Comprehensive view of the population history of Arabia as inferred by mtDNA variation

American Journal of Physical Anthropology

Volumn 159, Issue 4, 2016, Pages 607-616

  Černý, Viktor ^a   Čížková, Martina ^b   Poloni, Estella S ^c   Al Meeri, Ali ^d   Mulligan, Connie J ^e  

^a INSTITUTE OF MATHEMATICS   (Czech Republic)

^b CHARLES UNIVERSITY   (Czech Republic)

^c UNIVERSITY OF GENEVA   (Switzerland)

^d SANA A UNIVERSITY   (Yemen)

^e University of Florida ^*  (United States)

Author keywords

Arabian Peninsula; migrations; mtDNA variation

Indexed keywords

MITOCHONDRIAL DNA;

AFRICA; ARTICLE; ASIA; BALKAN PENINSULA; CONTROLLED STUDY; DNA SEQUENCE; DNA STRUCTURE; GENETIC DISTANCE; GENETIC VARIABILITY; GEOGRAPHIC DISTRIBUTION; HAPLOTYPE; HUMAN; HYPERVARIABLE SEGMENT 1; INDIA; MIGRATION; POPULATION GENETICS; POPULATION GROWTH; YEMEN; ETHNOLOGY; GENETIC VARIATION; GENETICS; HISTORY; MIDDLE EAST; PHYSICAL ANTHROPOLOGY;

AFRICA; ANTHROPOLOGY, PHYSICAL; ARABIA; DNA, MITOCHONDRIAL; GENETIC VARIATION; GENETICS, POPULATION; HISTORY, 21ST CENTURY; HISTORY, ANCIENT; HUMANS; INDIA; MIDDLE EAST;

EID: 84960315667     PISSN: 00029483     EISSN: 10968644     Source Type: Journal    
DOI: 10.1002/ajpa.22920     Document Type: Article

References (86)

1. Abu-Amero KK, Gonzalez AM, Larruga JM, Bosley TM, Cabrera VM., 2007. Eurasian and African mitochondrial DNA influences in the Saudi Arabian population. BMC Evol Biol 7: 32.
2. Abu-Amero KK, Larruga JM, Cabrera VM, Gonzalez AM., 2008. Mitochondrial DNA structure in the Arabian Peninsula. BMC Evol Biol 8: 45.
3. Al-Abri AR, Podgorná E, Rose JI, Pereira L, Mulligan CJ, Silva NM, Bayoumi R, Soares P, Černý V., 2012. Pleistocene-Holocene boundary in Southern Arabia from the perspective of human mtDNA variation. Am J Phys Anthropol 149: 291-298.
4. Al-Meeri A, Non AL, Lajoie TW, Mulligan CJ., 2011. Effect of different sampling strategies for a single geographic region in Yemen on standard genetic analyses of mitochondrial DNA sequence data. Mitochondrial DNA 22: 66-70.
5. Alshamali F, Pereira L, Budowle B, Poloni ES, Currat M., 2009. Local population structure in Arabian Peninsula revealed by Y-STR diversity. Hum Hered 68: 45-54.
6. Andrews RM, Kubacka I, Chinnery PF, Lightowlers RN, Turnbull DM, Howell N., 1999. Reanalysis and revision of the Cambridge reference sequence for human mitochondrial DNA. Nat Genet 23: 147.
7. Armitage SJ, Jasim SA, Marks AE, Parker AG, Usik VI, Uerpmann HP., 2011. The southern route "out of Africa": evidence for an early expansion of modern humans into Arabia. Science 331: 453-456.
8. Atkinson QD, Gray RD, Drummond AJ., 2008. mtDNA variation predicts population size in humans and reveals a major Southern Asian chapter in human prehistory. Mol Biol Evol 25: 468-474.
9. Badro DA, Douaihy B, Haber M, Youhanna SC, Salloum A, Ghassibe-Sabbagh M, Johnsrud B, Khazen G, Matisoo-Smith E, Soria-Hernanz DF, Wells RS, Tyler-Smith Ch, Platt DE, Zalloua PA, The Genographic Consortium. 2013. Y-chromosome and mtDNA genetics reveal significant contrasts in affinities of modern Middle Eastern populations with European and African populations. PLoS One 8: e54616.
10. Bendall KE, Sykes BC., 1995. Length heteroplasmy in the first hypervariable segment of the human mtDNA control region. Am J Hum Genet 57: 248-256.
11. Boivin N, Blench R, and, Fuller DQ., 2009. Archaeological, linguistic and historical sources on ancient seafaring: a multidisciplinary approach to the study of early maritime contact and exchange in the arabian peninsula. In:, Petraglia MD, and, Rose JI, editors. The evolution of human populations in Arabia Paleoenvironments, prehistory and genetics. New York: Springer. p 251-278.
12. Breton J-F., 1999. Arabia Felix from the time of the Queen of Sheba: eighth century B.C. to first century A.D. Notre Dame, IN: University of Notre Dame Press.
13. Brunner U., 2008. Les pièges de chasse antiques au Yémen. Arab Hum 15: 29-34.
14. Cabrera VM, Abu-Amero KK, Larruga JM, González AM., 2009. The Arabian Peninsula: gate for human migrations out of Africa or Cul-de-Sac? A mitochondrial DNA phylogeographic perspective. In:, Petraglia MD, Rose JI, editors. The evolution of human populations in Arabia Paleoenvironments, prehistory and genetics. New York: Springer. p 79-87.
15. Cadenas AM, Zhivotovsky LA, Cavalli-Sforza LL, Underhill PA, Herrera RJ., 2008. Y-chromosome diversity characterizes the Gulf of Oman. Eur J Hum Genet 16: 374-386.
16. Chaubey G, Metspalu M, Choi Y, Magi R, Romero IG, Soares P, van Oven M, Behar DM, Rootsi S, Hudjashov G, Mallick CB, Karmin M, Nelis M, Parik J, Reddy AG, Metspalu E, van Driem G, Xue Y, Tyler-Smith C, Thangaraj K, Singh L, Remm M, Richards MB, Lahr MM, Kayser M, Villems R, Kivisild T,. 2011. Population genetic structure in Indian Austroasiatic speakers: the role of landscape barriers and sex-specific admixture. Mol Biol Evol 28: 1013-1024.
17. Chiaroni J, King RJ, Myres NM, Henn BM, Ducourneau A, Mitchell MJ, Boetsch G, Sheikha I, Lin AA, Nik-Ahd M, Ahmad J, Lattanzi F, Herrera RJ, Ibrahim ME, Brody A, Semino O, Kivisild T, Underhill PA,. 2010. The emergence of Y-chromosome haplogroup J1e among Arabic-speaking populations. Eur J Hum Genet 18: 348-353.
18. Crassard R, Hilbert YH., 2013. A Nubian complex site from central Arabia: implications for Levallois taxonomy and human dispersals during the upper Pleistocene. PLoS One 8: e69221.
19. Crassard R, Petraglia MD, Parker AG, Parton A, Roberts RG, Jacobs Z, Alsharekh A, Al-Omari A, Breeze P, Drake NA, Groucutt HS, Jennings R, Régagnon E, Shipton C,. 2013. Beyond the Levant: first evidence of a pre-pottery Neolithic incursion into the Nefud Desert, Saudi Arabia. PLoS One 8: e68061.
20. Cremaschi M, Zerboni A, Charpentier V, Crassard R, Isola I, Regattieri E, Zanchetta G., 2015. Early-Middle Holocene environmental changes and pre-Neolithic human occupations as recorded in the cavities of Jebel Qara (Dhofar, southern Sultanate of Oman). Quat Int 382: 264-276.
21. Černý V, Mulligan CJ, Fernandes V, Silva NM, Alshamali F, Non A, Harich N, Cherni L, El Gaaied AB, Al-Meeri A, Pereira L,. 2011. Internal diversification of mitochondrial haplogroup R0a reveals post-last glacial maximum demographic expansions in South arabia. Mol Biol Evol 28: 71-78.
22. Černý V, Mulligan CJ, Rídl J, Žaloudková M, Edens CM, Hájek M, Pereira L., 2008. Regional differences in the distribution of the sub-Saharan, West Eurasian, and South Asian mtDNA lineages in Yemen. Am J Phys Anthropol 136: 128-137.
23. Černý V, Pereira L, Kujanová M, Vašíková A, Hájek M, Morris M, Mulligan CJ., 2009. Out of Arabia-the settlement of island Soqotra as revealed by mitochondrial and Y chromosome genetic diversity. Am J Phys Anthropol 138: 439-447.
24. de Maigret A., 2003. Arabia Felix. An exploration of the Archaeological History of Yemen. London: Stacey International.
25. Delagnes A, Tribolo C, Bertran P, Brenet M, Crassard R, Jaubert J, Khalidi L, Mercier N, Nomade S, Peigné S, Sitzia L, Tournepiche J-F, Al-Halibi M, Al-Mosabi A, Macchiarelli R,. 2012. Inland human settlement in southern Arabia 55,000 years ago. New evidence from the Wadi Surdud Middle Paleolithic site complex, western Yemen. J Hum Evol 63: 452-474.
26. Drechsler H., 2007. The Neolithic dispersal into Arabia. Proc Semin Arab Stud 37: 93-109.
27. Dupanloup I, Schneider S, Excoffier L., 2002. A simulated annealing approach to define the genetic structure of populations. Mol Ecol 11: 2571-2581.
28. Eaaswarkhanth M, Haque I, Ravesh Z, Romero IG, Meganathan PR, Dubey B, Khan FA, Chaubey G, Kivisild T, Tyler-Smith C, Singh L, Thangaraj K,. 2010. Traces of sub-Saharan and Middle Eastern lineages in Indian Muslim populations. Eur J Hum Genet 18: 354-363.
29. Excoffier L, Laval G, Schneider S., 2005. Arlequin ver. 3.0: An integrated software package for population genetics data analysis. Evol Bioinformatics Online 1: 47-50.
30. Excoffier L, Smouse PE, Quattro JM., 1992. Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics 131: 479-491.
31. Fernandes V, Alshamali F, Alves M, Costa MD, Pereira JB, Silva NM, Cherni L, Harich N, Černý V, Soares P, Richards MB, Pereira L,. 2012. The Arabian cradle: mitochondrial relicts of the first steps along the southern route out of Africa. Am J Hum Genet 90: 347-355.
32. Fernandes V, Triska P, Pereira JB, Alshamali F, Rito T, Machado A, Fajkošová Z, Cavadas B, Černý V, Soares P, Richards MB, Pereira L,. 2015. Genetic stratigraphy of key demographic events in Arabia. PLoS One 10: e0118625.
33. Forster P, Torroni A, Renfrew C, Rohl A., 2001. Phylogenetic star contraction applied to Asian and Papuan mtDNA evolution. Mol Biol Evol 18: 1864-1881.
34. Freitag U, Clarence-Smith W., 1997. Hadhrami traders, scholars and statesmen in the Indian Ocean, 1750s to 1960s. Leiden: Brill.
35. Fu YX., 1997. Statistical tests of neutrality of mutations against population growth, hitchhiking and background selection. Genetics 147: 915-925.
36. Gajda I., 2009. Le royaume de Himyar à l'époque monothéiste: L'histoire de l'Arabie du sud ancienne de la fin du IVe siècle. Paris: Mémoires de l'Académie des Inscriptions et Belles-Lettres, tome 40.
37. Galego LM, Jones ER, Eriksson A, Siska V, Arthur KW, Arthur JW, Curtis MC, Stock JT, Coltorti M, Pieruccini P, Stretton S, Brock F, Higham T, Park Y, Hofreiter M, Bradley DG, Bhak J, Pinhasi R, Manica A,. 2015. Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent. Science 350: 820-822.
38. Gonder MK, Mortensen HM, Reed FA, de Sousa A, Tishkoff SA., 2007. Whole-mtDNA genome sequence analysis of ancient African lineages. Mol Biol Evol 24: 757-768.
39. Groucutt HS, Petraglia MD., 2012. The prehistory of the Arabian peninsula: deserts, dispersals, and demography. Evol Anthropol 21: 113-125.
40. Groucutt HS, Petraglia MD, Bailey G, Scerri EM, Parton A, Clark-Balzan L, Jennings RP, Lewis L, Blinkhorn J, Drake NA, Breezem PS, Inglis RH, Devès MH, Meredith-Williams M, Boivin N, Thomas MG, Scally A,. 2015a. Rethinking the dispersal of Homo sapiens out of Africa. Evol Anthropol 24: 149-164.
41. Groucutt HS, Scerri EM, Lewis L, Clark-Balzan L, Blinkhorn J, Jennings RP, Parton A, Petraglia MD., 2015b. Stone tool assemblages and models for the dispersal of Homo sapiens out of Africa. Quat Int 382: 8-30.
42. Henn BM, Botigue LR, Gravel S, Wang W, Brisbin A, Byrnes JK, Fadhlaoui-Zid K, Zalloua PA, Moreno-Estrada A, Bertranpetit J, Bustamante CD, Comas D,. 2012. Genomic ancestry of North Africans supports back-to-Africa migrations. PLoS Genet 8: e1002397.
43. Hilbert Y, Parton A, Morley MW, Linnenlucke LP, Jacobs Z, Clark-Balzan L, Roberts RG, Galletti CS, Schwenninger JL, Rose J., 2015. Terminal Pleistocene and Early Holocene archaeology and stratigraphy of the southern Nejd, Oman. Quat Int 382: 250-263.
44. Hodgson JA, Mulligan CJ, Al-Meeri A, Raaum RL., 2014. Early back-to-Africa migration into the Horn of Africa. PLoS Genet 10: e1004393.
45. Karmin M, Saag L, Vicente M, Wilson Sayres MA, Jarve M, Talas UG, Rootsi S, Ilumae AM, Magi R, Mitt M, Pagani L, Puurand T, Faltyskova Z, Clemente F, Cardona A, Metspalu E, Sahakyan H, Yunusbayev B, Hudjashov G, DeGiorgio M, Loogväli EL, Eichstaedt C, Eelmets M, Chaubey G, Tambets K, Litvinov S, Mormina M, Xue Y, Ayub Q, Zoraqi G, Korneliussen TS, Akhatova F, Lachance J, Tishkoff S, Momynaliev K, Ricaut FX, Kusuma P, Razafindrazaka H, Pierron D, Cox MP, Sultana GN, Willerslev R, Muller C, Westaway M, Lambert D, Skaro V, Kovačevic L, Turdikulova S, Dalimova D, Khusainova R, Trofimova N, Akhmetova V, Khidiyatova I, Lichman DV, Isakova J, Pocheshkhova E, Sabitov Z, Barashkov NA, Nymadawa P, Mihailov E, Seng JW, Evseeva I, Migliano AB, Abdullah S, Andriadze G, Primorac D, Atramentova L, Utevska O, Yepiskoposyan L, Marjanovic D, Kushniarevich A, Behar DM, Gilissen C, Vissers L, Veltman JA, Balanovska E, Derenko M, Malyarchuk B, Metspalu A, Fedorova S, Eriksson A, Manica A, Mendez FL, Karafet TM, Veeramah KR, Bradman N, Hammer MF, Osipova LP, Balanovsky O, Khusnutdinova EK, Johnsen K, Remm M, Thomas MG, Tyler-Smith C, Underhill PA, Willerslev E, Nielsen R, Metspalu M, Villems R, Kivisild T,. 2015. A recent bottleneck of Y chromosome diversity coincides with a global change in culture. Genome Res 25: 459-466.
46. Kennedy H., 2008. The great Arab conquests: how the spread of Islam changed the world we live in. Boston: Da Capo Press.
47. Kitchen A, Ehret C, Assefa S, Mulligan CJ., 2009. Bayesian phylogenetic analysis of Semitic languages identifies an Early Bronze Age origin of Semitic in the Near East. Proc Biol Sci 276: 2703-2710.
48. Kivisild T, Reidla M, Metspalu E, Rosa A, Brehm A, Pennarun E, Parik J, Geberhiwot T, Usanga E, Villems R., 2004. Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears. Am J Hum Genet 75: 752-770.
49. Kivisild T, Shen P, Wall DP, Do B, Sung R, Davis K, Passarino G, Underhill PA, Scharfe C, Torroni A, Scozzari R, Modiano D, Coppa A, de Knijff P, Feldman M, Cavalli-Sforza LL, Oefner PJ,. 2006. The role of selection in the evolution of human mitochondrial genomes. Genetics 172: 373-387.
50. Kloss-Brandstatter A, Pacher D, Schonherr S, Weissensteiner H, Binna R, Specht G, Kronenberg F., 2011. HaploGrep: a fast and reliable algorithm for automatic classification of mitochondrial DNA haplogroups. Hum Mutat 32: 25-32.
51. Librado P, Rozas J., 2009. DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics 25: 1451-1452.
52. Macaulay V, Hill C, Achilli A, Rengo C, Clarke D, Meehan W, Blackburn J, Semino O, Scozzari R, Cruciani F, Taha A, Shaari NK, Raja JM, Ismail P, Zainuddin Z, Goodwin W, Bulbeck D, Bandelt HJ, Oppenheimer S, Torroni A, Richards M,. 2005. Single, rapid coastal settlement of Asia revealed by analysis of complete mitochondrial genomes. Science 308: 1034-1036.
53. Manger L., 2010. The Hadrami Diaspora: community-building on the Indian Ocean Rim. New York: Berghahn Books.
54. Mantel N., 1967. The detection of disease clustering and a generalized regression approach. Cancer Res 27: 209-220.
55. McCorriston J, Harrower M, Martin L, Oches E., 2012. Cattle cults of the Arabian Neolithic and early territorial societies. Am Anthropol 114: 45-63.
56. McDougall I, Brown FH, Fleagle JG., 2008. Sapropels and the age of hominins Omo I and II, Kibish, Ethiopia. J Hum Evol 55: 409-420.
57. Mellars P., 2006. Going east: new genetic and archaeological perspectives on the modern human colonization of Eurasia. Science 313: 796-800.
58. Mellars P, Gori KC, Carr M, Soares PA, Richards MB., 2013. Genetic and archaeological perspectives on the initial modern human colonization of southern Asia. Proc Natl Acad Sci USA 110: 10699-10704.
59. Metspalu M, Kivisild T, Metspalu E, Parik J, Hudjashov G, Kaldma K, Serk P, Karmin M, Behar DM, Gilbert MT, Endicott P, Mastana S, Papiha SS, Skorecki K, Torroni A, Villems R,. 2004. Most of the extant mtDNA boundaries in south and southwest Asia were likely shaped during the initial settlement of Eurasia by anatomically modern humans. BMC Genet 5: 26.
60. Musilová E, Fernandes V, Silva NM, Soares P, Alshamali F, Harich N, Cherni L, Gaaied AB, Al-Meeri A, Pereira L, Černý V., 2011. Population history of the Red Sea-genetic exchanges between the Arabian Peninsula and East Africa signaled in the mitochondrial DNA HV1 haplogroup. Am J Phys Anthropol 145: 592-598.
61. Oota H, Settheetham-Ishida W, Tiwawech D, Ishida T, Stoneking M., 2001. Human mtDNA and Y-chromosome variation is correlated with matrilocal versus patrilocal residence. Nat Genet 29: 20-21.
62. Pagani L, Kivisild T, Tarekegn A, Ekong R, Plaster C, Gallego Romero I, Ayub Q, Mehdi SQ, Thomas MG, Luiselli D, Bekele E, Bradman N, Balding DJ, Tyler-Smith C,. 2012. Ethiopian genetic diversity reveals linguistic stratification and complex influences on the Ethiopian gene pool. Am J Hum Genet 91: 83-96.
63. Pagani L, Schiffels S, Gurdasani D, Danecek P, Scally A, Chen Y, Xue Y, Haber M, Ekong R, Oljira T, Mekonnen E, Luiselli D, Bradman N, Bekele E, Zalloua P, Durbin R, Kivisild T, Tyler-Smith C,. 2015. Tracing the route of modern humans out of Africa by using 225 human genome sequences from Ethiopians and Egyptians. Am J Hum Genet 96: 986-991.
64. Peakall R, Smouse PE., 2012. GenAlEx 6.5: genetic analysis in Excel. Population genetic software for teaching and research-an update. Bioinformatics 28: 2537-2539.
65. Pennarun E, Kivisild T, Metspalu E, Metspalu M, Reisberg T, Moisan JP, Behar DM, Jones SC, Villems R., 2012. Divorcing the Late Upper Palaeolithic demographic histories of mtDNA haplogroups M1 and U6 in Africa. BMC Evol Biol 12: 234.
66. Pereira L, Freitas F, Fernandes V, Pereira JB, Costa MD, Costa S, Maximo V, Macaulay V, Rocha R, Samuels DC., 2009. The diversity present in 5140 human mitochondrial genomes. Am J Hum Genet 84: 628-640.
67. Petraglia MD, and, Rose JI., 2009. The evolution of human populations in Arabia. Paleoenvironments, prehistory and genetics. New York: Springer.
68. Poloni ES, Naciri Y, Bucho R, Niba R, Kervaire B, Excoffier L, Langaney A, Sanchez-Mazas A., 2009. Genetic evidence for complexity in ethnic differentiation and history in East Africa. Ann Hum Genet 73: 582-600.
69. Poloni ES, Semino O, Passarino G, Santachiara-Benerecetti AS, Dupanloup I, Langaney A, Excoffier L., 1997. Human genetic affinities for Y-chromosome P49a,f/TaqI haplotypes show strong correspondence with linguistics. Am J Hum Genet 61: 1015-1035.
70. R Core Team. 2013. R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. Available at: http://www.R-project.org/.
71. Ramos-Onsins SE, Rozas J., 2002. Statistical properties of new neutrality tests against population growth. Mol Biol Evol 19: 2092-2100.
72. Reynolds J, Weir BS, Cockerham CC., 1983. Estimation of the coancestry coefficient: basis for a short-term genetic distance. Genetics 105: 767-779.
73. Rídl J, Edens CM, and, Černý V., 2009. Mitochondrial DNA structure of Yemeni population: regional differences and the implications for different migratory contributions. In:, Petraglia MD, Rose JI, editors. The evolution of human populations in Arabia Paleoenvironments, prehistory and genetics. New York: Springer. p 69-78.
74. Richards M, Rengo C, Cruciani F, Gratrix F, Wilson JF, Scozzari R, Macaulay V, Torroni A., 2003. Extensive female-mediated gene flow from sub-Saharan Africa into near eastern Arab populations. Am J Hum Genet 72: 1058-1064.
75. Rose JI., 2010. New light on human prehistory in the Persian Gulf oasis. Curr Anthropol 51: 849-883.
76. Rose JI, Marks AE., 2014. " Out of Arabia" and the Middle-Upper Palaeolithic transition in the southern Levant. Quartär 61: 49-85.
77. Rose JI, Usik VI, Marks AE, Hilbert YH, Galletti CS, Parton A, Geiling JM, Černý V, Morley MW, Roberts RG., 2011. The Nubian complex of Dhofar, Oman: an African Middle stone age industry in southern Arabia. PLoS One 6: e28239.
78. Santos C, Fregel R, Cabrera VM, Gonzalez AM, Larruga JM, Lima M., 2010. Mitochondrial DNA patterns in the Macaronesia islands: variation within and among archipelagos. Am J Phys Anthropol 141: 610-619.
79. Scally A, Durbin R., 2012. Revising the human mutation rate: implications for understanding human evolution. Nat Rev Genet 13: 745-753.
80. Seielstad MT, Minch E, Cavalli-Sforza LL., 1998. Genetic evidence for a higher female migration rate in humans. Nat Genet 20: 278-280.
81. Soares P, Alshamali F, Pereira JB, Fernandes V, Silva NM, Afonso C, Costa MD, Musilová E, Macaulay V, Richards MB, Černý V, Pereira L,. 2012. The expansion of mtDNA haplogroup L3 within and out of Africa. Mol Biol Evol 29: 915-927.
82. Tajima F., 1989. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics 123: 585-595.
83. Tishkoff SA, Dietzsch E, Speed W, Pakstis AJ, Kidd JR, Cheung K, Bonne-Tamir B, Santachiara-Benerecetti AS, Moral P, Krings M., 1996. Global patterns of linkage disequilibrium at the CD4 locus and modern human origins. Science 271: 1380-1387.
84. Tishkoff SA, Williams SM., 2002. Genetic analysis of African populations: human evolution and complex disease. Nat Rev Genet 3: 611-621.
85. van Oven M, Kayser M., 2009. Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation. Hum Mutat 30: E386-E394.
86. Vyas D, Kitchen A, Miró-Herrans AT, Pearson LN, Al-Meeri A., Mulligan CJ,. 2015. Bayesian Analyses of Yemeni Mitochondrial Genomes Suggest Multiple Migration Events With Africa and Western Eurasia. Am J Phys Anthropol Early View.