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Table of Contents
Vol. 61, No. 3, 2006
Issue release date: August 2006
Hum Hered 2006;61:132–143
(DOI:10.1159/000093774)

Iran: Tricontinental Nexus for Y-Chromosome Driven Migration

Regueiro M.a · Cadenas A.M.a · Gayden T.a · Underhill P.A.b · Herrera R.J.a
aDepartment of Biological Sciences, Florida International University, Miami, Fla., and bDepartment of Genetics, Stanford University, Stanford, Calif., USA
email Corresponding Author

Abstract

Due to its pivotal geographic position, present day Iran likely served as a gateway of reciprocal human movements. However, the extent to which the deserts within the Iranian plateau and the mountain ranges surrounding Persia inhibited gene flow via this corridor remains uncertain. In order to assess the magnitude of this region’s role as a nexus for Africa, Asia and Europe in human migrations, high-resolution Y-chromosome analyses were performed on 150 Iranian males. Haplogroup data were subsequently compared to regional populations characterized at similar phylogenetic levels. The Iranians display considerable haplogroup diversity consistent with patterns observed in populations of the Middle East overall, reinforcing the notion of Persia as a venue for human disseminations. Admixture analyses of geographically targeted, regional populations along the latitudinal corridor spanning from Anatolia to the Indus Valley demonstrated contributions to Persia from both the east and west. However, significant differences were uncovered upon stratification of the gene donors, including higher proportions from central east and southeast Turkey as compared to Pakistan. In addition to the modulating effects of geographic obstacles, culturally mediated amalgamations consistent with the diverse spectrum of a variety of historical empires may account for the distribution of haplogroups and lineages observed. Our study of high-resolution Y-chromosome genotyping allowed for an in-depth analysis unattained in previous studies of the area, revealing important migratory and demographic events that shaped the contemporary genetic landscape.


 goto top of outline Key Words

  • Iran
  • Y-chromosome
  • SNP

 goto top of outline Abstract

Due to its pivotal geographic position, present day Iran likely served as a gateway of reciprocal human movements. However, the extent to which the deserts within the Iranian plateau and the mountain ranges surrounding Persia inhibited gene flow via this corridor remains uncertain. In order to assess the magnitude of this region’s role as a nexus for Africa, Asia and Europe in human migrations, high-resolution Y-chromosome analyses were performed on 150 Iranian males. Haplogroup data were subsequently compared to regional populations characterized at similar phylogenetic levels. The Iranians display considerable haplogroup diversity consistent with patterns observed in populations of the Middle East overall, reinforcing the notion of Persia as a venue for human disseminations. Admixture analyses of geographically targeted, regional populations along the latitudinal corridor spanning from Anatolia to the Indus Valley demonstrated contributions to Persia from both the east and west. However, significant differences were uncovered upon stratification of the gene donors, including higher proportions from central east and southeast Turkey as compared to Pakistan. In addition to the modulating effects of geographic obstacles, culturally mediated amalgamations consistent with the diverse spectrum of a variety of historical empires may account for the distribution of haplogroups and lineages observed. Our study of high-resolution Y-chromosome genotyping allowed for an in-depth analysis unattained in previous studies of the area, revealing important migratory and demographic events that shaped the contemporary genetic landscape.

Copyright © 2006 S. Karger AG, Basel


 goto top of outline References
  1. Cavalli-Sforza LL, Menozzi P, Piazza A: The history and geography of human genes. Princeton University Press, Princeton, NJ, 1994.
  2. Cinnioğlu C, King R, Kvisild T, Kalfoglu E, Atasoy S, Cavalleri GL, Lillie AS, Roseman CC, Lin AA, Prince K, Oefner PJ, Shen P, Semino O, Cavalli-Sforza LL, Underhill PA: Excavating Y-chromosome haplotype strata in Anatolia. Hum Genet 2004;114:127–148.
  3. Nasidze I, Ling EYS, Quinque D, Dupanloup I, Cordaux R, Rychkov S, Naumova O, Zhukova O, Sarraf-Zadegan N, Naderi GA, Asgary S, Sardas S, Farhud DD, Sarkisian T, Asadov C, Kerimov A, Stoneking M: Mitochondrial DNA and Y-chromosome variation in the Caucasus. Ann Hum Genet 2004;68:205–221.
  4. Luis JR, Rowold DJ, Regueiro M, Caeiro B, Cinnioğlu C, Roseman C, Underhill PA, Cavalli-Sforza LL, Herrera RJ: The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations. Am J Hum Genet 2004;74:32–544.

    External Resources

  5. Quintana-Murci L, Semino O, Bandelt HJ, Passarino G, McElreavy K, Santachiara-Benerecetti AS: Genetic evidence of an early exit of Homo sapiens sapiens from Africa through eastern Africa. Nat Genet 1999;23:437–441.
  6. Stringer C: Coasting out of Africa. Nature 2000;405:24–25, 27.
  7. Underhill PA, Passarino G, Lin AA, Shen P, Mirazon Lahr M, Foley RA, Oefner PJ, Cavalli-Sforza LL: The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations. Ann Hum Genet 2001;65:43–62.
  8. Kivisild T, Bamshad MJ, Kaldma K, Metspalu M, Metspalu E, Reidla M, Laos S, Parik J, Watkins WS, Dixon ME, Papiha SS, Mastana SS, Mir MR, Ferak V, Villems R: Deep common ancestry of Indian and western-Eurasian mitochondrial DNA lineages. Curr Biol 1999;9:1331–1334.
  9. Quintana-Murci L, Krausz C, Zerjal T, Sayar SH, Hammer MF, Mehdi SQ, Ayub Q, Qamar R, Mohyuddin A, Radhakrishna U, Jobling MA, Tyler-Smith C, McElreavey K: Y-chromosome lineages trace diffusion of people and languages in southwestern Asia. Am J Hum Genet 2001;68:537–542.
  10. Wells RS, Yuldasheva N, Ruzibakiev R, Underhill PA, Evseeva I, Blue-Smith J, Jin L, Bing S, Pitchappan R, Shanmugalakshmi S, Balakrishnan K, Read M, Pearson NM, Zerjal T, Webster MT, Zholoshvili I, Jamarjashvili E, Gambarov S, Nikbin B, Dostiev A, Aknazarov O, Zalloua P, Tsoy I, Kitaev M, Mirrakhimov M, Chariev A, Bodmer WF: The Eurasian heartland: A continental perspective on Y-chromosome diversity. Proc Natl Acad Sci USA 2001;98:10244–10249.
  11. Qamar R, Ayub Q, Mohyuddin A, Helgason A, Mazhar K, Mansoor A, Zerjal T, Tyler-Smith C, Mehdi SQ: Y chromosomal DNA variation in Pakistan. Am J Hum Genet 2002;70:1107–1124.
  12. Quintana-Murci L, Chaix R, Wells RS, Behar DM, Sayar H, Scozzari R, Rengo C, Al-Zahery N, Semino O, Santachiara-Benerecetti AS, Coppa A, Ayub Q, Mohyuddin A, Tyler-Smith C, Mehdi SQ, Torroni A, McElreavey K: Where west meets east: the complex mtDNA landscape of the southwest and central Asian corridor. Am J Hum Genet 2004;74:827–845.
  13. Antunez de Mayolo G, Antunez de Mayolo A, Antunez de Mayolo P, Papiha SS, Hammer M, Yunis JJ, Yunis EJ, Damodaran C, Martinez de Pancorbo M, Caeiro JL, Puzyrev VP, Herrera RJ: Phylogenetics of worldwide human populations as determined by polymorphic Alu insertions. Electrophoresis 2002;23:3346–3356.
  14. Shen P, Lavi T, Kivisild T, Chou V, Sengun D, Gefel D, Shpirer I, Woolf E, Hillel J, Feldman MW, Oefner PJ: Reconstruction of patrilineages and matrilineages of Samaritans and other Israeli populations from Y-chromosome and mitochondrial DNA sequence variation. Hum Mutat 2004;24:248–260.
  15. Sengupta S, Zhivotovsky LA, King R, Mehdi SQ, Edmonds CA, Chow CT, Lin AA, Mitra M, Sil SK, Ramesh A, Usha Rani MV, Thakur CM, Cavalli-Sforza LL, Majumder PP, Underhill PA: Polarity and temporality of high resolution Y-chromosome distributions in India identify both indigenous and exogenous expansions and reveal minor genetic influence of Central Asian pastoralists. Am J Hum Genet 2006;78:202–221.
  16. Martinez L, Reategui EP, Fonseca LR, Sierra-Montes JM, Terreros MC, Pereira-Simon S, Herrera RJ: Superimposing polymorphism: The case of a point mutation within a polymorphic Alu insertion. Hum Hered 2005;59:109–117.
  17. Hammer MF, Horai S: Y chromosomal DNA variation and the peopling of Japan. Am J Hum Genet 1995;56:951–962.
  18. Y Chromosome Consortium: A nomenclature system for the tree of human Y-chromosomal binary haplogroups. Genome Res 2002;12:339–348.
  19. Excoffier L, Smouse PE, Quattro JM: Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics 1992;131:479–491.
  20. Felsenstein J: PHYLIP-phylogeny inference package (version 3.2). Cladistics 1989;5:164–166.
  21. Rohlf F: NTSTSpc. Exter Publishing, Setauket, NY, 2002.
  22. Dupanloup I, Bertorelle G: Inferring admixture proportions from molecular data: extension to any number of parental populations. Mol Biol Evol 2001;18:672–675.
  23. Shepard EM, Herrera RJ: Iranian STR variation at the fringes of biogeographical demarcation. For Sci Int 2005;158:140–148.
  24. Nasidze I, Sarkisian T, Kerimov A, Stoneking M: Testing hypotheses of language replacement in the Caucasus: Evidence from the Y-chromosome. Hum Genet 2003;112:255–261.
  25. Passarino G, Semino O, Magri C, Al-Zahery N, Benuzzi G, Quintana-Murci L, Andellnovic S, Bullc-Jakus F, Liu A, Arslan A, Santachiara-Benerecetti AS: The 49a,f haplotype 11 is a new marker of the EU19 lineage that traces migrations from northern regions of the Black Sea. Hum Immunol 2001;62:922–932.
  26. Mallory JP: In search of the Indo-Europeans: Language, archaeology and myth. Thames and Hudson, London, 1989.
  27. Semino O, Passarino G, Oefner PJ, Lin AA, Arbuzova S, Beckman LE, De Benedictis G, Francalacci P, Kouvatsi A, Limborska S, Marcikiae M, Mika A, Mika B, Primorac D, Santachiara-Benerecetti AS, Cavalli-Sforza LL, Underhill P: The genetic legacy of Paleolithic Homo sapiens sapiens in extant Europeans: A Y chromosome perspective. Science 2000;290:1155–1159.
  28. Kivisild T, Rootsi S, Metspalu M, Mastana S, Kaldma K, Parik J, Metspalu E, Adojaan M, Tolk HV, Stepanov V, Golge M, Usanga E, Papiha SS, Cinnioğlu C, King R, Cavalli-Sforza L, Underhill PA, Villems R: The genetic heritage of the earliest settlers persists both in Indian tribal and caste populations. Am J Hum Genet 2003;72:313–332.
  29. Al-Zahery N, Semino O, Benuzzi G, Magri C, Passarino G, Torroni A, Santachiara-Benerecetti AS: Y-chromosome and mtDNA polymorphisms in Iraq, a crossroad of the early human dispersal and of post-Neolithic migrations. Mol Phylogenet Evol 2003;28:458–472.
  30. Hammer MF, Karafet T, Rasanayagam A, Wood ET, Altheide TK, Jenkins T, Griffiths RC, Templeton AR, Zegura SL: Out of Africa and back again: nested cladistic analysis of human Y chromosome variation. Mol Biol Evol 1998;15:427–441.
  31. Passarino G, Semino O, Quintana-Murci L, Excoffier L, Hammer M, Santachiara-Benerecetti AS: Different genetic components in the Ethiopian population, identified by mtDNA and Y-chromosome polymorphisms. Am J Hum Genet 1998;62:420–434.
  32. Scozzari R, Cruciani F, Santolamazza P, Malaspina P, Torroni A, Sellitto D, Arredi B, Destro-Bisol G, De Stefano G, Rickards O, Martinez-Labarga C, Modiano D, Biondi G, Moral P, Olckers A, Wallace DC, Novelletto A: Combined use of biallelic and microsatellite Y-chromosome polymorphisms to infer affinities among African populations. Am J Hum Genet 1999;65:829–846.
  33. Semino O, Magri C, Benuzzi G, Lin AA, Al-Zahery N, Battaglia V, Maccioni L, Triantaphyllidis C, Shen P, Oefner PJ, Zhivotovsky LA, King R, Torroni A, Cavalli-Sforza LL, Underhill P, Santachiara-Benerecetti AS: Origin, diffusion, and differentiation of Y-chromosome haplogroups E and J: Inferences on the neolithization of Europe and later migratory events in the Mediterranean area. Am J Hum Genet 2004;74:1023–1034.
  34. Cruciani F, Santolamazza P, Shen P, Macauley V, Moral P, Olckers A, Modiano D, Destro-Bisol G, Coia V, Wallace DC, Oefner PJ, Torroni A, Cavalli-Sforza LL, Scozzari R, Underhill P: A back migration from Asia to sub-saharan Africa is supported by high resolution analysis of human Y-chromosome haplotypes. Am J Hum Genet 2002;70:1197–1214.
  35. Qamar R, Ayub Q, Mohyuddin A, Helgason A, Mazhar K, Mansoor A, Zerjal T, Tyler-Smith C, Mehdi SQ: Y-chromosomal DNA variation in Pakistan. Am J Hum Genet 2002;70:1107–1124.
  36. ‘Janissaries.’ The Columbia Encyclopedia, 6th ed. New York: Columbia University Press, 2001–04. Retrieved November 14, 2005, from www.bartleby.com/65.
  37. Arredi B, Poloni ES, Paracchini S, Zerjal T, Fathallah DM, Makrelouf M, Pascali VL, Novelletto A, Tyler-Smith C: A predominantly Neolithic origin for Y-chromosomal DNA variation in North Africa. Am J Hum Genet 2004;75:338–345.
  38. Sanchez JJ, Hallenberg C, Borsting C, Hernandez A, Morling N: High frequencies of Y chromosome lineages characterized by E3b1, DYS19-11, DYS392-12 in Somali males. Eur J Hum Genet 2005;13:856–866.
  39. Semino O, Santachiara-Benerecetti AS, Falaschi F, Cavalli-Sforza LL, Underhill PA: Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny. Am J Hum Genet 2002;70:265–268.

 goto top of outline Author Contacts

Dr. Rene J. Herrera
Department of Biological Sciences, Florida International University
University Park, OE 304
Miami, FL 33199 (USA)
Tel. +1 305 348 1258, Fax +1 305 348 1259, E-Mail herrerar@fiu.edu


 goto top of outline Article Information

M. Regueiro and A.M. Cadenas contributed equally to the article.

Received: December 12, 2005
Accepted: March 23, 2006
Published online: June 12, 2006
Number of Print Pages : 12
Number of Figures : 5, Number of Tables : 3, Number of References : 39


 goto top of outline Publication Details

Human Heredity (International Journal of Human and Medical Genetics)

Vol. 61, No. 3, Year 2006 (Cover Date: August 2006)

Journal Editor: Devoto, M. (Philadelphia, Pa.)
ISSN: 0001–5652 (print), 1423–0062 (Online)

For additional information: http://www.karger.com/HHE


Copyright / Drug Dosage / Disclaimer

Copyright: All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, microcopying, or by any information storage and retrieval system, without permission in writing from the publisher or, in the case of photocopying, direct payment of a specified fee to the Copyright Clearance Center.
Drug Dosage: The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing research, changes in goverment regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any changes in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new and/or infrequently employed drug.
Disclaimer: The statements, opinions and data contained in this publication are solely those of the individual authors and contributors and not of the publishers and the editor(s). The appearance of advertisements or/and product references in the publication is not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or products referred to in the content or advertisements.

Abstract

Due to its pivotal geographic position, present day Iran likely served as a gateway of reciprocal human movements. However, the extent to which the deserts within the Iranian plateau and the mountain ranges surrounding Persia inhibited gene flow via this corridor remains uncertain. In order to assess the magnitude of this region’s role as a nexus for Africa, Asia and Europe in human migrations, high-resolution Y-chromosome analyses were performed on 150 Iranian males. Haplogroup data were subsequently compared to regional populations characterized at similar phylogenetic levels. The Iranians display considerable haplogroup diversity consistent with patterns observed in populations of the Middle East overall, reinforcing the notion of Persia as a venue for human disseminations. Admixture analyses of geographically targeted, regional populations along the latitudinal corridor spanning from Anatolia to the Indus Valley demonstrated contributions to Persia from both the east and west. However, significant differences were uncovered upon stratification of the gene donors, including higher proportions from central east and southeast Turkey as compared to Pakistan. In addition to the modulating effects of geographic obstacles, culturally mediated amalgamations consistent with the diverse spectrum of a variety of historical empires may account for the distribution of haplogroups and lineages observed. Our study of high-resolution Y-chromosome genotyping allowed for an in-depth analysis unattained in previous studies of the area, revealing important migratory and demographic events that shaped the contemporary genetic landscape.



 goto top of outline Author Contacts

Dr. Rene J. Herrera
Department of Biological Sciences, Florida International University
University Park, OE 304
Miami, FL 33199 (USA)
Tel. +1 305 348 1258, Fax +1 305 348 1259, E-Mail herrerar@fiu.edu


 goto top of outline Article Information

M. Regueiro and A.M. Cadenas contributed equally to the article.

Received: December 12, 2005
Accepted: March 23, 2006
Published online: June 12, 2006
Number of Print Pages : 12
Number of Figures : 5, Number of Tables : 3, Number of References : 39


 goto top of outline Publication Details

Human Heredity (International Journal of Human and Medical Genetics)

Vol. 61, No. 3, Year 2006 (Cover Date: August 2006)

Journal Editor: Devoto, M. (Philadelphia, Pa.)
ISSN: 0001–5652 (print), 1423–0062 (Online)

For additional information: http://www.karger.com/HHE


Copyright / Drug Dosage

Copyright: All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized in any form or by any means, electronic or mechanical, including photocopying, recording, microcopying, or by any information storage and retrieval system, without permission in writing from the publisher or, in the case of photocopying, direct payment of a specified fee to the Copyright Clearance Center.
Drug Dosage: The authors and the publisher have exerted every effort to ensure that drug selection and dosage set forth in this text are in accord with current recommendations and practice at the time of publication. However, in view of ongoing research, changes in goverment regulations, and the constant flow of information relating to drug therapy and drug reactions, the reader is urged to check the package insert for each drug for any changes in indications and dosage and for added warnings and precautions. This is particularly important when the recommended agent is a new and/or infrequently employed drug.
Disclaimer: The statements, opinions and data contained in this publication are solely those of the individual authors and contributors and not of the publishers and the editor(s). The appearance of advertisements or/and product references in the publication is not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any ideas, methods, instructions or products referred to in the content or advertisements.

References

  1. Cavalli-Sforza LL, Menozzi P, Piazza A: The history and geography of human genes. Princeton University Press, Princeton, NJ, 1994.
  2. Cinnioğlu C, King R, Kvisild T, Kalfoglu E, Atasoy S, Cavalleri GL, Lillie AS, Roseman CC, Lin AA, Prince K, Oefner PJ, Shen P, Semino O, Cavalli-Sforza LL, Underhill PA: Excavating Y-chromosome haplotype strata in Anatolia. Hum Genet 2004;114:127–148.
  3. Nasidze I, Ling EYS, Quinque D, Dupanloup I, Cordaux R, Rychkov S, Naumova O, Zhukova O, Sarraf-Zadegan N, Naderi GA, Asgary S, Sardas S, Farhud DD, Sarkisian T, Asadov C, Kerimov A, Stoneking M: Mitochondrial DNA and Y-chromosome variation in the Caucasus. Ann Hum Genet 2004;68:205–221.
  4. Luis JR, Rowold DJ, Regueiro M, Caeiro B, Cinnioğlu C, Roseman C, Underhill PA, Cavalli-Sforza LL, Herrera RJ: The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations. Am J Hum Genet 2004;74:32–544.

    External Resources

  5. Quintana-Murci L, Semino O, Bandelt HJ, Passarino G, McElreavy K, Santachiara-Benerecetti AS: Genetic evidence of an early exit of Homo sapiens sapiens from Africa through eastern Africa. Nat Genet 1999;23:437–441.
  6. Stringer C: Coasting out of Africa. Nature 2000;405:24–25, 27.
  7. Underhill PA, Passarino G, Lin AA, Shen P, Mirazon Lahr M, Foley RA, Oefner PJ, Cavalli-Sforza LL: The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations. Ann Hum Genet 2001;65:43–62.
  8. Kivisild T, Bamshad MJ, Kaldma K, Metspalu M, Metspalu E, Reidla M, Laos S, Parik J, Watkins WS, Dixon ME, Papiha SS, Mastana SS, Mir MR, Ferak V, Villems R: Deep common ancestry of Indian and western-Eurasian mitochondrial DNA lineages. Curr Biol 1999;9:1331–1334.
  9. Quintana-Murci L, Krausz C, Zerjal T, Sayar SH, Hammer MF, Mehdi SQ, Ayub Q, Qamar R, Mohyuddin A, Radhakrishna U, Jobling MA, Tyler-Smith C, McElreavey K: Y-chromosome lineages trace diffusion of people and languages in southwestern Asia. Am J Hum Genet 2001;68:537–542.
  10. Wells RS, Yuldasheva N, Ruzibakiev R, Underhill PA, Evseeva I, Blue-Smith J, Jin L, Bing S, Pitchappan R, Shanmugalakshmi S, Balakrishnan K, Read M, Pearson NM, Zerjal T, Webster MT, Zholoshvili I, Jamarjashvili E, Gambarov S, Nikbin B, Dostiev A, Aknazarov O, Zalloua P, Tsoy I, Kitaev M, Mirrakhimov M, Chariev A, Bodmer WF: The Eurasian heartland: A continental perspective on Y-chromosome diversity. Proc Natl Acad Sci USA 2001;98:10244–10249.
  11. Qamar R, Ayub Q, Mohyuddin A, Helgason A, Mazhar K, Mansoor A, Zerjal T, Tyler-Smith C, Mehdi SQ: Y chromosomal DNA variation in Pakistan. Am J Hum Genet 2002;70:1107–1124.
  12. Quintana-Murci L, Chaix R, Wells RS, Behar DM, Sayar H, Scozzari R, Rengo C, Al-Zahery N, Semino O, Santachiara-Benerecetti AS, Coppa A, Ayub Q, Mohyuddin A, Tyler-Smith C, Mehdi SQ, Torroni A, McElreavey K: Where west meets east: the complex mtDNA landscape of the southwest and central Asian corridor. Am J Hum Genet 2004;74:827–845.
  13. Antunez de Mayolo G, Antunez de Mayolo A, Antunez de Mayolo P, Papiha SS, Hammer M, Yunis JJ, Yunis EJ, Damodaran C, Martinez de Pancorbo M, Caeiro JL, Puzyrev VP, Herrera RJ: Phylogenetics of worldwide human populations as determined by polymorphic Alu insertions. Electrophoresis 2002;23:3346–3356.
  14. Shen P, Lavi T, Kivisild T, Chou V, Sengun D, Gefel D, Shpirer I, Woolf E, Hillel J, Feldman MW, Oefner PJ: Reconstruction of patrilineages and matrilineages of Samaritans and other Israeli populations from Y-chromosome and mitochondrial DNA sequence variation. Hum Mutat 2004;24:248–260.
  15. Sengupta S, Zhivotovsky LA, King R, Mehdi SQ, Edmonds CA, Chow CT, Lin AA, Mitra M, Sil SK, Ramesh A, Usha Rani MV, Thakur CM, Cavalli-Sforza LL, Majumder PP, Underhill PA: Polarity and temporality of high resolution Y-chromosome distributions in India identify both indigenous and exogenous expansions and reveal minor genetic influence of Central Asian pastoralists. Am J Hum Genet 2006;78:202–221.
  16. Martinez L, Reategui EP, Fonseca LR, Sierra-Montes JM, Terreros MC, Pereira-Simon S, Herrera RJ: Superimposing polymorphism: The case of a point mutation within a polymorphic Alu insertion. Hum Hered 2005;59:109–117.
  17. Hammer MF, Horai S: Y chromosomal DNA variation and the peopling of Japan. Am J Hum Genet 1995;56:951–962.
  18. Y Chromosome Consortium: A nomenclature system for the tree of human Y-chromosomal binary haplogroups. Genome Res 2002;12:339–348.
  19. Excoffier L, Smouse PE, Quattro JM: Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics 1992;131:479–491.
  20. Felsenstein J: PHYLIP-phylogeny inference package (version 3.2). Cladistics 1989;5:164–166.
  21. Rohlf F: NTSTSpc. Exter Publishing, Setauket, NY, 2002.
  22. Dupanloup I, Bertorelle G: Inferring admixture proportions from molecular data: extension to any number of parental populations. Mol Biol Evol 2001;18:672–675.
  23. Shepard EM, Herrera RJ: Iranian STR variation at the fringes of biogeographical demarcation. For Sci Int 2005;158:140–148.
  24. Nasidze I, Sarkisian T, Kerimov A, Stoneking M: Testing hypotheses of language replacement in the Caucasus: Evidence from the Y-chromosome. Hum Genet 2003;112:255–261.
  25. Passarino G, Semino O, Magri C, Al-Zahery N, Benuzzi G, Quintana-Murci L, Andellnovic S, Bullc-Jakus F, Liu A, Arslan A, Santachiara-Benerecetti AS: The 49a,f haplotype 11 is a new marker of the EU19 lineage that traces migrations from northern regions of the Black Sea. Hum Immunol 2001;62:922–932.
  26. Mallory JP: In search of the Indo-Europeans: Language, archaeology and myth. Thames and Hudson, London, 1989.
  27. Semino O, Passarino G, Oefner PJ, Lin AA, Arbuzova S, Beckman LE, De Benedictis G, Francalacci P, Kouvatsi A, Limborska S, Marcikiae M, Mika A, Mika B, Primorac D, Santachiara-Benerecetti AS, Cavalli-Sforza LL, Underhill P: The genetic legacy of Paleolithic Homo sapiens sapiens in extant Europeans: A Y chromosome perspective. Science 2000;290:1155–1159.
  28. Kivisild T, Rootsi S, Metspalu M, Mastana S, Kaldma K, Parik J, Metspalu E, Adojaan M, Tolk HV, Stepanov V, Golge M, Usanga E, Papiha SS, Cinnioğlu C, King R, Cavalli-Sforza L, Underhill PA, Villems R: The genetic heritage of the earliest settlers persists both in Indian tribal and caste populations. Am J Hum Genet 2003;72:313–332.
  29. Al-Zahery N, Semino O, Benuzzi G, Magri C, Passarino G, Torroni A, Santachiara-Benerecetti AS: Y-chromosome and mtDNA polymorphisms in Iraq, a crossroad of the early human dispersal and of post-Neolithic migrations. Mol Phylogenet Evol 2003;28:458–472.
  30. Hammer MF, Karafet T, Rasanayagam A, Wood ET, Altheide TK, Jenkins T, Griffiths RC, Templeton AR, Zegura SL: Out of Africa and back again: nested cladistic analysis of human Y chromosome variation. Mol Biol Evol 1998;15:427–441.
  31. Passarino G, Semino O, Quintana-Murci L, Excoffier L, Hammer M, Santachiara-Benerecetti AS: Different genetic components in the Ethiopian population, identified by mtDNA and Y-chromosome polymorphisms. Am J Hum Genet 1998;62:420–434.
  32. Scozzari R, Cruciani F, Santolamazza P, Malaspina P, Torroni A, Sellitto D, Arredi B, Destro-Bisol G, De Stefano G, Rickards O, Martinez-Labarga C, Modiano D, Biondi G, Moral P, Olckers A, Wallace DC, Novelletto A: Combined use of biallelic and microsatellite Y-chromosome polymorphisms to infer affinities among African populations. Am J Hum Genet 1999;65:829–846.
  33. Semino O, Magri C, Benuzzi G, Lin AA, Al-Zahery N, Battaglia V, Maccioni L, Triantaphyllidis C, Shen P, Oefner PJ, Zhivotovsky LA, King R, Torroni A, Cavalli-Sforza LL, Underhill P, Santachiara-Benerecetti AS: Origin, diffusion, and differentiation of Y-chromosome haplogroups E and J: Inferences on the neolithization of Europe and later migratory events in the Mediterranean area. Am J Hum Genet 2004;74:1023–1034.
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