations and Nubians in Northeast Africa. From left: cDe (R0), CDe (R1), cDE (R2), CDue, cDue, Cde, cde (r), cdE (Fleming (1965)).
Hiernaux’s claim is based on R0 frequency data gathered by Fourquet (1970), who analyzed the blood group means of Somalis from a single clan, the Gadaboursi and Issa Dir of Djibouti. However, this percentage appears to have been distorted by genetic drift. This is strongly implied by Iacovacci et al. (2017), who indicate that 100% of their Djiboutian Dir individuals belong to the paternal haplogroup T, while this Y-DNA clade normally occurs at moderate frequencies of ~15% among Somalis from other clans (cf. Sanchez et al. (2005); Immel and Kleiber (2009)). Furthermore, Fleming (1965) reports a much lower R0 frequency of 38% in a general (non-Sab) Somali sample, and percentages of 40% for the Amhara, 48.8% for the Bilen, and 53.8% for the Arsi Galla. Sistonen et al. (1987) likewise observed a ~38% R0 (cDe or DCe) frequency in their large sample of 1026 ethnic Somali individuals in Somalia (cf. Table II). According to Fleming (1979), the blood group data for the Chaha Gurage (which Hiernaux cites) also contains errors in that it mistakes cDE for cDe, thereby reporting lower cDe/R0 frequencies for the Gurage than this population actually has. On balance, going by Hiernaux’s way of thinking, this would suggest that there is greater Arabian serological influence among Somalis than among most other Afro-Asiatic speakers in the Horn. (*N.B. Sistonen et al.’s data is weighted more towards southern Somali individuals, with 59% of their cohort indicating that they were born to southern clans. The scientists, moreover, concede that the southern “Sab are generally considered to be of a more mixed descent, assimilating elements from virtually all of the other clans and, probably, also some from the Galla and Negroid populations the remnants of which still live around the rivers.” This greater Negroid influence explains why, in some of the other serological systems that they examined, their sample set appears more shifted toward hunter-gatherer populations than is usually the case.) Rather than recent Arabian introgression, though, it is more probable that the relatively low cDe/R0 frequencies carried by Somalis were simply bequeathed to them by their Cushitic ancestors (who presumably had an even lower cDe percentage). For one thing, similar cDe frequencies have been observed among some other Afro-Asiatic speakers in North Africa, whom we know from ancient DNA analysis are related to the early Cushites (e.g. Metri et al. (2012) report cDe/R0 frequencies of 31% and 37%, respectively, among Maghrebi individuals from Sid-El-Djilali and M’sirda in the Tlemcen region of western Algeria; Merghoub et al. (1997) likewise reveal a cDe or R0 (Dce) frequency of 32.9% among Mozabite Berbers from Mzab, Algeria; similarly, Skaik (2011) indicates a cDe frequency of 30.4% among Palestinian females in Gaza).
Blood group frequencies among populations in Northeast Africa, Southern Arabia and Southern Europe (Mourant et al. (1976)).
In his landmark thesis The Blood Groups of Somali Tribes published in 1959, K. L. G. Goldsmith collected serological data from over 1000 ethnic Somali individuals inhabiting the former British Somaliland protectorate. His large sample set was drawn from various clans, and is thus representative of the blood group characteristics of northern Somalis in general. Goldsmith observed that the Somalis had an average R0/cDe frequency of 48%, well within the typical range for this allele among the other Afro-Asiatic-speaking populations in the Horn. Mourant et al. (1976) highlight this in their Table 1 (shown to the right).
Blood group frequencies among the Tumaal and Midgaan (Goldsmith and Lewis (1958)).
In his defense, Hiernaux may not have been aware of Goldsmith’s study, as he does not include it among his bibliographical references in The People of Africa. He does, though, cite Goldsmith’s earlier paper from 1958, A Preliminary Investigation of the Blood Groups of the Sab Bondsmen of Northern Somaliland, which I. M. Lewis co-authored. This is quite interesting since Goldsmith and Lewis provide an average R0/cDe frequency for the northern Midgaan and Tumaal low-caste groups, who, along with the Yibir, are collectively known as the Gabooye (not to be confused with the southern Sab or Rahanweyn confederacy). At around 43%, it is almost identical to Fleming (1965)’s R0 value of 42.5% for the same Sab groups. Both are also within the typical allele percentage range of the other Afro-Asiatic-speaking groups in the Horn.
A Gabooye Somali woman. Serological analysis of the low-caste groups among the Somali, who are collectively known as the Gabooye, indicates that they are genetically similar to their higher-caste counterparts. This suggests that the Gabooye are of Cushitic origin like other ethnic Somalis, but probably became differentiated on the basis of their non-pastoralist occupations.
Thus, contrary to Hiernaux’s assertion, a value near 40% is the actual mean percentage for the R0 allele among northern Somalis, including the low-caste groups. Elizabeth W. Ikin, who co-wrote a series of influential blood group studies with Mourant on various populations in Ethiopia, further confirms this in her own 1959 study on blood group distribution in the Middle East. She therein remarks that the average Somali R0 frequency is around 43%, apparently in allusion to Goldsmith and Lewis’ analysis. In his 1980 book Human Biological Diversity co-authored with Amitabha Basu, Hiernaux himself would eventually acknowledge the relatively low R0 frequencies found among both ethnic Somalis and the Beja (perhaps he had since gotten wind of Goldsmith, Ikin and Mourant’s serological work).
With regard to the Rhesus frequencies among the Tutsi Bantus and their Hima kinsmen as well as the Maasai Nilotes and Ful West Africans, Hiernaux is both more and less straightforward. He writes that “moderate frequencies of R0 [are] found in some populations in which an exotic influence has been suspected, in the pastoral Ful of northern Nigeria and in the Hima of Ankole (Uganda)”, yet asserts elsewhere that “as a whole, the Ful and the non-Ful of West Africa are very similar in their allele frequencies for blood trait systems”. Hiernaux also notes that the Ful from Senegal, the Tutsi and the Maasai all have high R0/cDe frequencies like the Hutu and most other populations below the Sahara. This is in direct contrast to Excoffier et al. (1987), who, on the basis of a lack of the sickle-cell trait and apparently lower R0 frequencies than other Bantus (percentages which the authors never divulge), suggest that the “Tutsi and Hima [are] surrounded by Bantu populations but closer genetically to Cushites and Ethiosemites, [and] are known to have migrated from northern territories recently”. However, Hiernaux’s assertion is somewhat in agreement with Fleming (1979), who observes that:
In Rhesus and other systems, however, it is clear that none of the major contemporary Kenyan populations can be classified as Ethiopid, except for the Oromo and Somali. Neither Luo, nor Masai, nor Kikuyu emerge as anything but ordinary members of Nilotic and Bantu clumps.
Similarly, Lawrence Oschinsky (1960) observed:
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 indicates 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.
Despite emphasizing the importance of serology in understanding the origins of a given population, Hiernaux does not touch on the ABO blood group systems that are distributed in Northeast Africa. This is regrettable since the O allele that is found at especially high frequencies in Arabia is almost as common among Somalis, Beja and certain other Horn communities as well as among the Berbers of the Maghreb. Coupled with a low occurrence of the more typical African markers (such as the P1 blood group; see Mourant’s Tables 1 and 2 above), this altogether points to a strong serological connection between the Afro-Asiatic-speaking populations on either side of the Red Sea. Mourant (1983) writes:
The main feature of ABO distribution in the region as a whole is the marked westward diminution in the frequency of O from the high levels found on the east coast together with a rise of B and, in certain areas, especially Ethiopia, a rise of A. The high O and low B, especially in the Beja of the eastern Sudan and the Somalis, presumably indicate Arabian admixture in the population concerned, as might be expected from what we know of their history.
El Hassan et al. (1968), on which Mourant served as a co-author, similarly notes that:
In two respects the Beja differ from most of the surrounding peoples, namely in their low frequencies of B and of M. In their low frequency of B they resemble the southern Arabs and the Somalis, though they differ from both in their much higher frequency of all the typical African marker genes. In their lower frequency of M they resemble, on the other hand, some of the African peoples of north-east Africa who possess few of the typical Mediterranean genes, but also the Somalis, who possess many such genes.
Ikin (1959) goes further and asserts that the serological ties between Somalis and Southern Arabs are such that they form their own “Arabian” sub-class of the “Mediterranean race”:
The peoples of the Near East thus fall into two main classes. We have the Turks and Eti-Turks, whose relations appear to be mainly with Europe, and especially with the Mediterranean area. They have a high A gene frequency, a moderately high M gene frequency, together with a high R1, and a fairly high rh-negative frequency. Then we have the “Arabians”, a sub-class of the “Mediterranean” race. These are the Yemenite Arabs and Jews, the Zabidi Arabs and the Socotrans and, with reservations, the Somalis. These all have low A and high O gene frequencies, mostly high R1 and rather lower rh-negative gene frequencies. They also have a very high M frequency which is possibly distinct from the general high M of Asia. It is interesting to note that the Berbers, the “white” race of N. Africa, resemble the Arabs in having a high O frequency, but differ from them in having high N and high Rh-negative frequencies.
Froment (1994) likewise confirms that the blood group traits of Somalis, Afars and other Afro-Asiatic-speaking populations of the Horn, as cataloged in Mourant’s 1976 tome The Distribution of the Human Blood Groups and Other Polymorphisms, show considerable Arabian affinities.
Phylogenetic tree of the blood group affinities of the Afro-Asiatic, Khoisan, Niger-Congo and Nilo-Saharan-speaking populations in Africa. The Afro-Asiatic speakers form two clusters: the first joins Libyans, Berbers and Bedouins of North Africa with Somalis of the Horn at a bootstrap value of 75.6%; the second comprises Tigre, Amhara and Afar of the Horn, with Tigre and Amhara joined at a bootstrap value of 50.8% (López (2013)).
In 2013, Janire Allende López of the Universidad del País Vasco compared the blood group traits of a large selection of Afro-Asiatic, Khoisan, Niger-Congo and Nilo-Saharan-speaking communities inhabiting Africa. Like Mourant, El Hassan, Ikin and Froment before him, López observed that the populations generally grouped along linguistic lines. The Afro-Asiatic speakers formed two such phylogenetic clusters: the first grouping comprised the Libyans, Berbers and Bedouins of North Africa and the Somalis of the Horn of Africa, joined together at a bootstrap value of 75.6%; the second grouping consisted of the Tigre, Amhara and Afar of the Horn, with the Tigre and Amhara sharing a branch at a bootstrap value of 50.8%. The Nilo-Saharan-speaking Baria/Nara of Eritrea also fell into the latter Afro-Asiatic cluster, evidently due to their documented intermixture with local Cushitic- and Ethiosemitic-speaking peoples (cf. Trombetta et al. (2015), Supplementary Table 7; around 60% of Nara individuals today carry the E1b1b modal Afro-Asiatic paternal haplogroup on account of these contacts). On the other hand, the Maasai Nilotes in López’s sample set clustered with other Nilo-Saharan and Niger-Congo populations, just as Fleming had earlier remarked:
Las poblaciones de afroasiáticos, excepto los Funji, se encuentran agrupadas. Por un lado observamos las poblaciones de Libios y Bereberes unidos en una rama con un valor bootstrap de 51% y a los Libios, Bereberes y Beduinos con un valor bootstrap de 75,6%, todos ellos relacionados a los Somalies. Mientras que los Tigre, Amhara, Afar y Baria/Nara se encuentran en otra rama, las poblaciones Tigre y Amhara están unidas en una rama con un valor bootstrap de 50,8 %. Los pigmeos (Mbuti y Biaka) y los Hadza están relacionados con un valor bootstrap de 73%. En cuanto a la familia Khoi-Sanida, se observa que los Khoi y los San están en la misma rama. Por último las familias de Niger-Congo y Nilo-Saharianos se agrupan en el centro del árbol filogenético.
Afro-Asiatic populations, except the Funji, are grouped. On the one hand we observe the populations of Libyans and Berbers united in a branch with a bootstrap value of 51% and the Libyans, Berbers and Bedouins with a bootstrap value of 75.6%, all related to the Somalis. While the Tigre, Amhara, Afar and Baria / Nara are in another branch, the Tigre and Amhara populations are united in a branch with a bootstrap value of 50.8%. Pygmies (Mbuti and Biaka) and Hadza are related with a bootstrap value of 73%. As for the Khoi-San family, it is noted that the Khoi and the San are on the same branch. Finally, the families of Niger-Congo and Nilo-Saharan are grouped in the center of the phylogenetic tree.
Ecology, rock art and genetics
The final pillar of Hiernaux’s “Elongated African” theory focuses on ecology. Specifically, he argues that the linear physiques that are common among the populations inhabiting the southern rim of North Africa evolved as long-term morphological adaptations to hot and arid biotopes or habitats. He writes:
In the hot and dry zones of sub-Saharan Africa, selection has favoured a different body build, which also displays a low mass-to-surface ratio : a gracile, elongated physique, with long legs and narrow shoulders. Weight is relatively very low in such people of medium or tall stature, but not especially so in absolute value. This tendency of the human physique toward tallness in the hot and dry zone explains the failure of Bergmann’s rule in sub-Saharan Africa. Like size reduction, it represents an adaptation to heat, but this time in a dry climate in which cooling by sweating is the easiest.
Hiernaux stresses that head, face and nose form is also linked with environmental variables:
Head and face shape also are significantly correlated with air temperature and moisture in sub-Saharan Africa. The more extreme are heat and dryness, the narrower tends to be the head. Face width tends to increase with air moisture and climatic uniformity. Nose height tends to be lower in wetter regions. Of all measurements, nose width shows the closest correlations with climate: it tends to increase with the annual rainfall and to decrease with the temperature of the hottest month.
On the surface, the various anthropometric tables that Hiernaux provides in The People of Africa indeed appear to support his theory on the correlations between body dimensions and habitat. That is, until one closely examines his data table on the “pure” northern Nilotes of the Nile Valley:
As can be seen in the anthropometric means above, while all of Hiernaux’s Nilotic populations are indeed tall and possess very narrow heads and relatively narrow faces, their nose heights and nose breadths are not what one would expect from individuals whose ancestors supposedly evolved in a hot and dry habitat. Instead, they have wide and flat noses like the average Bantu and West African population — groups that Hiernaux insists evolved in a moist, jungle environment — with the platyrrhine nasal indices to match. This also contradicts Hiernaux’s assertion that “obviously a narrow face could not accommodate a very wide nose” since the Nilotes do, in fact, possess both morphological traits at once. Altogether, the anthropometric means suggest that the Nilotes evolved in a forested biotope similar to that which molded the Bantus, West Africans and Pygmies, with the main difference being that the Nilotes likely inhabited a sunnier portion of the same area.
On some level, Hiernaux appears to be aware of the common origin of the Nilotic, Bantu and West African populations, for he alludes to a “most numerous and most widely spread African stock which prevails in West and Central Africa”. He indicates that this “West Central African” ancestral stock (essentially a euphemism for the “Negroid” taxon of traditional physical anthropology) is best represented in the archaeological record by populations from the Mesolithic Wadi Halfa in Sudan and the Neolithic and protohistoric southern Sahara, such as the Asselar and Iwo Eleru specimens.
Hiernaux also concedes that the aforediscussed “East African skeletal material is not closely related to the Mesolithic Wadi Halfa remains”, yet speculates that “these people may have evolved from the stock represented by the Wadi Halfa population”. But is there any evidence to support this association? As discussed in detail on Ancient DNA from Sudan, the Mesolithic inhabitants of Wadi Halfa (the immediate ancestors of the Nilotes) are instead both genetically and morphologically distinct from the later A-Group, C-Group and Kerma and Meroitic, Post-Meroitic/X-Group and Christian period inhabitants of the Nile Valley (the ancestors of the Afro-Asiatic-speaking populations in Northeast Africa). Hiernaux is therefore mistaken about that as well.
So where, then, does Hiernaux believe his “Elongated African” morphology originated? He postulates that it developed in the Neolithic Sahara among early pastoralists, and subsequently spread from there during one of the region’s various dry periods. The migrating herders would then have undergone further local evolution in the arid zones of East Africa and the Nile Valley. As mentioned earlier in the essay, Hiernaux suggests that these Saharan pastoralists are represented by the C-Group people. This is quite ironic since, as we also saw, the C-Group herders were of Caucasoid physical type and had non-kinky hair form. They therefore were not the Tutsi- or Maasai-like “Elongated Africans” that Hiernaux had conjectured. On this ancient settlement of Saharan pastoralists, he indicates:
Quite possibly the migration of the Sahara pastoralists reached the Horn and East Africa. Settlements of village farmers, basically neolithic but also using some copper, are known from the plateau of north-western Ethiopia; their material culture shows affinities with that of the C-Group peoples who moved into Nubia from the western desert about 2500 BC. Rock paintings in the eastern parts of Ethiopia and Somalia are reminiscent of those of the eastern Sahara; they depict herdsmen and long-horned, humpless cattle.
Global racial distribution during the Middle Ages (Coon (1982)).
Additionally, in reference to his hypothetical “Elongated African” morphology, Hiernaux writes that “Coon believes that it was first concentrated in the cattle breeders of the neolithic Sahara, whose inhabitants dispersed because of progressive desiccation, giving rise to the herdsmen of East Africa”. However, this passage is rather misleading since Coon never made any such assertion or indeed even acknowledge an “Elongated African” entity. He instead maintained throughout his career that the first Afro-Asiatic-speaking settlers in the Horn region and environs were “Mediterranean Caucasoids”, who were ancestrally related to the “Hamitic” predynastic Egyptians and ancient Libyans. In 1982, Coon published Racial Adaptations, an update of his earlier Races: A Study of the Problems of Race Formation in Man from 1950. The latter was among the first scientific works to explore biological adaptation in human populations. Although Racial Adaptations was released after Hiernaux’s own book on the subject, Coon was not swayed by his colleague’s theories, as the enclosed racial distribution map makes clear.
Ultimately, Coon regarded the Middle East rather than the Sahara or East Africa as the likeliest area of origin of the Mediterranean stock. He explains it thusly in his The Races of Europe:
whatever the date of these specimens in years, East Africa was not, in Upper Palaeolithic times, the center of Mediterranean racial evolution. Neither, it would appear, was the Sahara; so far the archaeologists have not found evidences of the Upper Palaeolithic Capsian culture in the central zone of the desert itself, where there is at present a gap between the Levalloisian and what appears to be an early, arrow-chipping Neolithic in Capsian tradition. The Capsian apparently came to North Africa from the east, and the mid-Sahara may have served even during Pleistocene times as a dividing line between white and negroid humanity, just as it does today.
(For further details, see Did the Capsians of North Africa descend from the Iberomaurusians?, Have the Kiffians and Tenerians of the Gobero been genetically analysed yet?, and What are the genetic affinities of the Aterians?.)
Besides the foregoing, there are several additional key indications that Coon, as opposed to Hiernaux, is factually closer to the mark about the identity and provenance of the first Afro-Asiatic-speaking herders in the Horn region.
Firstly, ecological studies have established that the climate and lake levels in the Horn fluctuated during the Holocene (~10,000-6,000 ybp). The highlands were initially frigid and uninhabitable. This early wet phase was characterized by a glacial retreat, with a corresponding rise in the water levels of the Ethiopian and East African Rift Valley lakes as the snow melted. These lake levels would remain high throughout the Holocene, thereby precluding human occupation. As the climate gradually changed and the lake levels dropped, vegetation favoring pastoralism would begin to grow. Only after 5,000 ypb, with the start of a new dry phase, would the first Neolithic pastoralists arrive from the Sahara. Oba (2013) writes:
In the region of the Horn of Africa, both the highlands and the lowlands experienced climatic fluctuations. It is possible that during the early phase of the wet Holocene climate, the highlands were too cold for human settlement. Finnevan reports: “The …wet phase was accompanied by a glacial retreat” from the highlands in both Ethiopia and East Africa. Between 12,000 and 5,000 years ago snow retreat resulted in a rise in the levels of the Ethiopian and the East African Rift Valley lakes. At the regional level, the fluctuation in the East African Rift Valley lakes was synchronized, suggesting close links between changes in lake levels and regional climate..[…]
In East Africa, during the full Holocene period from around 10000 through 6000 BP, the high water levels in the Rift Valley lakes did not allow human settlement in the moist environments. The wet and the dry phases alternated between 4450 BP and 2700 BP, with lake levels dropping, followed by recession of the sub-humid vegetation towards arid conditions in the lowlands and the retreat of forest cover in higher elevations. From the Afar region of Ethiopia, lakebed changes depict the different phases of the paleoclimate. Grove, referring to carbon dating of the sediment cores of the lakes (i.e. Shala, Abiyata and Langano), confirms higher lake levels around 9200 BP, which retreated to the present levels by 4000 BP. The decline in the lake levels is an indication that climate conditions had shifted to a dry phase ca. 5000 BP[…]
The Neolithic herders arrived from the Sahara. The period between 5000 BP and 3000 BP represents the mid-Holocene, when aridity induced migrations of prehistoric pastoralists into the Horn of Africa.
The Horn was thus considerably less arid in the past than it is now. This perhaps should be obvious given the presence of certain fauna on the region’s ancient cave paintings, which can only survive in habitats where there is abundant vegetation (e.g. the ibex). Indeed, the very type of cattle depicted in the earliest pastoral rock art, the long-horned humpless cattle or Bos taurus, itself serves as a reliable indicator of the kind of climate that prevailed at the time. Oba explains:
During the Neolithic period called Bovidian, the styles of the rock art enable various interpretations based on the depiction of livestock species such as ovids (goats) and capris (sheep) and Bovin (cattle). By about 8000 BP, the depictions in the rock art were mainly of ovicaprids and cattle. The rock artists were predominantly concerned with the sources of their food during each period. Thus, the shift from the Paleolithic phase, which represented the hunter foragers, to Neolithic developments, which represented pastoralists, show shifts in food sources and society responses to climate and environmental changes. The domestication of Bos primigenius (primitive cow) expanded into the Nile valley about 7000 BP; this was the ancestor of the cattle breeds Bos taurus (the humpless long horned) and Bos indicus (the humped short horned zebu). Bos taurus was common during the humid phase of the climate, while Bos indicus became prominent with increased aridity. The depictions show that the rock art serves as a slow cinematic sequence of pastoral evolution to climate change, even when data from a single site is scrutinized. In particular, the gradual disappearance of Bos taurus and the popularity of Bos indicus in rock art sites strongly imply environmental adaptation since Bos indicus had better thirst tolerance than its predecessor. The camel was introduced later than the bovids during periods of increased aridity. From the rock art across the region archaeologists have pieced together the regional migration patterns of the rock art pastoralists.
An anthropomorphic Puntite crocodile figurine excavated in northern Somalia (Ibrahim (2013)).
Recent archaeological excavations in northern Somalia, during which the first actual artifacts from the Land of Punt were unearthed (as explained in detail here), provide a similar indication. Among the discovered objects was an anthropomorphic crocodile figurine of apparent religious significance. This suggests that the ancient Puntites inhabited a more lush environment than today because crocodiles usually congregate in freshwater biotopes.
Hiernaux was therefore mistaken when he proposed that the Horn was once dry enough for the “Caucasoid” morphology to have, at least in part, evolved in situ there. Since the region would not become arid until relatively recently, the ancient pastoralists with that phenotype had clearly arrived from elsewhere.
Secondly, the pastoral-themed cave paintings at Laas Geel, Dhambalin and Karin Hegane in northern Somalia and at other sites in Northeast Africa are in a singular “Ethiopian-Arabian” style. The earliest examples of this rock art tradition are found in Saudi Arabia, and they date back to around 4,500 BCE. On this point, Rodolfo Fattovich of the Università l’Orientale di Napoli, who in 2001 led archaeological excavations at Mersa/Wadi Gawasis on the Red Sea (the ancient Egyptian port of Saww used for commercial expeditions to the Land of Punt), notes:
Finally, a strong interregional interaction, if not a proper movement of people from Arabia to Ethiopia, in the 3rd-2nd millennia B.C. is supported by the occurrence of rock pictures in a typical style in both regions. This style is characterized by painted or engraved bovines with the body in profile and the head and horns in plan. It is recorded as «Jubba Style» in northern Saudi Arabia, «Dahthamani Style» in central Arabia, and «Karora Style» or «Ethiopian-Arabian Style» in the Horn of Africa.
The earliest evidence of this style has been traced in northern Saudi Arabia and dated to the mid-5th millennium B.C. By the mid-3rd millennium B.C., most likely, it spread to the southern Hidjaz (Saudi Arabia), northern Hararge (eastern Ethiopia), and – along the Ethiopian Rift Valley – to Sidamo (southern Ethiopia). Then, figures dating back to the 2nd-early lst millennia B.C. have been recorded at Jebel Qara (central Arabia), Hararge (eastern Ethiopia), northern Somalia, and Eritrea. By the lst millennium B.C. this style spread from Eritrea to Nubia, southern Upper Egypt and the Sahara.
Thirdly, the pre-Islamic religion of the Cushitic Somalis is related to that adhered to in the pre-Islamic Middle East. Mukhtar (2003) notes that the Somali word for spirit, hoobal, is derived “from the Arabic hubal, a pre-Islamic Arabian god whose effigy was once venerated in the Ka’ba in Mecca.” Ali (2006) similarly indicates:
the Biblical YAHWE (later turned Yehova and Jehova) was evidently the same as the Somali YAHU – traditionally invoked to ward off evil or danger. While the Cananite god ‘Pal’ was still present in Somalia in the same sense in one or two words, the ancient Aramaic name for the almighty, EBBE, was to this day the most commonly used names for God besides the Islamic ‘Allah’. The Biblical TUBAN-CAIN, whose profession was to make instruments (Genesis 4:22) was obviously a Greek mispronunciation of TUMAL, the Somali iron-monger.
Fourthly, Tishkoff et al. (2010) reports that the most common lactase persistence allele carried by East Cushitic individuals is G-13915. This variant peaks among Somalis at a frequency of 50%. It is closely associated with camel pastoralism and is believed to have evolved in the Arabian peninsula, where the mutation is today prevalent. To this end, Fattovich (1997) asserts that “an introduction of camel from Arabia into the Horn at an early time, perhaps between 2500 and 1500 B.C., might be also supported by the use of an archaic South Arabian type of camel saddle in Somalia and Socotra.”
Neolithic cave paintings at Laas Geel depicting long-horned humpless cattle (the “Hamitic Longhorn”).
Fifthly, analysis by Decker et al. (2014) of 134 cattle breeds from around the world has established that the long-horned humpless cattle depicted in the Neolithic cave paintings of Northeast Africa — cattle that is often colloquially referred to as the “Hamitic Longhorn” because of its ubiquity in Ancient Egypt — is, in fact, related to cattle that was first domesticated in the Middle East at an earlier date (~10,000 ypb). The cattle was therefore brought to Africa by migrating herders, who not long afterwards began drawing the “Ethiopian-Arabian” style rock art in the areas in which they settled:
Geneticist and anthropologists previously suspected that ancient Africans domesticated cattle native to the African continent nearly 10,000 years ago. Now, a team of University of Missouri researchers has completed the genetic history of 134 cattle breeds from around the world. In the process of completing this history, they found that ancient domesticated African cattle originated in the “Fertile Crescent,” a region that covered modern day Iraq, Jordan, Syria and Israel. Lead researcher Jared Decker, an assistant professor of animal science in the MU College of Agriculture, Food and Natural Resources, says the genetics of these African cattle breeds are similar to those of cattle first domesticated in the Middle East nearly 10,000 years ago, proving that those cattle were brought to Africa as farmers migrated south. Those cattle then interbred with wild cattle, or aurochs, which were native to the region, and changed their genetic makeup enough to confuse geneticists[…]
“In many ways, the history of cattle genetics mirrors human history,” Decker said. “In the case of African cattle, anthropologists and geneticists used to suspect that domesticated African cattle were native to the continent, when in fact, they were brought by migrating peoples thousands of years ago. By better understanding the history of the animals we domesticate, we can better understand ourselves.”
Mwai et al. (2015) provide a useful map showing how the cattle likely spread after their initial introduction from the Fertile Crescent:
Lastly, genetic analysis of modern and ancient human populations in Northeast and Northwest Africa also supports Middle Eastern affinities for the continent’s first Afro-Asiatic-speaking settlers. Hodgson et al. (2014) found a West Eurasian ancestral component that defines the Afro-Asiatic speakers in the Horn, with a frequency peak among ethnic Somalis. Dobon et al. (2015) observed an analogous element among Egyptian Copts, Beja and other Afro-Asiatic speakers in the Nile Valley and Ethiopia, as well as among many present-day Nubians. Henn et al. (2012) in turn identified a West Eurasian ancestral component that defines the Afro-Asiatic-speaking populations in the Maghreb, with a frequency peak among Tunisian Berbers. In addition, as explained in detail in the Ancient DNA from Sudan thread, Sirak et al. (2015) noted a similar Middle Eastern affiliation for the ancient DNA of a specimen from the medieval site of Kulubnarti near the Nile. (*N.B. In 2023, researchers from Uppsala University analysed the DNA of Middle Neolithic pastoralists buried at Skhirat, an archaeological site in northern Morocco. The scientists report that the Skhirat individuals bear Levantine ancestry (cf. Simões (2023a); Simões (2023b)). Our genome analysis of modern Cushitic, Ethiosemitic and North Omotic speakers from the Horn of Africa has found that these individuals also carry varying degrees of the Skhirat Levantine ancestry, with a percentage climax among the Cushitic-speaking pastoralist groups. The Cushites of the Pastoral Neolithic and the medieval Nubians likewise harbor this same Levantine ancestry. Hence, the Middle Neolithic specimens from Skhirat, who originated from the Middle East, appear to belong to the Afro-Asiatic-speaking population that first introduced pastoralism to Africa. For further details, see Ancient DNA from Sudan.)
In conclusion, Hiernaux was initially on the right path when he acknowledged an Afro-Asiatic influence of varying degrees on other populations in Africa. Where he went wrong was in allowing unfortunate post-colonial circumstances, both in his place of residence in eastern Central Africa and in academe, to force him to sacrifice empirical truth for political correctness and conjecture. As techniques on how to successfully extract ancient DNA continue to be refined and more old human remains are analyzed, other speculative fallacies will similarly fall by the wayside while robust traditions will instead receive confirmation.
11 Saturday Mar 2023
Tags
A-Group, Afro-Asiatic, ancient DNA, C-Group, Christian Nubia, Kerma, Meroe, Northeast Africa, Savanna Pastoral Neolithic
Linguists often hypothesize that the Sudan area in Northeast Africa may have been the urheimat or “original homeland” of the Afro-Asiatic (Hamitic-Semitic) language family. Certain old cultures in the vicinity — namely, the A-Group, C-Group, Kerma and Meroitic civilizations — also have close ties with that of ancient Egypt and the Cushitic-associated Savanna Pastoral Neolithic. As such, the biological affinities of early populations from this region are of considerable interest to scientists.
Ancient DNA (aDNA) analysis has become increasingly common in Europe and parts of Asia. This is due to the generally temperate climate in these geographical areas, which in turn facilitates the molecular preservation of DNA extracted from ancient human specimens. Conversely, much of the northern half of Africa as well as the Middle East falls under a hot desert climate (BWh) in the global Köppen climate classification:
As a result, the typically hot and arid environmental conditions in these regions have until recently made the collection of ancient DNA difficult.
Neolithic and Meroitic, Post-Meroitic & Christian periods
In 1997, C. Lalueza Fox published the first ancient DNA study on Nubia. He sampled Meroitic period individuals excavated at the Amir-Abdallah site near Abri in northern Sudan, located between the Second and Third Cataracts of the Nile river. The scientist examined the specimens for the Hpa I (np3,592) mitochondrial marker, which occurs at elevated frequencies among most contemporary Sub-Saharan African populations (~69% on average). He notes that therefore “the widespread presence of this marker in the African continent would suggest its ancestral character (Denaro et al. 1981), perhaps attributable to a proto-khoisan stock (Scozzari et al. 1988), prior to the divergence of Khoisan and Negroids (Scozzari et al. 1994).” Of the 15 Meroitic individuals whose DNA Fox was able to successfully amplify, he reports that 4 carried the Hpa I (np3,592) mutation. Hence, while most of the Meroitic samples did not harbor this marker (~73%), a significant minority of these ancient specimens were Hpa I (np3,592) bearers (~27%). This Hpa I (np3,592) percentage is lower than that which Ritte et al. (1993) found among their Ethiopian Jew samples (44.12%), but higher than that which has been observed among Somali individuals (~20%). According to Fox, this suggests that there was some gene flow from Sub-Saharan Africa into the Nile Valley, which served to dilute the Meroitic population’s originally Eurasian mitochondrial base.
In 2009, Hisham Yousif and Muntaser Eltayeb with the University of Khartoum managed to successfully extract ancient DNA from bone samples stored at the Sudan National Museum. The specimens dated from the Neolithic, Meroitic, Post-Meroitic and Christian periods, with the Neolithic specimens belonging to the Kadruka culture:
The researchers observed that the Neolithic Kadruka samples generally belonged to different paternal lineages than the later Meroitic, Post-Meroitic and Christian period samples. Most of the Kadruka individuals were haplogroup A-M13 bearers, whereas the Meroitic, Post-Meroitic and Christian era samples were instead mainly haplogroup DE (haplogroup YAP, which includes the D-M174 and E-M96 clades) and haplogroup F-M89 carriers:
Haplogroups A-M13 was found at high frequencies among Neolithic samples. Haplogroup F-M89 and YAP appeared to be more frequent among Meroitic, Post-Meroitic and Christian periods. Haplogroup B-M60 was not observed in the sample analyze.
Since haplogroup A is today the most common paternal lineage among Nilotic groups in the Nile Valley, the researchers propose that the Neolithic Kadruka population was likely ancestral to present-day Nilotes residing in the area. By contrast, the higher haplogroup E and F frequencies found among the Meroitic, Post-Meroitic (X-Group) and Christian era populations prompted the scientists to assert that these other early Sudanese groups instead had closer ties with communities ancestral to the modern Afro-Asiatic-speaking (Beja, Sudanese Arabs) and Nubian-speaking populations:
In Y-chromosome terms this mean in simplest terms introgression of the YAP insertion (haplogroups E and D), and Eurasian Haplogroups which are defined by F-M89 against a background of haplogroup A-M13. The data analysis of the extant Y-chromosomes suggests that the bulk of genetic diversity appears to be a consequence of recent migrations and demographic events mainly from Asia and Europe, evident in a higher migration rate for speakers of Afro-Asiatic as compared to the Nilo-Saharan family of languages, and a generally higher effective population size for the former.
Osteological analysis of the ancient A-Group culture bearers of Nubia, ancient and modern Nilo-Saharan and Niger-Congo speakers, and modern Cushitic-speaking Somali individuals. Among the modern comparative samples, the A-Group makers cluster with the Somali cohort, at a classification accuracy of 97.8%. This suggests that the A-Group peoples were likely Afro-Asiatic speakers (Becker (2011)).
Although specimens belonging to the A-Group culture of Lower Nubia (which postdates the Neolithic Kadruka culture of Upper Nubia) were not examined, existing skeletal analysis of these individuals suggests that they too were ancestral to the modern Afro-Asiatic-speaking and Nubian groups. This is because the A-Group specimens share close osteological ties with the latter populations, as well as with ancient and contemporary Egyptian samples. Conversely, the A-Group people are markedly unlike the modern Nilotic groups. Becker (2011) compared ancient skeletons belonging to the A-Group makers with those of modern Somali individuals from the Horn of Africa, modern Niger-Congo speakers from eastern and western Africa, modern Nilo-Saharan speakers from South Sudan and the Sahel, ancient Nilo-Saharan specimens excavated from the Wadi Hawar site in Sudan, and ancient Sub-Saharan African specimens exhumed from the Jebel Sahaba site also in Sudan. The scientist observed that the A-Group specimens could only be grouped with the Somali samples and vice versa, and at a high classification accuracy of 97.8%. He ascribes this close affinity to shared non-sub-Saharan African morphological attributes. Becker writes:
All relevant results of the osteological analyses clearly showed that the Wadi Howar sample consisted of the remains of people of biologically sub-Saharan ancestry (see IV.A.9. and V.C.1.h.). Additionally, not a single Wadi Howar individual or group was found to be most similar to either the A-Group or the Somali sample, the only core comparative samples with appreciable frequencies of biologically non-sub-Saharan morphological characteristics (see IV.D.). Therefore, the Wadi Howar’s prehistoric inhabitants were evidently members of a biologically sub-Saharan population complex.
Altogether, this indicates that the A-Group culture bearers were in all likelihood Afro-Asiatic speakers, having different ancestral origins from the Nilo-Saharan and Niger-Congo-speaking communities (cf. Billy (1981b)).
Inner coffins of the ancient Egyptian aristocrats Nakht-Ankh (right) and Khnum-Nakht (left), dating from the Middle Kingdom (c. 1985-1773 BCE). Forensic analysis of the bodies of these Two Brothers revealed profound anatomical differences, which indicates that the siblings likely had been placed in the wrong coffins. Nakht-Ankh was found to have had a “Caucasoid” skull of a delicate character, an orthognathous craniofacial profile, a slender build, an olive complexion (based on examination of residual skin tissue), and soft-textured dark brown hair, whereas Khnum-Nakht had a “Negroid” skull of a robust character, a considerably prognathous craniofacial profile, and a dark complexion (cf. Murray (1910); David (2007); Riggs (2014); Forshaw (2019)). Along with some hieroglyphic evidence, this marked physical divergence further suggested to researchers that the brothers shared a mother but had different fathers (Drosou et al. (2018); Forshaw (2019)). Ancient DNA analysis confirmed these suspicions: the men bore different paternal markers but belonged to the same maternal lineage, the M1a1 haplogroup, which is a signature mtDNA clade of the Afro-Asiatic-speaking populations inhabiting the Nile Valley and Horn of Africa (Drosou et al. (2018)). While Khnum-Nakht’s specific Y-DNA haplogroup could not be identified, Nakht-Ankh was, after some initial uncertainty, successfully assigned to the H2m clade (cf. HaploTree; Open Genomes). The H2 patrilineage has been tentatively detected in Christian-era Nubian individuals (Yousif and Eltayeb (2009); see discussion below), consistent with the proposed ancestral ties between the “red” Nubians of the ancient Egyptian monuments and the Egyptians themselves.
After the publication of Yousif and Eltayeb (2009)’s study, it was recognized that the M282 mutation, which at the time was classified under F-M89, had been misclassified. M282 was subsequently reassigned to its proper lineage, the paternal haplogroup H; specifically, as the defining mutation of the H2 sublineage. In the archaeogenetic record, H2 mainly has been found among Neolithic European specimens (e.g. Cassidy et al. (2020)). The oldest example of this subhaplogroup in Africa has been detected in an ancient Egyptian individual, the Middle Kingdom nobleman Nakht-Ankh (HaploTree; Open Genomes). Ergo, if the instances of F-M89 which Yousif and Eltayeb (2009) identified among their Christian-era Nubian samples belong to the M282 mutation, they probably were indeed inherited from Afro-Asiatic-speaking ancestors. However, if these mutations belong to other F-M89 derivatives, they were likely instead inherited from Neolithic Europeans. This is because, as of 2023, the oldest example of the F-M89 paternal clade has been observed among Early Neolithic specimens excavated at Kahf Taht Al Ghar, an archaeological site located in the Maghreb region in northwestern Africa. These ancient individuals were reported to bear Anatolian Neolithic ancestry, which today peaks among Sardinians and other Southern Europeans.
(*N.B. As of 2023, three distinct Eurasian ancestries have been observed among ancient Afro-Asiatic-speaking individuals in Africa: a Levantine Natufian-associated component, which has been detected in an ancient Egyptian individual from Nuerat, who lived during the Old Kingdom (c. 2868-2492 BCE); a Levantine Skhirat-associated component, which has been detected among Middle Neolithic pastoralists buried at Skhirat, an archaeological site in northern Morocco; and a European Neolithic-associated component, which has been detected among Early Neolithic agriculturalists excavated at the Kahf Taht Al Ghar archaeological site in the Maghreb. It is not yet clear which of these Eurasian ancestries, if any, the A-Group, C-Group and Meroitic culture bearers of Nubia carried because these ancient individuals have not yet been genomically analysed. However, it is possible that the C-Group peoples, who are among the first attested pastoralists in the Nile Valley, may have borne significant Levantine Skhirat-related ancestry. Besides these three primary Eurasian ancestries, it is likely that certain ancient specimens in Africa linked to early Afro-Asiatic speakers also carried European Steppe and East Asian ancestries. We have detected the latter ancestral elements in genome analysis of ancient and modern Afro-Asiatic-speaking individuals. For more details on this, refer to Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa. Also see our study Autosomal STR Analysis of the Ancient Egyptian Amarna Royal Family, Pharaoh Ramesses III, and Unknown Man E (Prince Pentawere.)
Mesolithic, Neolithic and MXCH (Meroitic, Post-Meroitic & Christian periods)
In 2013, Francigny Vincent and colleagues attempted to extract ancient DNA from skeletal remains associated with the Ancient Kerma (2500-2050 BCE), C-Group (2300-1600 BCE), Meroitic (350 BCE-350 CE), and Christian (550-1500 CE) cultural periods of Nubian history. However, the scientists were unsuccessful in this endeavour, owing to excessive DNA degradation of the examined specimens.
The following year, in 2014, a genetic research team led by Kendra Sirak of Emory University’s Department of Anthropology managed to successfully extract endogenous DNA from other ancient individuals from Sudan. The analyzed specimens consisted of six samples spanning from the Mesolithic through to the Meroitic, Post-Meroitic/X-Group and Christian periods (MXCH) in Wadi Halfa, Lower Nubia/northern Sudan:
As of August 2015, the researchers have only published the aDNA results of one of these ancient specimens, a Christian-period infant dubbed NUB04b in the sample set:
NUB04b was found to belong to the mtDNA haplogroup L5 (formerly known as L1e), a small maternal clade that today is most common in East Africa:
To gauge NUB04b’s autosomal DNA affinities, the researchers also ran a principle component analysis, which compared 300 of the specimen’s single nucleotide polymorphisms (SNPs) spatially against those of various modern individuals from the Omni 2.5 dataset. For reasons I shall explain in a future post, such biogeographical testing/admixture testing, including SNP genotyping and whole genome analysis, is extremely flawed with regard to present-day populations alone (this in large part has to do with the difference between the genealogical tree and the genetic tree). It can, however, be informative for ancient individuals when they are compared against modern individuals. Unfortunately, NUB04b’s SNP genotyping was apparently compromised by the presence of some extraneous DNA:
The sample is found to be located between European and African populations, suggesting the presence of some European contaminants extracted together with endogenous DNA.
So what can be gleaned from this ancient DNA analysis? For starters, certain mtDNA haplogroups that today are primarily concentrated below (or above) the Sahara may not always have been. Such appears to be the case with the L5 clade to which the NUB04b specimen apparently belongs. More importantly, these lineages may originally have had quite different population affinities than their present-day distribution would suggest.
As the earliest ancient specimen so far found to carry haplogroup L5, it would be interesting to know just what are NUB04b’s general phenotypic affinities. To this end, the researchers provide a graphic illustrating the marked difference in skull form between the later Meroitic, Post-Meroitic/X-Group and Christian (MXCH) period inhabitants of Wadi Halfa and the earlier Mesolithic period dwellers:
The Mesolithic period inhabitants were clearly a more robust people, with a rugged osteological framework. Comparative craniofacial and anthropometric analyses of these ancient specimens versus later MXCH samples, modern Nubians and ancient and modern Egyptian samples have been conducted by various anthropologists. These analyses have consistently grouped the Mesolithic Wadi Halfa crania with those of present-day “Negroid” populations, while the MXCH and modern Nubian samples have instead typically clustered with Egyptians and other surrounding populations with a generally “Caucasoid” craniofacial pattern.
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. Notice how the Egyptian cranium closely resembles the MXCH skull above, in accordance with the archaeogenetic evidence (discussed below) indicating that Egyptian-related peoples inhabited areas of Nubia (Elliot Smith and Jones (1910)).
In April 2016, Sirak and colleagues published an abstract on morphological and genetic analysis, which they conducted on ancient Nubian individuals. The sample set consisted of 150 adults excavated in Upper and Lower Nubia, with the specimens dating from the Mesolithic through to the Christian periods. The researchers observed a major difference in craniofacial form between the Mesolithic and Neolithic individuals, in accordance with their haplogroup and SNP analysis. While the briefing does not specify if the Neolithic samples belonged to the earlier Kadruka culture of Upper Nubia or the ensuing A-Group civilization of Lower Nubia, we can assume from the morphological patterns that they were associated with the latter cultural sphere. All in all, the data further underlines the distinct ancestral origins of the Mesolithic period inhabitants of Nubia versus the later A-Group, C-Group, Kerma, Meroitic, X-Group (Post-Meroitic) and Christian era populations:
The biological history of the occupants of Nubia throughout the Holocene remains a topic of debate; specifically, there is some tendency to assume a discontinuous history of occupation and cultural development based on traditional anthropological and archaeological evidence. Population movements, a shift in subsistence strategies and in-situ evolution have all been used to explain the biological variation observed in Nubian remains, particularly in the cranium. Here we investigate craniofacial and mandibular shape patterns, as well as ancient DNA variation, among populations from Upper and Lower Nubia spanning 12,000 years and reflecting a transition from hunting-gathering to intensive farming.
Our sample includes 150 adult specimens from six archaeological sites along the Nile River in Egypt and Sudan that belong to eight chrono-cultural groups spanning from the Late Mesolithic through the Christian periods. All individuals were digitized with a surface scanner, then 397 and 120 three-dimensional landmarks and semilandmarks respectively were extracted on skulls and mandibles of each specimen. Landmark configurations were subjected to generalized Procrustes analysis, tangent space projection, principal component analysis, discriminant analysis, and MANOVA. Endogenous DNA was extracted and sequenced from a subsample in order to explore variation in Nubian population genetics throughout time and space using informative SNPs and haplogroup determination.
Our results highlight a strong distinction between Mesolithic and the Neolithic samples. Craniofacial patterns underline the importance of gene flow and give some support to the hypothesis of regional continuity among more recent groups, while patterns of mandibular morphology show high correlation with subsistence strategy.
Approximate location of the Kadruka archaeological site, situated north of Dongola and south of Kerma along the Nile River in Upper Nubia (Geographic.org). During the early Neolithic period, Kadruka was inhabited by carriers of the archaic African haplogroup A, a paternal lineage that is common today among Nilotic groups in Sudan (cf. Yousif and Eltayeb (2009)‘s ancient DNA analysis). Subsequent population movements in the late Neolithic/Bronze Ages would introduce the first Afro-Asiatic speakers into the area. These new arrivals were instead found to be genetically indistinguishable from the Cushitic settlers of the Early Pastoral Neolithic, supporting the theory that these pastoralists had migrated southward to eastern Africa from a center in the Nile Valley (Wang et al. (2022)).
Wang et al. (2022) sequenced DNA from an ancient individual excavated at Kadruka, Upper Nubia. The specimen is 4000 years old and is thus coeval with the Mota hunter-gatherer from southwestern Ethiopia, as well as with Djehutynakht, the oldest Egyptian individual thus far analyzed (who belongs to the mtDNA haplogroup U5b2b5), and the Afro-Asiatic-speaking Kerma and C-Group populations of Nubia. Unsurprisingly, considering the time period in question, the researchers observed that their Middle Kingdom Kadruka specimen was genetically indistinguishable from the Cushitic settlers of the Early Pastoral Neolithic in the Great Lakes region. This reflects the spread of Afro-Asiatic speakers into Upper Nubia, an area which, as seen above (cf. Yousif and Eltayeb (2009)), was formerly in early Neolithic times principally inhabited by haplogroup A carriers.
Christian period
In 2015, Sirak et al. published another conference paper on ancient DNA from the Sudan region. The bone samples this time were collected from the medieval site of Kulubnarti, located between the Second Cataract and Dal Cataract beside the Nile. Three individuals in total, they all dated from the early Christian era i.e., the historical period when the Alodia/Alwah, Nobatia and Makuria kingdoms were extant:
The scientists then ran an analysis on the sample that yielded the highest endogenous DNA, KulR17. Like NUB04b from Wadi Halfa, this early Kulubnarti inhabitant also happened to be an infant. 523 of KulR17’s SNPs were compared against various global reference samples from the Human Genome Diversity Project dataset. However, there was apparently no DNA contamination this time. The researchers instead remarked that “the authenticity of KulR17 seems to show that even in hot and arid climatic conditions it is possible to extract endogenous DNA from archaeological samples.”
KulR17’s biogeographical ancestry turned out to be most similar to that of modern populations in the Middle East and environs:
From this analysis, the geographic ancestry of this individual was estimated to be closer to Middle Eastern/and Central and South Asian populations than to any African population.
Medieval illustration of a Nubian king.
This finding is consistent with the Coptic ancestral component that Dobon et al. (2015) observed to be the defining element among Egyptian Copts and other Afro-Asiatic speakers in the Nile Valley and Ethiopia, as well as among many present-day Nubians. Hodgson et al. (2014) found an analogous West Eurasian ancestral component among Afro-Asiatic speakers in the Horn region, with a frequency peak among ethnic Somalis. Since it is unlikely that there was a population replacement among Nubians in the intervening centuries after the medieval Christian period, we can safely assume that the Coptic/Ethio-Somali ancestral component that defines modern Nubians is the same West Eurasian-affiliated ancestral component that defines the KulR17 specimen from Kulubnarti.
Additionally, this aDNA result is in keeping with the aforenoted osteological affinities of MXCH period skeletons from Sudan, as well as medieval iconography of Nubian royalty (see illustration on right). In future posts, we shall see that this finding is also in alignment with the linguistic affiliations of the earlier C-Group and Kerma Afro-Asiatic-speaking cultures, and possibly also that of the Meroitic civilization.
*Update #1
Location of Meroë and other key sites in Nubia (Khalil and Miller (1996)).
Exciting news, folks! It took a few years, but we finally have some ancient DNA analysis on pre-Christian Nubia (better late than never).
In early 2020, Yahia Mehdi Seddik Cherifi of the Laboratoire d’Anthropométrie, du Patrimoine et d’Archéologie and Selma Amrani of the Institute of Archaeology at the University of Algiers released a study on the first mitochondrial DNA extracted from individuals belonging to the X-Group (Post-Meroitic) culture. This civilization, also known as the Ballanean Culture, existed in Upper Nubia between 300-600 CE, arising after the end of the Meroitic Kingdom. The X-Group Nubians were culturally different from the preceding Meroites. They also, apparently, did not speak the Meroitic language. They are thought to have spoken instead the Nobiin or Nubian language, a Nilo-Saharan tongue, which most Nubians today speak.
(*N.B. Nubians are a distinct people from the Nuba/Noba. Nubians today speak Nobiin, a Nilo-Saharan language which they are believed to have adopted. Their Meroitic ancestors appear to have originally spoken an Afro-Asiatic language like the neighboring Egyptians (see discussion below). Nubians inhabit the area between central Sudan and southern Egypt. They are thought to descend from the Egyptian-related inhabitants of ancient Nubia (the “red Nubians”) because they share close physical, genetic and cultural ties with local Afro-Asiatic-speaking populations. By contrast, the Nuba/Noba today speak Kordofanian languages, which belong to the Niger-Congo family. They inhabit the Nuba Mountains/Kordofan Mountains/Nuba Hills to the south of the Nubians. The Noba/Nuba are thought to descend from the Nilotic-related inhabitants of ancient Nubia (the “black Nubians”) because they share close physical, genetic and cultural ties with local Nilo-Saharan-speaking populations.)
The researchers obtained mitochondrial sequences from eight X-Group skeletons buried at the Missiminia Necropolis, as well as from two Meroitic (including Late Meroitic) specimens and one Christian period individual interred at the site. They observed that the X-Group representatives carried mtDNA haplogroups that are nowadays associated with both Africa and Asia, including the X (twice), T1a, N, L3b, L3e, L2 and L1b clades. By contrast, both of the Meroitic samples belonged to the H2 lineage and the Christian era specimen was assigned to the W1 haplogroup, maternal clades that are of non-African origin. The scientists therefore postulate that there was in Nubia an “influx of sub-Saharan African ancestry after the Meroitic Period.” This is indeed suggested by the presence of the L1b and L2 haplogroups in the X-Group/Post-Meroitic sample set. It is also supported by the fact that most modern Sudanese “Arabs” (Gaalien, Meseria, Arakien) carry mtDNA derivatives of the L macroclade (Hassan et al. (2009)). However, the L3 subclades are not necessarily indicative of Sub-Saharan introgression since the earliest occurrence of the L3 macrohaplogroup has, in fact, been found among specimens belonging to the Pre-Pottery Neolithic culture of the Levant (cf. Fernández et al. (2014)). Babalini et al. (2002) and Yatsishina et al. (2021) also each detected the L3 macroclade in, respectively, an ancient Libyan individual excavated in the Fezzan (dated to c. 3000 to 1500 BCE) and an ancient Egyptian mummy examined at the Kurchatov Institute. Additionally, Farrell et al. (2013) identified certain L3-rich areas in the Arabian Peninsula, where the local populations do not show any apparent Sub-Saharan genomic influence. Ergo, some of the L3 sublineages borne by the ancient X-Group individuals as well as many modern Afro-Asiatic and Nubian speakers in Africa could actually have been imported from the Middle East. (*N.B. For the latest evidence on the suggested non-African origin of the mtDNA macrohaplogroup L3, see Cabrera (2022) and Cabrera et al. (2017).)
mtDNA haplogroups of ancient X-Group (MIS-83, MIS-93, MIS-3/1, MIS-C, MIS-171, MIS-309B, MIS-466, MIS-474), Meroitic (MIS-TM, MIS-TMT), and Christian (MIS-TC) period specimens buried at Missiminia, Upper Nubia (Cherifi and Amrani (2020)).
Now that we have both Y-DNA and mtDNA data on the X-Group/Post-Meroitic population, we can try and ascertain the prevailing direction of gene flow into this ancient community. More specifically, we can find out if:
1) The X-Group was originally a Nilo-Saharan-speaking population related to Nilotes, which later interbred with Afro-Asiatic speakers.
or
2) The X-Group was originally an Afro-Asiatic-speaking population related to Egyptians, which later interbred with Nilo-Saharan speakers and adopted from the latter the Nubian language.
mtDNA haplogroups of modern Nubians and Afro-Asiatic, Nilo-Saharan and Bantu-speaking populations in East Africa. 50%-60% of the Nubian and Afro-Asiatic-speaking individuals belong to maternal lineages derived from the Eurasian M and N macroclades; the remainder carry L(xM,N) haplogroups primarily consisting of the Pre-Pottery Neolithic-associated L3 clade (Non (2010)).
The uniparental markers would appear to favor scenario #2 since most of the X-Group individuals thus far analyzed (2 out of 3 samples or 66.67%) belong to paternal lineages derived from the YAP (DE) clade, like the preceding Meroites and most other neighboring Afro-Asiatic speakers. A minority (1 out of 3 samples or 33.33%) were assigned to haplogroup A, which is the modal patrilineal clade among the ancient and modern Nilotes of the Nile Valley (see Yousif and Eltayeb (2009)’s analysis above). If we assume that the L3 mtDNA sublineages that the X-Group population bears are of Middle Eastern extraction, this implies that most of the Post-Meroitic individuals had Afro-Asiatic-affiliated maternal lineages. This would, in turn, suggest that the Sub-Saharan influence primarily entered the X-Group population through assimilation of Nilotic men. In further support of this male-mediated admixture, Hrdy (1978), who analyzed the hair morphology of Meroitic and X-Group mummies buried at Semna South in Sudanese Nubia, reports that:
Although there is not a consistent statistically significant difference between the X-group and Meroitic samples, it is interesting that the X-group sample, especially the males, had higher curling variables, indicating more of an African element. [*N.B. Hrdy’s African comparison sample consists of modern individuals from Kenya, Uganda, Mozambique and Zambia, who generally have similar hair texture as the Nilotes of the Nile Valley; this “East Africa” sample was first used in his 1973 study.]
Distribution of blond, brown, black and red pigmentation in hair samples from Meroitic, X-Group/Post-Meroitic and Christian era mummies buried at Semna, Sudanese Nubia. Most of the ancient specimens have brown hair. Also notice how reddish and blond hair are most common during the Meroitic period and gradually decline in the later X-Group/Post-Meroitic and Christian epochs, likely due to foreign contacts (Hrdy (1978)). (*N.B. Lazaridis et al. (2022) analyzed ancient Levantine individuals dating from the Bronze and Chalcolithic Ages and did not observe any examples of red-haired specimens. However, the scientists did find some ancient European samples with red hair. This suggests that the occurrence of red hair among the Meroites and ancient Egyptians is specifically attributable to intermingling with early European populations (cf. Supplementary Materials).)
The Meroites, for their part, show an even more frankly Egyptian uniparental marker profile. According to Yousif and Eltayeb (2009), 3 out of 4 or 75% of their Meroitic period samples culled from the Haraz Cemetery carry derivatives of the YAP/DE paternal haplogroup. Furthermore, Sirak et al. (2021) observed a prevalence of the Y125054 subclade of E1b1b-V12 among Christian-era Nubian individuals inhumed at the Kulubnarti site in Sudan. Since the medieval Kulubnarti specimens appear to descend from the Post-Meroitic or X-Group peoples that immediately preceded them, and the Post-Meroitic folks in turn seem to descend from the Meroites, it is likely that the YAP/DE derivatives which Yousif and Eltayeb found among their Meroitic cohort similarly comprise E1b1b sublineages. (*N.B. Trombetta et al. (2015) report that 74.5% of their modern southern Egyptian samples are haplogroup E1b1b-V12 carriers, with 78.7% total E1b1b bearers (cf. Supplementary Table 7). This points to close ties between Egyptians and the Christian-era, Post-Meroitic and Meroitic populations of Nubia.)
On the other hand, Cherifi and Amrani posit that the H2 mitochondrial clade, which both of their sampled Meroitic specimens belong to, was likely of Middle Eastern introduction. However, the H lineage also occurs among modern Nubians and contemporary Afro-Asiatic speakers in the Nile Valley and Horn of Africa (cf. Non (2010), Table 3-3). For example, Comas et al. (1999) note that a number of the Somali individuals they examined bore the Cambridge Reference Sequence (CRS), which belongs to the mtDNA haplogroup H. The lineage is particularly common among the Tuareg Berbers of the southern Maghreb, though here most individuals bear the H1 subclade (Ottoni et al. (2009)); the H clade is also widespread among the Asni, Bouhria and Figuig Berbers of Morocco (Coudray et al. (2009)). Among other ancient specimens, Breidenstein et al. (2018) indicate that all of the medieval Nubians they analysed at the El-Kurru site in Sudan carried Eurasian mtDNA lineages, with 2 of the 5 examined individuals (40%) bearing the H2 subhaplogroup. Similarly, Gad et al. (2020a) report that the 18th Dynasty ancient Egyptian Pharaoh Amenhotep III carries the H2b maternal clade. Drosou et al. (2020), moreover, divulge that the 25th Dynasty (c. 660 BCE) ancient Egyptian mummy Takabuti, an aristocrat from Upper Egypt, belongs to the H4a1 mtDNA sublineage. The scientists indicate that this is “a predominantly European haplogroup.” Accordingly, this clade has been observed in a Neolithic Sardinian specimen (Marcus et al. (2020)). Rayo et al. (2022) likewise state that one of the canopic jars they analyzed, which contains the vital organs of a mummified ancient Egyptian individual, yielded the mitochondrial haplogroup H. The other container the researchers examined held viscera associated with the R0a1 clade, a maternal lineage that is today relatively common among Afro-Asiatic speakers in the Nile Valley, Horn of Africa and Arabian peninsula. (*N.B. Rayo et al. (2022) do not indicate any dates for their canopic jars, so it is unclear which period(s) these mitogenomes belong to.)
These findings only reinforce the Egyptian-like nature of the ancient Meroitic population — an aspect that was already suggested by artwork depicting Meroitic figures with a “Caucasoid” physiognomy, the Meroites’ pyramid-building culture, their ancient Egyptian-derived writing system, and craniometric affinities (cf. Batrawi (1946); Mukherjee (1955); Billy (1981b); Rösing (1990)). As Breidenstein et al. cogently put it, “Nubian genetic makeup before Arab expansion is speculative, inferred from marker typing, or based on non-genetic markers, but thought to be closely related to Egyptians.”
Genetic distance analysis of modern Kababish “Arabs” from Sudan using the Vahaduo Admixture JS program. Most of the examined Sudanese “Arab” individuals show a clear preference for the medieval Kulubnarti Nubian specimens. This supports the view that Sudanese “Arabs” trace most of their proximal or recent ancestry to these earlier Kulubnarti Nubian individuals.
Genome analysis of Sudanese “Arab” individuals and other modern samples from Africa. The examined Sudanese “Arabs” (labeled here “Sudan Nile 2”) derive most of their Eurasian autosomal DNA from an Egypt-related source (light red), followed by a Somali-related source (pink), and then a Saudi Arab-related source (dark red). This is consistent with our genome analysis, which indicates that contemporary Sudanese “Arabs” and Nubians trace most of their proximal or recent ancestry to the medieval Kulubnarti Nubians, who in turn descend from the ancient Egyptian-related X-Group and Meroitic populations of Sudan. The scientists note that “in contrast, inferred Arabic-related admixture in the Beni-Amer involves an African source more closely related to Somalis” (pink component). Hollfelder et al. (2017) detected a similar Cushitic-associated West Eurasian component in their Beja samples, observing that “Beja groups, who generally reside in eastern areas of Sudan close to the sea, show high non-African admixture in all tests[…] these old admixture events into the Beni Amer could be driven by admixture from the Cushitic-speaking populations of the Horn of Africa.” This confirms that the Beja trace most of their proximal or recent ancestry to the Cushites of the Pastoral Neolithic (Bird et al. (2023)).
(*N.B. Sirak et al. (2021), who analysed Christian-era individuals buried at Kulubnarti, similarly assert that “the rise (~300 BCE) and collapse (~350 CE) of the Meroitic Kingdom in Nubia provides a possible historical context for admixture between Egyptian peoples carrying West Eurasian-related ancestry and local Nubians.” However, Sirak et al. argue that modern Nubians don’t descend from the Christian-era Kulubnarti individuals (and, by extension, from the Meroites), but instead derive most of their West Eurasian ancestry from recent Arab Muslim settlers. This claim is rendered doubtful by the fact that a) the researchers utilized a Eurasian-admixed Dinka Nilote sample to arrive at their conclusions (cf. Skoglund et al. (2017); Scheinfeldt et al. (2019)), and b) Dobon et al. (2015), among other scientists, have observed that contemporary Nubians and Sudanese “Arabs” actually share most of their West Eurasian ancestry with Copts and other Afro-Asiatic speakers in Northeast Africa rather than with peninsular Arabs. Thus, it would make more sense to use ancient Egyptian samples, as opposed to Levantine or Arabian samples, as a proxy for the West Eurasian ancestry in modern Nubians; especially since, by Sirak et al.’s own admission, the ancient Egyptian specimens that have thus far been sequenced have just ~5% Sub-Saharan admixture (similar to the small amount of Eurasian admixture that has been detected in the much-used Mota sample) and are the best-fitting surrogate for the Kulubnarti individuals’ West Eurasian ancestry.)
Craniometric analysis of ancient and modern populations from North Africa, the Horn of Africa, and the Middle East. Among the ancient samples, the predynastic Egyptian Badarians cluster nearest to the post-Neolithic populations of Nubia (C-Group, Kerma, Meroitic, X-Group, Philae) and the ancient Egyptians excavated at Deir el-Medineh, Upper Egypt. Among the recent samples, the Badarians cluster nearest to the Galla & Somali lumped sample (Rösing (1990)).
Hrdy (1978) likewise observed, with respect to the Meroitic and X-Group hair samples he examined, that:
In component I, which is heavily loaded on general curling variables and scale count, the total sample centroid was significantly different from European and African samples, though it was definitely more European than African.
Estimated non-African ancestry proportions among Afro-Asiatic-speaking groups of the Horn region. The analysed Cushitic and Ethiosemitic-speaking individuals have a predominant non-African ancestry, averaging nearly 70% (right-most column) (Hodgson et al. (2014), Supplementary Text S1). This is close to the estimate for total non-African/Eurasian ancestry that we found in our genome analysis of these populations. (*N.B. Hodgson et al. (2014)’s Somalia sample was taken from Pagani et al. (2012). Pagani et al. in turn indicate that their Somalia sample was collected by their co-author S. Qasim Mehdi (S. Q. M.) for the earlier Pakistan-based study Li et al. (2007). According to the D.I.P., most Somalis here originate from the Mogadishu area in south-central Somalia. As such, Hodgson’s/Pagani’s Somalia sample primarily consists of Hawiye Somali individuals.)
As with the Meroites of Nubia, the anatomist Grafton Elliot Smith similarly remarks that most of the predynastic Egyptian specimens that he forensically analyzed — which number in the thousands, exponentially more than any other Egyptologist — had soft-textured wavy, straight or curly hair of a very dark brown or black hue:
The hair of the Proto-Egyptian was precisely similar to that of the brunet South European or Iberian people of the present day. It was a very dark brown or black colour, wavy or almost straight, and sometimes curly ; but it presented no resemblance whatever to the so-called “woolly” appearance and peppercorn-like arrangement of the Negro’s hair.
Mummy of the ancient Egyptian Queen Tiye (1389-1338 BCE), “The Elder Lady”, with her natural wavy red hair still attached. Forensic examination of ancient Egyptian specimens indicates that a significant minority had red hair. However, most predynastic Egyptian individuals possessed dark brown or black hair, like the related Meroites (cf. Elliot Smith (1911)).
Mural from the Tomb of Amenemonet at Luxor, Egypt, depicting the ancient Egyptian Pharaoh Amenhotep III and his wife Queen Tiye (Dreamstime).
This also brings to the fore the question of the Meroitic language’s origin. Is it an Afro-Asiatic idiom, as Kirsty Rowan (2006) contends? Or is it, as Claude Rilly (2010) argues, a Nilo-Saharan tongue, which we may presume an Egyptian-related, originally Afro-Asiatic-speaking population later adopted? Either hypothesis is plausible in view of the existing ancient DNA and morphological analyses.
A Meroitic statuette of a bound Noba ruler (50-1 BCE). The figurine has an engraving on it in the Meroitic language, which reads: qo qore Nob-o-l-o (“this one is the king of the Noba”). This inscription suggests that the Meroites were an ancestrally distinct people from the Kordofanian-speaking Noba. It, in turn, lends credence to the idea that either the Meroitic language belongs to the Afro-Asiatic family or the Meroites were originally Afro-Asiatic speakers who later adopted a Nilo-Saharan tongue (Donsmaps).
The Necropolis at Meroë has over 200 pyramids, exceeding in number those erected in ancient Egypt (Sacred Sites).
In April 2020, Abigail Breidenstein of the Institute of Evolutionary Medicine at the University of Zürich and colleagues published an intriguing abstract on paleogenetics from Nubia. Although very brief, it nicely complements Yousif and Eltayeb’s earlier work and Cherifi and Amrani’s latest analysis. The researchers examined the mitogenomes of specimens dating from the Meroitic period through to the Christian era. They observed that most of the individuals belonged to Eurasian haplogroups (4 out of 6 samples or 66.7%), with the rest carrying maternal lineages derived from Sub-Saharan Africa. This is close to the percentage of Meroitic individuals which Lalueza Fox (1997) found to carry the Hpa I (np3,592) mutation, a Sub-Saharan African mitochondrial marker (see discussion above). From the few details Breidenstein et al. provide, it is, however, uncertain just what the identified haplogroups are or whether they include the Pre-Pottery Neolithic-associated L3 mtDNA clade. Nonetheless, it is interesting to note once more the close ties between the Meroitic, Post-Meroitic and Christian era populations of Nubia and the Afro-Asiatic speakers.
(For further details, see Who were the ancient Nubians?, If Nubians are ancestrally related to Egyptians, why do they now speak a Nilo-Saharan language instead of an Afro-Asiatic language?, Are Sudanese “Arabs” really peninsular Arabs?, and Which Afro-Asiatic-speaking population of the Horn region has the most non-African ancestry?.)
*Update #2
Using the Vahaduo Admixture JS program, genome analysis of Christian-era individuals excavated at the Kulubnarti site in Sudan indicates that they bear a predominant non-African ancestry (over 70%), consisting of majority West Eurasian elements (ancient Egyptian, European-related Steppe, and Levantine Natufian components) and a minority East Eurasian element (East Asian component). Furthermore, these ancient specimens have minor Sub-Saharan African admixture (under 27%) and a minute North African Iberomaurusian/Taforalt admixture (under 3%). The Cushites of the Pastoral Neolithic have the same ancestral composition, as do modern Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn region and Kababish “Arabs” from Sudan. Contemporary Muslim Egyptians and Libyans also share a similar ancestral makeup, but experienced some extra recent gene flow from western Asia. By contrast, Coptic Egyptians and northern Egyptians from Cairo and Mansoura appear to descend directly from earlier Dynastic period ancient Egyptians (as represented by the aristocrat Nakht-Ankh discussed above), with little apparent influences from other population sources. Maghrebi groups have an ancestral composition comparable to that of Libyans, with greater Iberomaurusian ancestry (~29% on average) as well as a significant Anatolian Neolithic admixture (~23%), especially in northern coastal areas opposite Iberia.
(*N.B. This ~70% Eurasian ancestry estimate for Cushitic, Ethiosemitic and North Omotic-speaking individuals of the Horn of Africa is also roughly comparable to that of Hodgson et al. (2014) (see Supplementary Text S1) and Dobon et al. (2015) (viz. “the main component (~70%) is that detected in North Africa and Middle East”).)
For the full details, refer to Genetic affinities of the Cushitic, Ethiosemitic and North Omotic-speaking populations of the Horn of Africa.
*Update #3
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. For starters, in her more detailed 2023 dissertation, the lead researcher Luciana G. Simões indicates that the Middle Neolithic individuals from Skhirat harbored 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].
Furthermore, two of the three examined Skhirat individuals were found to carry the paternal haplogroup T, and one bore the mtDNA haplogroup M1a1b. As discussed in greater depth on Ancient DNA from Ethiopia, 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). This 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 the Middle Neolithic Skhirat individuals. These ancient pastoralists primarily bear the T paternal clade and also carry the M1 maternal lineage (Simões (2023a)). These Eurasian haplogroups are signature uniparental markers of the contemporary Afro-Asiatic-speaking populations in the Horn and Nile Valley.
Taken together, the above appears to corroborate the traditional interpretation — supported by genetic analysis of modern African cattle breeds (cf. Decker et al. (2014)), as well as a singular “Ethiopian-Arabian” style of rock art distributed in the Arabian peninsula, Horn of Africa, Nile Valley and Sahara (Fattovich (1997)) — 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, as extensively described above, hints that E1b1b-V32 may have been first brought to the Horn from the Nile Valley during the Meroitic period. A later such expansion of E1b1b-V32 over a haplogroup T substrate is also supported by genome analyses of modern Afro-Asiatic-speaking individuals of the Horn. Osman and Jonasson (2022) report that Northern Somalis born in the northeastern Puntland region of Somalia (who mostly belong to the Majerteen Darod clan) carry a predominant Eurasian ancestry, which on average is comparable in frequency to that borne by North African individuals (cf. Fig. S1). Although these Northern Somali individuals have not been examined for Y-DNA, Trombetta et al. (2015) report that the Afro-Asiatic-speaking Eritrean samples in their dataset mostly belong to the E1b1b lineage, with many falling under the V32 subclade (observed total E1b1b percentages for Afro-Asiatic-speaking Eritreans: Tigre=100%; Saho=98.9%; Tigray-Tigrinya=71.9%; cf. Supplementary Table 7). This is instructive because, after Northern Somalis from Puntland, Cushitic and Ethiosemitic-speaking Eritreans have the next highest reported frequency of Eurasian ancestry in the Horn of Africa (e.g. refer to the Tigray-Tigrinya samples in Hodgson et al. (2014), Supplementary Text S1; Llorente et al. (2015)). Altogether, this suggests that there indeed was a recent migration into the Horn from the Nile Valley, which brought with it an additional layer of Eurasian ancestry that is still carried today by the V32-bearing northern groups.
An Afar pastoralist woman. Archaeogenetic analysis of Middle Neolithic specimens excavated at the Skhirat site in the Maghreb has found that these pastoralists bore Levantine ancestry (cf. Simões (2023a); Simões (2023b)). Since both ancient and modern Afro-Asiatic speakers in the Horn vicinity and Nubians in the Nile Valley bear this Levantine ancestry, this suggests that pastoralism was first introduced to Africa during the Neolithic period, by herders who originated from the Middle East.
To confirm whether or not modern Cushitic, Ethiosemitic and North Omotic-speaking individuals of the Horn carry any Skhirat-derived Levantine ancestry, we performed a Vahaduo Single analysis on the official Global25 samples. It appears that most of these Afro-Asiatic speakers do bear this Middle Neolithic Levantine component, at frequencies ranging from a high of 35.2% in a Southern Somali individual to a low of 0% among a few of the Tigrayan samples. Perhaps unsurprisingly, the Middle Neolithic Skhirat component of the Levant, which again is linked to pastoralism, seems to be more salient among the contemporary pastoralist groups (especially Cushitic speakers), whereas the Epipaleolithic Natufian component of the Levant, which is linked to foraging and early kinds of agriculture, seems to be more pronounced among the modern agricultural groups (especially Ethiosemitic speakers). Overall, the Cushitic, Ethiosemitic and North Omotic-speaking individuals on average bear the same proportion of Eurasian ancestry as before (~70%), as do the Cushites of the Pastoral Neolithic and the medieval Kulubnarti Nubians (for the latter, see here and here). That includes European-related Steppe and East Asian components. The latter ancestral elements have not yet officially been identified in the archaeogenetic record; however, ancient haplogroup evidence supporting their existence has been detected (see the mitogenome charts above on the Meroitic period Kwieka and Christian era Ghazali samples).
Although one of the Skhirat samples was found to belong to the R1 paternal haplogroup, preliminary investigation has assigned this individual to the L2 subclade. This is a mainly Italian sublineage, distinct from both the Steppe-associated M269 subclade of R1b which has been detected among ancient Egyptian mummies (Gad et al. (2020a); Gad et al. (2020b); iGENEA; Yatsishina et al. (2021)) and the Anatolian Neolithic-associated V88 subclade of R1b which has been detected among Middle/Late Neolithic Sardinian specimens (Marcus et al. (2018), Supplementary Information). Thus, while the main Levantine ancestries of the Afro-Asiatic speakers and Nubians in the Horn and Nile Valley have now been detected in the paleogenetic record, the European Steppe and East Asian elements, which these individuals also seem to bear, have yet to be identified. Watch this space for an update when the original disseminators of these components have been found.
11 Saturday Mar 2023
Tags
ancient Azanians, artifact, Azania, Azanians, Caucasoid, Cushitic, maiden, Nikon, Periplus of the Erythraean Sea, Rhapta, Sarapion
The artifact below was excavated from the Merca area in coastal southern Somalia. It dates from the 6th century CE and shows a beautifully attired and bejeweled maiden carrying water:
A Cushitic Azanian maiden (6th century CE).
According to the geographer Ibn Said, during the 13th century, Merca was a stronghold of the “Hadiye” (cf. Lewis (1965); Reimer (1993)). The latter designation has usually been taken to refer to the Hawiye clan of the Somali, who still live in the area, rather than the Hadiyya group of the Sidamo, another Cushitic-speaking population today inhabiting southwestern Ethiopia (Reimer (1993)).
Given the Merca artifact’s pre-Islamic date and place of excavation, it therefore appears to depict an Azanian individual — the first visual representation of its kind to be discovered. Besides the textual evidence above, this is also attested by the female figure’s high stature, “Caucasoid” facial features, long soft-textured hair, almond-shaped eyes, agropastoral activity, and traditional attire. A physiognomy like that of the late Somali poet Hawa Jibril, shown below.
An Oromo woman of “pure” Cushitic type, showing little Sub-Saharan African influence. Note the long, flowing soft-textured hair, similar to that of the excavated “Azanian” female figure. Also notice the shared almond-shaped eyes, a common trait of the Cushitic peoples.
But just who were the Azanians? They were an early Cushitic people mentioned in the 1st century CE Periplus of the Erythraean Sea, a travelogue penned by a Greek merchant based in Alexandria, Ptolemaic Egypt. Traditional evidence on the Azanians describes them as “tall, bearded, long-haired, and ‘red’ in colour” (cf. Oliver (1988)). This is consistent with the Cushitic Somalis’ cosmogony (creation myth), which asserts that:
God created the white people and was quite pleased, then he created black people and was also quite pleased. Then God created Somalis and he laughed.
The traditional accounts on the Azanians also concur with testimony left by the explorer James Bruce, who visited Ethiopia between 1768 and 1773. Bruce indicates:
The Agow [Agaw]… are not negroes, but are of a coarse tawny red colour.[…] The Galla [Oromo]… speak a peculiar language, are very numerous, and are not negroes.[…] Native black Africans [are] called in Abyssinia Shankala[…] They are jet black, strong made, wooly-haired and thick-lipped. They are the aborigines of that part of Africa”.
According to the Periplus, the Azanians were “very great in stature” and inhabited the southern Somalia littoral and the territory just below that i.e., in ancient commercial ports like Nikon, Sarapion and Rhapta on the Azania coast.
Location of Azania according to the Periplus of the Erythraean Sea (c. 1st century CE). The Periplus indicates that the market-towns of Nikon, Sarapion and Rhapta along this southern coast were at the time inhabited by the ancient Azanians, Cushitic peoples who were “very great in stature” (Wickramasinghe (2018)).
Although the Periplus only alludes to the Azanians’ height, we can infer from it that they were not “Negroes” but instead a Cushitic people. Had the Azanians been Nilotes, the author of the Periplus surely would have also noted their pitch-black skin color since he does elsewhere remark on both the great stature and very dark complexion of the inhabitants of a certain part of India (viz. “Beyond the gulf of Baraca is that of Barygaza and the coast of the country of Ariaca, which is the beginning of the Kingdom of Nambanus and all of India[…] Very many cattle are pastured there, and the men are of great stature and black in color”). George Peter Murdock (1959) therefore concludes that: “it is significant that the [Periplus of the Erythraean Sea] author remarks only on [the Azanian’s] stature. He seems, therefore, to assume that they were Caucasians, for, to a Greek of his day, Negroes would have been strange beings whose characteristics would certainly have been noted.[…] The inhabitants can therefore have been no other than Megalithic Cushites who had descended the few miles from the Kenya highlands to the coast and there turned to maritime pursuits.”
In Somalia, the Cushitic Azanians would intermarry with early Arab and Persian merchants, thereby forming the modern Benadiri and Bravanese communities. The Periplus indicates:
Two days’ sail beyond, there lies the very last market-town of the continent of Azania, which is called Rhapta; which has its name from the sewed boats (rhapton ploiarion) already mentioned; in which there is ivory in great quantity, and tortoise-shell. Along this coast live men of piratical habits, very great in stature, and under separate chiefs for each place. The Mapharitic chief governs it under some ancient right that subjects it to the sovereignty of the state that is become first in Arabia. And the people of Muza now hold it under his authority, and send thither many large ships; using Arab captains and agents, who are familiar with the natives and intermarry with them, and who know the whole coast and understand the language.
The foregoing has been confirmed by haplogroup analysis, which has unveiled that most Benadiri individuals today carry the E1b1b and T paternal haplogroups like other ethnic Somalis (~52%; 35.48% E1b1b and 16.13% T), albeit with notable frequencies of the Arab-mediated J clade (~29%) and the Persian-affiliated R lineage (~6%) (cf. Immel and Kleiber (2009)). (*N.B. Among general Somalis, the J haplogroup has been observed at a low frequency of 3% (Sanchez et al. (2005)).) Genome analysis also shows a similar pattern:
— Genome analysis of a modern Benadiri Somali individual by the 23andme company. 23andme, AncestryDNA and other genetic testing labs indicate that Benadiris mostly bear Cushitic-related ancestry, averaging nearly 50%. The majority of this ancestry, approximately 40%, appears to be shared with Cushitic groups inhabiting Somalia (i.e., other ethnic Somalis); a minority of this ancestry, under 10%, seems to be more closely associated with Cushitic groups inhabiting Ethiopia & Eritrea (Oromos, Agaws, etc.). Additionally, Benadiris have some admixture derived from western Asia, averaging just under 25%. Most of this gene flow can be traced to contact with peninsular Arabs. Benadiri individuals, furthermore, bear under 15% Central/South Asian-related admixture, which likely dates to the Muslim period. Moreover, Benadiris have minor Bantu-associated admixture (~10%). This gene flow appears to be linked with Bantus from remote areas (e.g. Angola & Congo) rather than adjacent locales (Kenya); its presence is therefore primarily attributable to the recent Islamic slave trade rather than interaction with early Bantu settlers.
The Tanzanian scholars Felix Chami and Amandus Kwekason (2003) argue that the Azanians domiciled at Rhapta, the southernmost of the Periplus‘ market towns, were not Cushitic but instead early Bantu agriculturalists. They base this principally on Lionel Casson’s translation of the Periplus, which describes the Azanians as “tillers of the soil” rather than pastoralists. However, we can be reasonably certain that this identification is wrong and that the Azanians were indeed Cushitic peoples since:
A pillar tomb at Port Dunford, southern Somalia. The site marks the location of the old emporium of Nikon, as described in the Periplus of the Erythraean Sea. Such stone edifices were typical constructions of the ancient Azanians or “Megalithic Cushites.”
Descriptions of the early coastal people in classical documents had been considered to refer to Southern Cushites, with the origins of Tana Tradition lying in the Pastoral Neolithic wares of the Rift Valley of Kenya (Horton, 1996: 411). Support for this theory came from the similarity in shape and pattern in cooking vessels found at early coastal sites and those of Cushitic speaking pastoralists and hunter gatherers from the coastal hinterland (Kusimba, 1999: 108). The presence of Tana Tradition ceramics at virtually all sites during the 1st millennium AD on the coast suggests commonalities among the settlements, but there must also have been economic and social variation derived from the different access to imports as illustrated by excavations at Dar es Salaam and Mkiu (La Violette et al., 1999: 76) (Pollard (2008)).
TT pottery is our best marker for proto-Swahili society. From where does it derive? Typological study of forms and decorative schemes are currently being undertaken in a number of PhD theses at the University of Upsalla, but it is clear that we cannot link it with Early Iron pottery, known as the Mwitu tradition. Regional phases of these include Kwale Ware and Maore Ware. The Mwitu pottery is generally thought to be representative of the first Bantu speaking agriculturalists, who reached the coast by the 2nd century AD, with coastal hinterland sites known in particular from the Kwale, Pare and Usambara Hills. The recent redating of the Periplus to c. 40 AD) and the recognition that this described agriculturalists on the coast has raised the question of pre-Bantu agriculturalists in the late first millenium BC. The pottery of the pastoral neolithic (PN) provides a closer comparison to TT pottery. Finds from the Eastern Rift Valley, such as Maringishu and Narosura are relatively close in form and decoration. This material dates to the late 1st millenium BC or early centuries AD, and would be contemporary within the revised TT chronology. Placing TT pottery within a PN context has important implications. Reconstruction of language families and ceramics indicates that a Cushitic — probably Southern Cushitic — correlation for early TT pottery is likely. The implication is that proto-Swahili societies were Cushitic speaking and only adopted a Bantu language during the 1st millenium AD (Parkin (1994)).
(*N.B. The above is also supported by Prendergast et al. (2018), who analysed Iron Age remains from agricultural and pastoral sites in Kenya’s Rift Valley. In contrast to Wang et al. (2020)’s and Lipson et al. (2022)’s coastal samples from the same time period, these inland specimens had a preponderance of Bantu/Nilotic ancestry instead of forager ancestries, with ancillary West Eurasian admixture. Hence, the Cushites of the Pastoral Neolithic were primarily assimilated by Bantu/Nilotic peoples in the Rift Valley. Later Cushites (the Azanians), who dwelled along the Indian Ocean during the Iron Age, were absorbed by indigenous hunter-gatherers. Some of the Cushitic-admixed Bantus would subsequently leave the interior to settle along the southern Azanian coast, below the Horn. There, during the Middle Ages, these new Bantu arrivals would intermingle with Persians and other foreign Islamic settlers to form the Swahili culture.)
Admixture analysis of a Bajuni individual of Swahili type from Somalia. The person (labeled S25) was found to mainly harbor Bantu-related ancestry (light green component) with minor West Eurasian admixture, like other Swahili speakers from the Comoros and East Africa. However, unlike the Comorian Swahilis, the West Eurasian admixture borne by this Bajuni individual and the other East Africa Swahili samples is primarily associated with Cushitic-speaking populations of the Horn region (dark grey component, totaling under 20%) rather than Arabs (red component) or Iranians (light pink component) (Brucato et al. (2019)).
![Genetic modeling of medieval Swahili individuals. Most of the examined individuals derive about 60% of their ancestry from the earlier Bantu individuals from Makwasinyi, with the rest of their ancestry primarily derived from Persian sources and secondarily from Indian sources. Brielle et al. (2022) further note that "Makwasinyi associated [ancestry] itself is ~80% Bantu-related and ~20% pastoralist-related," where pastoralist includes ancestry related to the ancient Cushites of the Pastoral Neolithic. Ergo, before the admixture events between Persian settlers and Bantus that established the medieval Swahili culture, the Bantus would have moved from the continental interior, where they were originally based, to the seaboard. There, in their new place of settlement, the Bantus would have encountered and eventually absorbed the Cushitic Azanians, who, as we already know from archaeogenetic analysis of Iron Age skeletons, preceded them on the Swahili coast (Brielle et al. (2022)).](https://landofpunt.files.wordpress.com/2023/03/brielle2022-3.png?w=529)
Comparison of uniparental haplogroups (Y-DNA and mtDNA) carried by medieval Swahili individuals and modern Swahili individuals. Consistent with their mixed genome ancestry, the medieval Swahilis’ paternal clades are mostly associated with the Near East, whereas their maternal clades are mostly associated with African populations. Conversely, almost all of the modern Swahilis’ paternal and maternal clades are associated with African populations, in line with their own mostly Bantu-related genome ancestry (Brielle et al. (2022)).
Craniometric affinities of ancient and modern populations worldwide. Zanzibaris/Swahilis group with the Bantu and West African samples, despite having some Eurasian admixture. By contrast, the Cushitic-speaking Somalis cluster with the North African, ancient Egyptian, Bronze age Jericho and European samples. This is consistent with the ancient DNA analysis discussed above, which has found that modern Swahilis immediately descend from the medieval Bantus excavated at the inland site of Makwasinyi, Kenya (Brace (1993)).
Thus, on the coast immediately south of the Horn region, the Cushitic Azanians of the Iron Age would by the 10th century CE be completely absorbed or driven out by incoming populations; first by indigenous foragers and then by Bantu/Nilotic groups expanding from the hinterland:
1. “Rhapta is described as a place inhabited by big-bodied men. Many authorities remark that there is no specific mention of negroid people in the Periplus or of any distinction between the fair-skinned people of the Somali coast and the dark-skinned people south of the Juba. The implication is that Bantu-speaking people had not at this stage moved north of Rhapta. The phrase ‘big-bodied’ has also been taken to refer to Cushitic-speaking people (there are survivals of Cushitic languages in East Africa).”
Hilton, J. (1993). “Peoples of Azania“. Electronic Antiquity, 1(5)
2. “The Periplus records the names of ports, gives advice on how to comport oneself with local leaders, and describes the goods for trade. The Africans were tall and described as “red men” who fish, hunt, and herd cattle, sheep, and goats. These Azanians were most probably Cushitic speakers who had migrated into eastern Africa from Ethiopia, and were not the darker-complexioned inhabitants described by Muslim traders on the East African coast several centuries later. They had valuable items to trade — ivory, rhinoceros horn, tortoiseshell, spices, particularly cinnamon (Cinnamomum zeylanicum), the most profitable spice in the trade, and perfumes (frankincense, myrrh, and ambergris) — that were exchanged for iron, wheat, cloth, and porcelains.”
Collins, Robert O., and James McDonald Burns. A History of Sub-Saharan Africa. Cambridge: Cambridge UP, 2007.
3. “it is significant that the [Periplus of the Erythraean Sea] author remarks only on [the Azanian’s] stature. He seems, therefore, to assume that they were Caucasians, for, to a Greek of his day, Negroes would have been strange beings whose characteristics would certainly have been noted. In this assumption, of course, he was right, since, as noted above, the archeological evidence demonstrates indisputably the complete absence of Negroes in this part of Africa for centuries to come. The Cushitic peoples, however, are noted for their tall stature. The inhabitants can therefore have been no other than Megalithic Cushites who had descended the few miles from the Kenya highlands to the coast and there turned to maritime pursuits. This is attested by the numerous megalithic remains, including stone phalli, which still dot the Azanian coast.”
Murdock, George Peter. Africa: Its Peoples and Their Culture History. New York: McGraw-Hill Book, 1959.
In connection with the ancient DNA data and other evidence above, we may also note that historical texts link the Azanians with Cushitic peoples. The 9th century Chinese scholar Duan Chengshi mentions in his treatise Yu yang tsa tsu a territory in eastern Africa, which he refers to as Po-Pa-Li. The cultural and physical description he gives of this land’s inhabitants most closely matches Cushites. Duan Chengsi states that the denizens of Po-Pa-Li were pastoralists that raised cattle, whom they would bleed for sustenance. Traditionally, this was a common practice among Cushitic herders. The ancient Greek historian Agatharchides wrote that the Troglodytes (earlier Cushitic pastoralists) subsisted on a mixture of milk and blood, as they were disinclined to slay their cattle for meat (Hurskainen (1984)). Duang Chengsi further asserts, with regard to the inhabitants of Po-Pa-Li, that “their women are immaculately white, straight and tall.” This is a clear allusion to the cad (“olive-skinned”) phenotype that is found among the Cushitic peoples (see Ancient DNA from Ethiopia for details). Additionally, he notes that the territory was at the time “an independent country with an infantry of over 200 thousand men, strong enough to defy the powerful Tazi (the Arab Empire)” (cf. 袁武 (2006)). Ergo, the occupants of Po-Pa-Li were distinct from the Arabs. Duan Chengshi explicitly differentiates them from the ancient Persians, too, as he asserts that “when Po-saï (Persian) traders wish to enter this country, they form a caravan of several thousand men, and after having made (the natives) a present of strips of cloth, all of them both young and old draw blood by pricking themselves and take an oath, after which they trade their goods” (cf. Hasan (2017)).
Map of Trogloditica or the Horn of Africa, published in 1589 by the geographer Stefano Bonsignori. This document explicitly locates ancient Po-Pa-Li (Populi) in coastal southern Somalia, confirming that Po-Pa-Li was the Azania of yore and that its Cushitic inhabitants were the Azanians.
What’s more, the geographer Stefano Bonsignori published a map in 1589 of Trogloditica or the Horn of Africa (shown above), which locates Po-Pa-Li (Populi) in coastal southern Somalia. Coupled with Duan Chengshi’s testimony, we may conclude from this that the Po-Pa-Li of old was ancient Azania, and that, before the spread of Bantu/Nilotic peoples from the interior beginning in the 10th century CE (i.e. around 100 years after Duan Chengsi wrote his opus), the primary inhabitants of this territory would indeed have been none other than the Cushitic Azanians.
A Majerteen Darod Somali woman from Puntland. Genome analysis of modern Somali individuals born in the northeastern Puntland region of Somalia, who belong to the Darod clan (mostly the Majerteen subclan), indicates that they carry a predominant non-African ancestry. This Eurasian ancestry is on average comparable in frequency to that borne by North African individuals. Hence, the similarity in physiognomy which we have noted above between Benadiri/Bravanese Somalis and other ethnic Somalis from the northern clans is not coincidental. This likeness is instead grounded in the fact that both groups have elevated Eurasian ancestry, which was mostly inherited from their Cushitic ancestors. (For further details, see Which Afro-Asiatic-speaking population of the Horn region has the most non-African ancestry?.)
11 Saturday Mar 2023
It was mentioned in part 1 that Horn Africans with more angular features often get highest matches with European types within the Pictriev algorithm. Individuals with some neotenic or child-like characteristics like the woman below also sometimes get linked with folks with a similar Alpine phenotype (i.e. with a short, small nose & round baby-face, but still also Caucasoid in overall structure).
Sadia Shegow:
Highest matches with other non-European individuals with a Caucasoid phenotype within the algorithm system are more typical, though.
Ordinary Somalis:
And some prominent Somalis:
Mohamed Jawari:
Amina Mohamed:
Aar Maanta:
Abdihakim Abdullahi Haji Omar:
Khadra Ahmed:
Haji Mohamed Yasin Ismail:
Leila Abukar:
Mohamed Yusuf:
Asha Geele:
Abdulkadir Ali:
Mohamed Omar Arte:
Hussein Adan Igeh:
Asha Abdalla:
Asad Osman Abdullahi:
Hassan Mohamed Hussein:
As of January 2022, the Pictriev website appears to be down. Good news, though! A better alternative has since emerged: Betaface.
Overall, the Betaface algorithm works quite similarly to the Pictriev facial recognition system. One key difference between these services is that Betaface can compare the similitude of custom uploaded faces to those from its own internal database, as well as to celebrity profiles and images from the wider web. Betaface can also sort profiles according to race based on craniofacial attributes.
As with Pictriev, Afro-Asiatic-speaking individuals from Northeast Africa typically show highest similarity with persons sharing a “Caucasoid” craniofacial pattern. The race sorting for such individuals on Betaface’s detailed algorithm is usually as “white” (or sometimes “Middle Eastern”).
Below are examples of how Betaface works, using a prominent ethnic Somali individual.
Shadya Yasin:
(Betaface basic algorithm)
(Betaface detailed algorithm)
11 Saturday Mar 2023
Pictriev is a fun algorithm where a person can plug in any human face, including his/her own. It then matches that face on a percentage basis with the closest famous lookalikes of either gender within its system. The match algorithm is based on general facial feature and head shape recognition. However, some non-biological factors such as whether the individual is wearing glasses, has his/her head tilted or is facing a certain way, has stubble or a beard, is smiling or squinting, etc. can also affect the overall matches (mainly at the lower percentages).
For the average individual from the Horn region, the highest lookalike matches are other individuals with Caucasoid features. Since there are few other HOA folks in it at the moment, that’s the best it can do. For individuals with especially angular features, he/she may get highest matches with certain European types.
Here are some examples with ordinary Somalis:
And some prominent Somalis:
Haji Bashir Ismail:
Abdullahi Ahmed Irro:
Abdullahi Ahmed Addow:
Ismail Ali Ismail:
Abdulqadir Hashi:
Halima Ahmed:
Farah Ali Jama:
Barni Ahmed Qaasim (right) and her Somali friend:
Ahmed Hussen:
Ali Said Hassan:
Hamdi Bilan:
Abdullahi Yusuf Ahmed:
Mohamed Abdi Yussuf:
Fartun Abdulqadir:
Mohamed Abdullahi Mohamed:
MorphThing is another cool algorithm that allows you to morph between two to four faces, including fantastical creatures. I morphed several older and younger Somali male and female images and got the following composites:
Males:
Females:
Continued at Morphing & matching software pt. 2.
11 Saturday Mar 2023
Tags
Afro-Asiatic, Archaeology, excavations, Gol Waraabe, Isotopic, Land of Punt, Puntites, Pwene, Pwenet
The Mystery of the Land of Punt Unravelled is a fascinating book written by Ahmed Ibrahim Awale. He is a Somali scholar who led archaeological excavations in 2013 in Gol Waraabe, a site in Hargeisa valley in northwestern Somalia, where he and his team unearthed what appear to be the first actual artifacts belonging to the ancient Land of Punt. Based on these finds and their close similarity with antiquities from Egypt, Ibrahim hypothesizes that the ancient Egyptian culture (predynastic and dynastic alike) may have evolved from Puntite prototypes.
The first edition of the book seems to be out-of-stock in many online retailers. However, a second edition is now available for roughly the same price (a little over $10 USD). It contains more than 40 additional pages worth of new material.
The author, an environmentalist by training, begins by giving a brief but informative summary of the various scholarly theories on where the ancient Land of Punt was located. What’s great is that he also takes into account the recent isotopic work in Eritrea. Ibrahim then presents his hypothesis that the kingdom was situated in the Horn region, with its sociopolitical center in the Somali territories in particular. The rest of the book is devoted to proving that theory through both his own archaeological excavations and by pointing out similarities in culture, physiognomy, language, mythology and rock art.
Below are some of the excavated Puntite artifacts, which are currently housed at the Hargeisa national museum. Notice the commonalities between the Puntite statuette in the book’s Figure 6 and the ancient Egyptian Pharaoh Tutankhamun’s coffin in Figure 7. They include a similar headdress, oval face shape and narrow facial features, Puntite beard (Osiris beard), cobra and vulture headdress emblems, and crook and flail across the chest symbolism:
Another Puntite sculpture wearing the royal headdress with the cobra and vulture emblems, and possessing the standard oval face shape and narrow facial features; he also has the Puntite/Osiris beard:
These statues also resemble full-size shabti/shawabti, which are funerary figures that the ancient Egyptians and related early Afro-Asiatic-speaking groups in the Sudan area would often carve for the dead:
An Egyptian Faience Shabti for Ankh-Netcher-Hapy, Late Period, Dynasty XXVI (664-525 BCE).
Interestingly, among the excavated Puntite artifacts were several statuettes with elongated heads. This too is consistent with ancient Egyptian tradition, wherein the custom of artificial cranial deformation was practiced by some Pharaohs and royal officials:
Some of the excavated Puntite sculptures also bear similarities with ancient Egyptian religious figurines. For example, the Puntite artifact in Figure 11 and the ancient Egyptian bronze oil lamp in Figure 10, which both appear to have been made in honor of the crocodile deity Sobek:
The Puntite statuette in Figure 11 is actually the same one as on the cover, albeit shown in profile:
Furthermore, the lead archaeologist suggests that the Neolithic rock art at Laas Geel and elsewhere in the northwest and northeast Somali territories often depicts worshipers honoring the ancient Egyptian deity Hathor:
Note that the Puntite figures themselves are much bigger than this; the heads are actually larger than life-size. Besides these statuettes (which include several others alongside the ones above), the archaeologists also excavated some stone bowls, among other artifacts. This too is rather interesting since there happens to be an early Cushitic culture called the Savanna Pastoral Neolithic or Stone Bowl Culture, which is characterized by similar stone bowls.
It is interesting to note at this juncture that in 1899, during his final expedition at the old stone ruins in Zimbabwe (formerly Rhodesia) — a territory which Ibrahim (2013) suggests was a distant sourcing area for the Puntites, where they would often sojourn to obtain gold and some exotic trading products — the explorer Carl Peters announced that he had discovered a similar ancient Egyptian statue. This sculpture was inscribed with what appeared to be hieroglyphic symbols. Shortly afterwards, Peters submitted the object to the Egyptologist Flinders Petrie for inspection and appraisal. Petrie confirmed that “the figure is certainly genuinely ancient.” He noted further that on the statuette’s chest was an engraved cartouche of the Pharaoh Thutmoses III (Tahutmes III), the nephew of the Queen Hatshepsut; Petrie opined that the item therefore probably belonged to a courtier of this ruler (Keane (1901)). Peters’ find was met with ambivalence among the intelligentsia of his day. Although most pundits recognized the authenticity of the object itself, they questioned how to begin with such a statue had wound up at its remote place of excavation (cf. Keane (1901); The Independent (1901)).
On a related note, J. H. Patterson (1907) also reported the finding of comparable Egyptian-related antiquities in Mombasa, a coastal town in Kenya. Before the arrival of Bantus/Nilotes from the interior, this area was part of the realm of the Cushitic Azanians (see Who were the ancient Azanians?): “The town of Mombasa[…] is supposed to have been founded about A.D. 1000, but the discovery of ancient Egyptian idols, and of coins of the early Persian and Chinese dynasties, goes to show that it must at different ages have been settled by people of the very earliest civilisations.”
Overall, thanks to the newly excavated Puntite artifacts, the book succeeds in establishing the Horn region as the most likely location of the mysterious Land of Punt. It is easily among the most important works yet to be published on the ancient territory.
(For more details, refer to Could the ancient “Puntite” statues that were excavated in Somalia actually be forgeries?.)
In 2018, Robert Kluijver, curator of the Museum of Contemporary Ancient Arabia, announced the discovery of a second batch of ancient Puntite sculptures. This hoard was exhumed from private property in the vicinity of Berbera, situated on the coast of northwestern Somalia. The statues have been tentatively dated to 2000 years ago. This would make them contemporaneous with the old “Berber” port city of Malao, which the 1st century CE travelogue the Periplus of the Erythraean Sea indicates was located near present-day Berbera. See Punt: an ancient civilization rediscovered for details on this latest excavation.
Land of Punt 
