Genetic history of Egypt

This article's lead section may be too short to adequately summarize the key points. Please consider expanding the lead to provide an accessible overview of all important aspects of the article. (May 2023)

The genetic history of Egypt reflects its geographical location at the crossroads of several major biocultural areas:

Egyptologist Barry Kemp has noted that DNA studies can only provide firm conclusions about the population of ancient Egypt if the sample results are of a significant number of individuals and represent a broad geographical and chronological range.^[1] According to historian William Stiebling and archaeologist Susan N. Helft, conflicting DNA analysis on recent genetic samples such as the Amarna royal mummies has led to a lack of consensus on the genetic makeup of the ancient Egyptians and their geographic origins.^[2]

2012 study of Ramesses III

In 2012, two mummies of two 20th dynasty individuals, Ramesses III and "Unknown Man E" believed to be Ramesses III's son Pentawer, were analyzed by Albert Zink, Yehia Z Gad, and a team of researchers under Zahi Hawass. Genetic kinship analyses revealed identical haplotypes in both mummies; using the Whit Athey's haplogroup predictor, the Y chromosomal haplogroup E1b1a was predicted.^[3]

In another study by the same authors in 2020, which once again deals with the paternal lineage of Ramesses III and the "Unknown Man E" (possibly Pentawer), E1b1a is said to show its highest frequencies in modern West African populations (~80%) and Central Africa (~60%).^[4]

2017 DNA study of Mummies at Abusir el-Meleq

A study published in 2017 by Schuenemann et al. extracted DNA from 151 Egyptian mummies, whose remains were recovered from Abusir el-Meleq in Middle Egypt. The samples are from the time periods: Late

The absolute estimates of sub-Saharan African ancestry in these three ancient Egyptian individuals ranged from 6 to 15%, and the absolute estimates of sub-Saharan African ancestry in the 135 modern Egyptian samples ranged from 14 to 21%, which show an 8% increase in African component. The age of the ancient Egyptian samples suggests that this 8% increase in African component occurred predominantly within the last 2000 years.[5] The 135 modern Egyptian samples were: 100 from modern Egyptians taken from a study by Pagani et al., and 35 from el-Hayez Western Desert Oasis taken from a study by Kujanova et al.^[5] The 35 samples from el-Hayez Western Desert Oasis, whose population is described by the Kujanova et al. study as a mixed, relatively isolated, demographically small but autochthonous population, were already known from that study to have a relatively high sub-Saharan African component,^[6] which is more than 11% higher than the African component in the 100 modern Egyptian samples.^[7]

Verena Schuenemann and the authors of this study suggest a high level of genetic interaction with the Near East since ancient times, probably going back to Prehistoric Egypt although the oldest mummies at the site were from the New Kingdom: "Our data seem to indicate close admixture and affinity at a much earlier date, which is unsurprising given the long and complex connections between Egypt and the Middle East. These connections date back to Prehistory and occurred at a variety of scales, including overland and maritime commerce, diplomacy, immigration, invasion and deportation".^[5]

Responses to the 2017 DNA study

The 2017 study has generated academic responses from scholars from other related disciplines, remarking on the conclusions of the study from a multi-disciplinary approach.

In 2021, Gourdine et al disputed Scheunemann et al's claim, in an unpublished article, that the increase in the sub-Saharan component in the modern Egyptian samples resulted from the trans-Saharan slave trade. Instead they argued that the sub-Saharan "genetic affinities" may be attributed to "early settlers" and "the relevant sub-Saharan genetic markers" do not correspond with the geography of known trade routes".^[12]

In 2022, biological anthropologist S.O.Y. Keita argued that there were problems with the study's approaches and conclusions such as over-generalizations and a failure to consider alternative explanations. Particularly, he raised issues with the comparative samples from West Africa as a proxy group and generalisations about geographical Egypt and population origins from the sample results. He also drew attention to the fact that the authors draw inference on migrations in line with their Bayesian statistical approach rather than integrate other data into their explanations about the population history.^[13]

In 2022, archaeologist Danielle Candelora stated that there were several limitations with the 2017 Scheunemann et al. study such as “new (untested) sampling methods, small sample size and problematic comparative data”.^[14]

In 2023, Stiebling and Helft acknowledged that the 2017 study had performed the largest study on ancient Egyptians but noted that the findings still derived from a small sample of mummies from one site in Middle Egypt dating to the New Kingdom and later periods. They also stated that this study could not represent earlier populations or Egyptians from Upper Egypt who were geographically closer to Sub-Saharan populations.^[2]

In 2023, Christopher Ehret argued that the conclusions of the 2017 study were based on insufficiently small sample sizes, and that the authors had a biased interpretation of the genetic data.^[15] Ehret also criticised the Schuenemann article for asserting that there was “no sub-Saharan genetic component” in the Egyptian population and cited previous genetic analysis suggesting that the E-M35 paternal haplogroup originated in the Horn of Africa.^[16]

Later findings

A unpublished, follow-up study by Schuenemann & Urban et al. (2021) was carried out collecting samples from six excavation sites along the entire length of the Nile valley spanning 4000 years of Egyptian history. Samples from 17 mummies and 14 skeletal remains were collected, and high quality mitochondrial genomes were reconstructed from 10 individuals. According to the authors the analyzed mitochondrial genomes matched the results from the 2017 study at Abusir el-Meleq.^[17]

A study by Mussauer et al., presented at the 2023 ISBA10 conference (10th Meeting of the International Society for Biomolecular Archaeology), analysed the mtDNA of 25 Egyptian individuals dating from the Predynastic Period to the Coptic Period (c. 3500 BCE – 650 CE), from the archaeological sites of Asyut, Akhmim, Deir el-Bahari, Deir el-Medina, Thebes, the Valley of the Queens, and Gebelein. These samples displayed an mtDNA haplogroup diversity similar to the samples published by Schuenemann et al. 2017, providing "further evidence for shared maternal ancestries between western Eurasian or northern African populations and ancient Egyptians".^[18][5]

A 2020 study was conducted on ancient samples from Lebanon. Two individuals who lived in Lebanon around 500 BCE did not cluster with their contemporary Lebanese population. The study used the same Egyptian samples from the 2017 Schuenemann et al. study to further test these two individuals. One of these two individuals was a female who formed a clad with the three ancient Egyptian individuals from Schuenemann et al., implying that she shared all of her ancestry with them or a genetically equivalent population. The other one was a male who derived ~70% of his ancestry from a population related to the female and ~30% from a population related to ancient Levantines. Further testing suggests that the female was an Egyptian woman and the male was her son from a man who himself had both Egyptian and Lebanese ancestries.^[19]

2018 study of Nakht-Ankh and Khnum-Nakht

The tomb of two high-status Egyptians, Nakht-Ankh and Khnum-Nakht, was discovered by Egyptian workmen directed by Sir William

In 2018 the mummified head of Djehutynakht was analysed for mitochondrial DNA. Djehutynakht was the nomarch of the Hare nome in Upper Egypt during the 11th or 12th Dynasty in the early Middle Kingdom period, c. 2000 BC. Two laboratories independently analysed Djehutynakht's DNA and found that he belonged to the mtDNA haplogroup U5b2b5,^[23] described by the lead author Odile Loreille as "a European haplogroup".^[24] U5 is thought to have originated in Europe,^[25][26] and U5b2b5 has been found in ancient European samples dating from the Neolithic onwards.^[27][28][29] U5b2b5 has also been found in 10 samples from Christian Period Nubia,^[30] and a related European sequence (U5b2c1) has been observed in an ancient sample from Carthage (6th century BC).^[23][25] Among ancient Egyptian samples the Djehutynakht sequence resembles a U5a lineage from sample JK2903, a 2000-year-old skeleton from the Abusir el-Meleq site in Egypt.^[5] Haplogroup U5 is found in modern Egyptians,^[31][32] and is found in modern Egyptian Berbers from the Siwa Oasis in Egypt. A 2009 study by Coudray et al. recorded haplogroup U5 at 16.7% in the Siwa Oasis in Egypt, whereas haplogroup U6 is more common in other Berber populations to the west of Egypt.^[33]

2020 study on two Egyptian child mummies

A study on male child mummies from the Greco-Roman period originating in the Memphite or Luxor area, revealed that the mtDNA for one was T2c1a and the other HV. Identical or phylogenetically close derivatives of these lineages are present in both ancient and modern Egyptians, as well as among several present-day populations of the Near East and North Africa. The researchers noted that mtDNA alone is not enough to reach any precise conclusion about the origin of an individual, but the results are in accordance with an Egyptian origin. The ages of the two mummified corpses ranged from 11–15 years old, and 2–4 years old.^[34]

2020 study of Tutankhamun and other mummies of the 18th Dynasty

A 2020 study by Gad, Hawass, et al. analysed mitochondrial and Y-chromosomal haplogroups from

• Amenhotep III: YDNA R1b & mtDNA H2b.
• Tutankhamun: YDNA R1b & mtDNA K.
• Akhenaten: YDNA R1b & mtDNA K.
• Tiye: mtDNA K.
• Yuya: YDNA G2a & mtDNA K.
• Thuya: mtDNA K.

Both Y-DNA haplogroups R1b and G2a, as well as both mtDNA haplogroups H and K, are carried by modern Egyptians.^[4]

2020 study of Takabuti

In 2020 the mummy of Takabuti was tested for mitochondrial DNA. Takabuti was a noblewoman from Thebes in Upper Egypt who lived during the 25th Dynasty, c. 660 BC. Analysis of her DNA revealed that she belonged to the mtDNA haplogroup H4a1.^[37] The study states that “the H super-haplogroup is the most common mtDNA lineage in Europe and is found also in parts of present-day Africa and western Asia”.^[37] Haplogroup H is also carried by modern Egyptians, the subgroup H4 in particular, along with H2a1 and H13a1, account for 42% of H lineages in modern Egypt, with H4 accounting for 16.7% of H lineages in modern Egypt.^[38]

2020 study of mummies at the Kurchatov Institute

In 2020, three mummies, dating from the 1st millennium BCE, from the Pushkin Museum of Arts collection were tested at the Kurchatov Institute of Moscow for their mitochondrial and Y-chromosomal haplogroups. Two of the mummies were found to belong to the Y-chromosomal haplogroup R1b1a1b (R1b-M269),^[39] which originated either in Eastern Europe or in the Near East, and to the Y-chromosome haplogroup E1b1b1a1b2a4b5a,^[39] which originated in North Africa.^[40][5][41] They also belonged to mtDNA haplogroups L3h1 and N5, common in Africans and Middle Easterners, respectively. The third mummy was found to belong to mtDNA haplogroup N, which is widely distributed across Eurasia as well as eastern and northeastern Africa.^[39]

2022 comments on mummies of the 18th and 20th Dynasties

In a comment on Hawas et al. (2010^[42]& 2012^[3]), the anthropologist S.O.Y. Keita pointed out, based on inserting the data into the PopAffiliator online calculator, which only calculates affinity to East Asia, Eurasia, and sub-Saharan Africa, but not to North Africa or the Near East, for instance,^[43] that Ramesses III and the Amarna ancient royal family (including Tutankhamun) showed "an affinity with sub-Saharan Africans in one affinity analysis, which does not mean that they lacked other affiliations — an important point that typological thinking obscures. Also, different data and algorithms might give different results, which would illustrate the complexity of biological heritage and its interpretation."^[44]

Genetic studies on modern Egyptians

A 2004 mtDNA study of 58 upper Egyptian individuals included 34 individuals from

A 2005 genetic study found close affinities of eastern sub-Saharan populations with Egypt in the phylogenetic trees through analysis of the short DNA sequences. The authors suggested that the influential role of the Nile River served as a migratory route and an agent of genetic flow which contributed to present-day heterogeneity in Egypt.^[55]

Though there has been much debate of the origins of haplogroup M1 a 2007 study had concluded that M1 has West Asia origins not a Sub Saharan African origin, although the majority of the M1a lineages found outside and inside Africa had a more recent East African origin, as a result of "the first M1 backflow [from Asia] to Africa, dated around 30,000 [years ago]". The study states that "the most ancient dispersals of M1 occurred in northwestern Africa, reaching also the Iberian Peninsula, instead of Ethiopia", and states that the evidence points to either "that the Near East was the most probable origin of the primitive M1 dispersals, West into Africa and East to Central Asia ... [with] the Sinai Peninsula as the most probable gate of entrance of this backflow to Africa" or "that M1 is an autochthonous North African clad that had its earliest spread in northwestern areas marginally reaching the Near East and beyond".^[56]

However, other authors have proposed the view that the M haplogroup developed in Africa before the 'Out of Africa' event around 50,000 years ago, and dispersed from North Africa or East Africa 10,000 to 20,000 years ago. Quintana-Murci et al. stated in reference to the M haplogroup that "Its variation and geographical distribution suggest that Asian haplogroup M separated from eastern-African haplogroup M more than 50,000 years ago. Two other variants (489C and 10873C) also support a single origin of haplogroup M in Africa".^[57][58]

A 2003 Y-chromosome study was performed by Lucotte on modern Egyptians, with haplotypes V, XI, and IV being most common. Haplotype V is common among all

Babiker, H et al. (2011) examined the genotypes of 15 STRs for 498 individuals from 18 Sudanese populations and featured comparative genotype data with Egypt, Somalia and the Karamoja population from Uganda. Overall, the results showed that the genotypes of individuals from northern Sudan clustered with those of Egypt, the Somali population was found to be genetically distinct and individuals from southern Sudan clustered with those from the Karamoja population. The study determined that similarity of the Nubian and Egyptian populations suggested that migration, potentially bidirectional, occurred along the Nile river Valley, which is consistent with the historical evidence for long-term interactions between Egypt and Nubia.^[62]

An allele frequency comparative study led by the Egyptian Army Major General Doctor Tarek Taha conducted STR analysis in 2020 between the two main Egyptian ethnic groups, Muslims and Christians, each group represented by a sample of 100 unrelated healthy individuals, supported the conclusion that Egyptian Muslims and Egyptian Christians genetically originate from the same ancestors.^[63]

A Nov 2023 study by Hammarén et al isolated the non-african parts of the genomes of modern day northeast Africans found that Sudanese Copts and Egyptian muslims from Cairo bore most similarities to Levantines, unlike other populations in the region which had predominant genetic contributions from the Arabian peninsula rather than Levant for their Non-African genetic component. The study also found that Egyptian muslims and Sudanese Copts are genetically most similar to Middle Eastern groups rather than the other African populations, they estimated the admixture date for modern Egyptians with Eurasians to have occurred around the 14th century, however the authors noted that "most, if not all, of the populations in this study have or have had admixture with populations from the Middle East during the Arab expansion, and this newer admixture is obscuring older admixture patterns". The study overall points that the distribution of Eurasian ancestry in modern Eastern and Northeast Africa is the result of more recent migrations, many of which are recorded in historical texts rather than ancient one.^[64]

Y-DNA haplogroups

E1b1b and J, that are carried by both ancient and modern Egyptians.^[36][5][40] The subclade E-M78 of E1b1b is suggested to have originated in Northeast Africa in the area of Egypt and Libya, and is more predominant in Egypt.^[40] These two haplogroups and their various subclades in general are distributed in high frequencies in the Middle East and North Africa.^[65][66]

A study by Arredi et al., which analyzed 275 samples from five populations in Algeria, Tunisia, and Egypt, as well as published data from Moroccan populations, suggested that the North African pattern of Y-chromosomal variation, including in Egypt, is largely of Neolithic origin. The study analyzed North African populations, including North Egyptians and South Egyptians, as well as samples from southern Europe, the Middle East, and sub-Saharan Africa, and revealed the following conclusions about the male-lineage variation in North Africa: "The lineages that are most prevalent in North Africa are distinct from those in the regions to the immediate north and south: Europe and sub-Saharan Africa ... two haplogroups predominate within North Africa, together making up almost two-thirds of the male lineages: E3b2 and J* (42% and 20%, respectively). E3b2 is rare outside North Africa, and is otherwise known only from Mali, Niger, and Sudan to the immediate south, and the Near East and Southern Europe at very low frequencies. Haplogroup J reaches its highest frequencies in the Middle East".^[46]

A study by Lucotte using the

Sub-Saharan populations.^[60] Remarking on Lucotte's Y-chromosome study, which found that haplotypes V, XI, and IV are most common, Keita states that "a synthesis of evidence from archaeology, historical linguistics, texts, distribution of haplotypes outside Egypt, and some demographic considerations lends greater support to the establishment, before the Middle Kingdom, of the observed distributions of the most prevalent haplotypes V, XI, and IV. It is suggested that the pattern of diversity for these variants in the Egyptian Nile Valley was largely the product of population events that occurred in the late Pleistocene to mid-Holocene through the First Dynasty".^[59] Keita later states "Later, mid-Holocene climatic-driven migrations led to a major settlement of the valley in Upper Egypt and Nubia, but less so in Lower Egypt, by diverse Saharans with haplotypes IV, XI, and V. These people fused with the indigenous valley peoples, as did Near Easterners with VII and VIII, but perhaps also some V".^[67]

The major downstream mutations within the M35 subclade are M78 and M81. There are also other M35 lineages, e.g., M123. In Egypt, haplotypes VII and VIII are associated with the J haplogroup, which is predominant in the Near East.[59]

Population	Nb	A/B	E1b1a	E1b1b1 (M35)	E1b1b1a (M78)	E1b1b1b1 (M81)	E1b1b1b2 (M123,M34)	F	K	G	I	J1	J2	R1a	R1b	Other	Study
Egyptians	110	0	3.5%	0	36%	0	8.5%	0	0	7.5%	0	24.5%	2%	2.8%	8.4%	6.5%	Fadhloui-Zid et al. (2013)
Egyptians	370	1.35%	2.43%	3.24%	21.89%	11.89%	6.76%	1.08%	0.27%	5.68%	0.54%	20.81%	6.75%	2.16%	5.94%	9.21%	Bekada et al. (2013)[38]
Egyptians	147	2.7%	2.7%	0	18.4%	5.4%	0	0	8.2%	8.8%	0	19.7%	12.2%	3.4%	4.1%	2.1%	Luis et al. (2004)[36]
Egyptians from El-Hayez Oasis (Western Desert)	35	0	5.70%	5.7%	28.6%	28.6%	0	0	0	0	0	31.4%	0	0	0	0	Kujanová et al. (2009)[6]
Berbers from Siwa Oasis (Western Desert)	93	28.0%	6.5%	2.2%	6.5%	1%	0	0	0	3.2%	0	7.5%	6.5%	0	28.0%	8.3%	Dugoujon et al. (2009)[68]
Egyptians	87	1%	3%	10%	31%	0	2.5%	0	0	2%	0	20%	15%	5%	2%	8.5%	Pagani et al. (2015)
Northern Egyptians	44	2.3%	0	4.5%	27.3%	11.3%	0	6.8%	2.3%	0	0	9.1%	9.1%	2.3%	9.9%	6.8%	Arredi et al. (2004)
Southern Egyptians	29	0.0%	0	0	17.2%	6.8%	0	17.2%	10.3%	0	3.4%	20.7%	3.4%	0	13.8%	0	Arredi et al. (2004)

Distribution of E1b1b1a (E-M78) and its subclades

Population	N	E-M78	E-M78*	E-V12*	E-V13	E-V22	E-V32	E-V65	Study
Egyptians (sample includes people labeled as "berber" and people from the oases)	370	21.89%	0.81%	7.03%	0.81%	9.19%	1.62%	2.43%	Bekada et al. (2013)[38]
Southern Egyptians	79	50.6%		44.3%	1.3%	3.8%		1.3%	Cruciani et al. (2007)[40]
Egyptians from Bahari	41	41.4%		14.6%	2.4%	21.9%		2.4%	Cruciani et al. (2007)
Northern Egyptians (Delta)	72	23.6%		5.6%	1.4%	13.9%	2.8%		Cruciani et al. (2007)
Egyptians from Gurna Oasis	34	17.6%	5.9%	8.8%			2.9%		Cruciani et al. (2007)
Egyptian from Siwa Oasis	93	6.4%		2.1%				4.3%	Cruciani et al. (2007)

Mitochondrial DNA

In 2009 mitochondrial data was sequenced for 277 unrelated Egyptian individuals^[69] by Jessica L Saunier et al. in the journal Forensic Science International, as follows

• R0 and its subgroups (31.4%)
• L3 (12.3%); and Asian origin (n = 33)

including M (6.9%)

• T (9.4%)
• U (9.0%)
• J (7.6%)
• N (5.1%)
• K (4.7%)
• L2 (3.6%)
• L1 (2.5%)
• I (3.2%)
• W (0.7%)
• X (1.4%); African origin (n = 57) including L0 (2.2%)

Autosomal DNA

Mohamed, T et al. (2009) in their study of nomadic Bedouins featured a comparative study with a worldwide population database and a sample size of 153 Bedouin males. Their analysis discovered that both Muslim Egyptians and Coptic Christians showed a distinct North African cluster at 65%. This is their predominant ancestral component, and unique to the geographic region of Egypt.^[70]

In a 2019 study that analyzed the autosomal make-up of 21 modern North African genomes and other populations using Ancient DNA reference populations, this sample of Egyptian genomes were found to share more affinity with Middle Eastern populations compared to other North Africans. Egyptians carry more of the Caucasus hunter gatherer / Iran Neolithic component compared to other North Africans, more of the Natufian related component and less of the Iberomaurusian related component than other North Africans, and also less of the Steppe / European hunter gatherer component, consistent with Egypt's geographical proximity to southwest Asia.^[71]

Coptic Christians

A Y-DNA sample of Copts from Egypt was analyzed in Éric Crubézy et al. 2010. The Y-DNA profile was:

• 74% E1b1
• 7% G
• 3% T
• 1% J1
• 15% ambiguous.^[72]

Cruzby further added that “The haplotype E1b1 has a fairly wide distribution across Africa, but never had been described with such frequency in this area”. He also noted that Ethiopia was one of the areas in the world where the haplotype was most common. Cruzby suggested that this shared lineage could be attributed to the contacts between Coptic Christian communities in both Egypt and Ethiopia although the overlaps between the religious communities were rare.^[72]

In Sudan

According to Y-DNA analysis by Hassan et al. (2008), 45% of

Somalis.^[74] The next most common haplogroups borne by Copts are R1b (15%), most common in Europe, and the widespread African haplogroup B (15%). According to the study, the presence of haplogroup B may also be consistent with the historical record in which southern Egypt was colonized by Nilotic populations during the early state formation.^[73]

Maternally, Hassan (2009) found that the majority of Copts in Sudan (of a sample of 29) carried descendants of the

L1c.^[75]

A 2015 study by Dobon et al. identified an ancestral autosomal component of West Eurasian origin that is common to many modern Afroasiatic-speaking populations in Northeast Africa. Known as the Coptic component, it peaks among Egyptian Copts who settled in Sudan over the past two centuries, they also formed a separated group in PCA, a close outlier to other Egyptians, Afroasiatic-speaking Northeast Africans and Middle East populations. The Coptic component evolved out of a main North African and Middle Eastern ancestral component that is shared by other Egyptians and also found at high frequencies among other Afroasiatic-speaking populations in Northeast Africa (~70%), who carry a Nilo-Saharan element as well. The scientists suggest that this points to a common origin for the general population of Egypt.^[76] They also associate the Coptic component with Ancient Egyptian ancestry, without the later Arabic influence that is present among other Egyptians, especially people of the Sinai.^[76]

In another 2017 study that genotyped and analyzed the same populations including Sudanese Copts and Egyptians, The ADMIXTURE analyses and the PCA displayed the genetic affinity of the Copts to the Egyptian population. Assuming few clusters, the Copts appeared admixed between Near Eastern/European populations and northeastern Sudanese and look similar in their genetic profile to the Egyptians. Assuming greater number of clusters (K≥18), the Copts formed their own separate ancestry component that was shared with Egyptians but can also be found in Arab populations. This behavior in the admixture analyses is consistent with shared ancestry between Copts and Egyptians and/or additional genetic drift in the Copts. The Egyptians and Copts showed low levels of genetic differentiation (FST = 0.00236), lower levels of genetic diversity and greater levels of RoH compared to other northeast African groups, including Arab and Middle Eastern groups that share ancestry with the Copts and Egyptians. A formal test did not find significant admixture into the Egyptians from other tested groups (X), and the Copts and Egyptians displayed similar levels of European or Middle Eastern ancestry (Copts were estimated to be of 69.54% ± 2.57 European ancestry, and the Egyptians of 70.65% ± 2.47 European ancestry). Taken together, these results point to that the Copts and the Egyptians have a common history linked to smaller population sizes, and that Sudanese Copts have remained relatively isolated since their arrival to Sudan with only low levels of admixture with local northeastern Sudanese groups.^[77]

See also

Wikimedia Commons has media related to Ancient Egyptian DNA.

Wikimedia Commons has media related to Genetic studies on Egyptians.

• Ancient Egyptian race controversy
• Demographics of Egypt
• Genetic history of Africa
• Genetic history of North Africa
• Genetic history of the Middle East
• Genetic studies on Arabs
• Population history of Egypt

References

1. ISBN 9781134563883. Archived
   from the original on 14 April 2023. Retrieved 14 April 2023.
2. ^
   ISBN 978-1-000-88066-3
   .
3. ^
   S2CID 206896841
   .
4. ^
   ISBN 978-80-7308-979-5
   .
5. ^
   PMID 28556824
   .
6. ^
   PMID 19425100
   .
7. PMID 26027499
   .
8. PMID 28556824
   .
9. PMID 28556824
   .
10. PMID 28556824
    .
11. PMID 28556824
    .
12. ISBN 978-0-521-84067-5
    .
13. ISBN 978-0367434632.{{cite book}}: CS1 maint: location missing publisher (link
    )
14. ISBN 9780367434632.{{cite book}}: CS1 maint: location missing publisher (link
    )
15. ISBN 978-0-691-24409-9. Archived
    from the original on 22 March 2023. Retrieved 20 March 2023.
16. ISBN 978-0-691-24410-5
    .
17. ^ "Human mitochondrial haplogroups and ancient DNA preservation across Egyptian history (Urban et al. 2021)" (PDF). ISBA9, 9th International Symposium on Biomolecular Archaeology, p.126. 2021. Archived (PDF) from the original on 16 October 2021. Retrieved 14 January 2023. In a previous study, we assessed the genetic history of a single site: Abusir el-Meleq from 1388 BCE to 426 CE. We now focus on widening the geographic scope to give a general overview of the population genetic background, focusing on mitochondrial haplogroups present among the whole Egyptian Nile River Valley. We collected 81 tooth, hair, bone, and soft tissue samples from 14 mummies and 17 skeletal remains. The samples span approximately 4000 years of Egyptian history and originate from six different excavation sites covering the whole length of the Egyptian Nile River Valley. NGS 127 based ancient DNA 8 were applied to reconstruct 18 high-quality mitochondrial genomes from 10 different individuals. The determined mitochondrial haplogroups match the results from our Abusir el-Meleq study.
18. ^ "Genetic study of ancient Egyptian human remains dating from the Predynastic Period to the early Islamic Period (ca. 4000 cal. BCE - 800 cal. CE) (Mussauer et al.)". ISBA10: 10th Meeting of the International Society for Biomolecular Archaeology. 2023. p. Abstract Book, p.222.
19. PMID 32470374
    .
20. doi:10.1016/j.jasrep.2017.12.025. Archived
    from the original on 17 January 2022. Retrieved 12 November 2021.
21. ^ "The kinship of two 12th Dynasty mummies revealed by ancient DNA sequencing" (PDF).
22. ^ "FamilyTreeDNA - Notable Connections - Nakht-Ankh". FamilyTreeDNA Discover (Beta). Retrieved 15 September 2023.
23. ^
    PMID 29494531
    .
24. ^ "Tomb secrets: The FBI cracks the DNA code on an ancient Egyptian mummy". Independent. 21 June 2018.
25. ^
    PMID 27224451
    .
26. PMID 11032788
    .
27. PMID 20178764
    .
28. PMID 30872528
    .
29. PMID 32313080
    .
30. PMID 34907168
    .
31. PMID 23910099
    .
32. PMID 19414160
    .
33. S2CID 21826485
    .
34. PMID 31945091
    .
35. PMID 33059357. Archived
    from the original on 2 May 2021. Retrieved 19 December 2022.
36. ^
    PMID 14973781
    .
37. ^
    PMID 33046824
    .
38. ^
    PMID 23431392
    .
39. ^
    S2CID 232315321
    . Haplogroup R1b1a1b isolated in the studied samples is also uncharacteristic of the modern Egyptian population; its frequency in the population is less than 1%, while it is found in approximately half of the male population of Western Europe.
40. ^
    PMID 17351267
    .
41. PMID 34907168
    .
42. PMID 20159872. Archived
    from the original on 14 April 2020. Retrieved 21 March 2023.
43. S2CID 11740334
    .
44. ^ "Analysis of the short tandem repeat (STR) data published on Ramesses III and the Amarna ancient royal family (including Tutankhamun) showed a majority to have an affinity with “sub-Saharan” Africans in one affinity analysis, which does not mean that they lacked other affiliations—an important point that typological thinking obscures".Keita, S. O. Y. (September 2022). "Ideas about "Race" in Nile Valley Histories: A Consideration of "Racial" Paradigms in Recent Presentations on Nile Valley Africa, from "Black Pharaohs" to Mummy Genomest". Journal of Ancient Egyptian Interconnections. Retrieved 11 May 2023.
45. ^
    S2CID 26741827
    .
46. ^
    PMID 15202071
    .
47. PMID 10090902
    .
48. ISBN 978-1-902937-20-5
    .
49. ^
    PMID 17351267
    .
50. ISBN 978-0-691-02905-4
    .
51. ^
    S2CID 44901197
    .
52. Cavalli-Sforza, L.L., P. Menozzi, and A. Piazza (1994). The History and Geography of Human Genes Archived 4 April 2023 at the Wayback Machine
    . Princeton:Princeton University Press. p. 174. ISBN 0-691-08750-4
53. ISBN 978-0-691-08750-4. Archived
    from the original on 4 April 2023. Retrieved 19 March 2023.
54. PMID 15457403
    .
55. S2CID 36880409
    .
56. PMID 17620140
    .
57. PMID 12536373
    .
58. S2CID 2000627. Archived
    from the original on 19 April 2022. Retrieved 19 April 2022.
59. ^
    S2CID 33076762
    .
60. ^
    PMID 12687584
    .
61. ISBN 978-90-272-8985-8.{{cite book}}: CS1 maint: multiple names: authors list (link
    )
62. S2CID 7390979
    .
63. S2CID 219586129. Archived
    from the original on 3 June 2021. Retrieved 14 April 2023.
64. PMID 37939042
    .
65. ^ "Phylogeographic Analysis of Haplogroup E3b (E-M215) Y-Chromosomes Reveals Multiple Migratory Events Within and Out Of Africa" (PDF). 26 June 2008. Archived from the original (PDF) on 26 June 2008. Retrieved 13 April 2023.
66. S2CID 20279122. Archived
    (PDF) from the original on 24 September 2016. Retrieved 24 September 2016.
67. S2CID 33076762. Archived
    from the original on 11 June 2022. Retrieved 11 June 2022.
68. ^ Dugoujon J.M., Coudray C., Torroni A., Cruciani F., Scozzari F., Moral P., Louali N., Kossmann M. The Berber and the Berbers: Genetic and linguistic diversities
69. PMID 19414160
    .
70. PMID 19639002
    .
71. S2CID 204972040
    .
72. ^
    S2CID 248278173
    .
73. ^
    PMID 18618658
    .
74. PMID 26108492
    .
75. ^ Yousif, Hisham; Eltayeb, Muntaser (July 2009). Genetic Patterns of Y-chromosome and Mitochondrial DNA Variation, with Implications to the Peopling of the Sudan (Thesis). Archived from the original on 12 November 2021. Retrieved 12 November 2021.
76. ^
    PMID 26017457
    .
77. PMID 28837655
    .