History of click-speaking populations of Africa inferred from mtDNA and Y chromosome genetic variation.

Little is known about the history of click-speaking populations in Africa. Prior genetic studies revealed that the click-speaking Hadza of eastern Africa are as distantly related to click speakers of southern Africa as are most other African populations. The Sandawe, who currently live within 150 km of the Hadza, are the only other population in eastern Africa whose language has been classified as part of the Khoisan language family. Linguists disagree on whether there is any detectable relationship between the Hadza and Sandawe click languages. We characterized both mtDNA and Y chromosome variation of the Sandawe, Hadza, and neighboring Tanzanian populations. New genetic data show that the Sandawe and southern African click speakers share rare mtDNA and Y chromosome haplogroups; however, common ancestry of the 2 populations dates back >35,000 years. These data also indicate that common ancestry of the Hadza and Sandawe populations dates back >15,000 years. These findings suggest that at the time of the spread of agriculture and pastoralism, the click-speaking populations were already isolated from one another and are consistent with relatively deep linguistic divergence among the respective click languages.

[1]  Ecological Studies in Southern Africa. , 1965 .

[2]  J. L. Newman The ecological basis for subsistence change among the Sandawe of Tanzania : foreign field research program , 1970 .

[3]  E. G. Pulleyblank Language and History. , 1972 .

[4]  G. Bräuer The morphological differentiation of anatomically modern man in Africa, with special regard to recent finds from East Africa. , 1978, Zeitschrift fur Morphologie und Anthropologie.

[5]  A Modern History of Tanganyika , 1979 .

[6]  E. D. Elderkin On the Classification of Hadza , 1982 .

[7]  C. Stringer,et al.  Preparation and further study of the Singa skull from Sudan , 1985 .

[8]  J. Weiner,et al.  The Peoples of Southern Africa and Their Affinities , 1986 .

[9]  M. Ruhlen A Guide to the World’s Languages , 1987 .

[10]  N. Saitou,et al.  The neighbor-joining method: a new method for reconstructing phylogenetic trees. , 1987, Molecular biology and evolution.

[11]  A Piazza,et al.  Reconstruction of human evolution: bringing together genetic, archaeological, and linguistic data. , 1988, Proceedings of the National Academy of Sciences of the United States of America.

[12]  S. Pääbo,et al.  Extensive mitochondrial diversity within a single Amerindian tribe. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[13]  K. Hawkes,et al.  African populations and the evolution of human mitochondrial DNA. , 1991, Science.

[14]  Ken Hale,et al.  Language endangerment and the human value of linguistic diversity , 2015 .

[15]  Colin Renfrew,et al.  ARCHAEOLOGY, GENETICS AND LINGUISTIC DIVERSITY* , 1992 .

[16]  K. Hawkes,et al.  Demography of the Hadza, an increasing and high density population of Savanna foragers. , 1992, American journal of physical anthropology.

[17]  Hunters and Herders of Southern Africa: The G/wi and G//ana of the central Kalahari , 1992 .

[18]  Richard Lee,et al.  The Dobe Ju/'Hoansi , 1993 .

[19]  R. Cann The history and geography of human genes , 1995, The Journal of Asian Studies.

[20]  A. Piazza History and Geography of Human Genes , 1994 .

[21]  M W Feldman,et al.  Genetic absolute dating based on microsatellites and the origin of modern humans. , 1995, Proceedings of the National Academy of Sciences of the United States of America.

[22]  N. Takahata,et al.  Recent African origin of modern humans revealed by complete sequences of hominoid mitochondrial DNAs. , 1995, Proceedings of the National Academy of Sciences of the United States of America.

[23]  D. F. Roberts,et al.  The History and Geography of Human Genes , 1996 .

[24]  H. Bandelt,et al.  Origin and evolution of Native American mtDNA variation: a reappraisal. , 1996, American journal of human genetics.

[25]  David S. G. Thomas,et al.  Multiple episodes of aridity in southern Africa since the last interglacial period , 1997, Nature.

[26]  S. Tavaré,et al.  Computational Methods for the Coalescent , 1997 .

[27]  P. Donnelly,et al.  Progress in population genetics and human evolution , 1997 .

[28]  H. Bandelt,et al.  Mitochondrial footprints of human expansions in Africa. , 1997, American journal of human genetics.

[29]  Language, identity, and conceptualization among the Khoisan , 1998 .

[30]  Language, Identity, and Conceptualization among the Khoisan , 1998 .

[31]  R. Foley,et al.  Towards a theory of modern human origins: geography, demography, and diversity in recent human evolution. , 1998, American journal of physical anthropology.

[32]  A. von Haeseler,et al.  Pattern of nucleotide substitution and rate heterogeneity in the hypervariable regions I and II of human mtDNA. , 1999, Genetics.

[33]  M. Feldman,et al.  Population growth of human Y chromosomes: a study of Y chromosome microsatellites. , 1999, Molecular biology and evolution.

[34]  H. Bandelt,et al.  Median-joining networks for inferring intraspecific phylogenies. , 1999, Molecular biology and evolution.

[35]  A. Torroni,et al.  Combined use of biallelic and microsatellite Y-chromosome polymorphisms to infer affinities among African populations. , 1999, American journal of human genetics.

[36]  Li Jin,et al.  Y chromosome sequence variation and the history of human populations , 2000, Nature Genetics.

[37]  B Brinkmann,et al.  A short tandem repeat-based phylogeny for the human Y chromosome. , 2000, American journal of human genetics.

[38]  John P. Hutchison,et al.  African Languages: An Introduction , 2000 .

[39]  M. Hitchcock,et al.  The Peopling of Africa , 2000 .

[40]  L. Excoffier,et al.  SIMCOAL: a general coalescent program for the simulation of molecular data in interconnected populations with arbitrary demography. , 2000, The Journal of heredity.

[41]  D. Wallace,et al.  mtDNA variation in the South African Kung and Khwe-and their genetic relationships to other African populations. , 2000, American journal of human genetics.

[42]  Bernard Comrie Is there a single time depth cut-off point in historical linguistics? , 2000 .

[43]  A. Torroni,et al.  Prehistoric and historic traces in the mtDNA of Mozambique: insights into the Bantu expansions and the slave trade. , 2001, Annals of human genetics.

[44]  P. Forster,et al.  Phylogenetic star contraction applied to Asian and Papuan mtDNA evolution. , 2001, Molecular biology and evolution.

[45]  K. Crandall,et al.  Intraspecific gene genealogies: trees grafting into networks. , 2001, Trends in ecology & evolution.

[46]  R. Nielsen,et al.  Distinguishing migration from isolation: a Markov chain Monte Carlo approach. , 2001, Genetics.

[47]  L. Excoffier,et al.  An extensive analysis of Y-chromosomal microsatellite haplotypes in globally dispersed human populations. , 2001, American journal of human genetics.

[48]  L L Cavalli-Sforza,et al.  The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations , 2001, Annals of human genetics.

[49]  Scott M. Williams,et al.  Genetic analysis of African populations: human evolution and complex disease , 2002, Nature Reviews Genetics.

[50]  P. Underhill,et al.  Ethiopians and Khoisan share the deepest clades of the human Y-chromosome phylogeny. , 2002, American journal of human genetics.

[51]  Á. Carracedo,et al.  The making of the African mtDNA landscape. , 2002, American journal of human genetics.

[52]  P. Underhill,et al.  A back migration from Asia to sub-Saharan Africa is supported by high-resolution analysis of human Y-chromosome haplotypes. , 2002, American journal of human genetics.

[53]  A. Morris Isolation and the origin of the khoisan: Late pleistocene and early holocene human evolution at the southern end of Africa , 2002 .

[54]  M. Feldman,et al.  Genetic Structure of Human Populations , 2002, Science.

[55]  Jonathan Scott Friedlaender,et al.  A Human Genome Diversity Cell Line Panel , 2002, Science.

[56]  P. Mitchell The Archaeology of Southern Africa , 2004 .

[57]  Siavash Vahdati Daneshmand,et al.  On Steiner trees and minimum spanning trees in hypergraphs , 2003, Oper. Res. Lett..

[58]  Daniel S. Myers,et al.  Necessity is the mother of invention: a simple grid computing system using commodity tools , 2003, J. Parallel Distributed Comput..

[59]  C. Habel,et al.  Language , 1931, NeuroImage.

[60]  Merritt Ruhlen,et al.  African Y Chromosome and mtDNA Divergence Provides Insight into the History of Click Languages , 2003, Current Biology.

[61]  R. Villems,et al.  Ethiopian mitochondrial DNA heritage: tracking gene flow across and around the gate of tears. , 2004, American journal of human genetics.

[62]  Paul Proulx,et al.  Time depth in historical linguistics , 2004 .

[63]  P. Underhill,et al.  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. , 2004, American journal of human genetics.

[64]  R. Villems,et al.  MtDNA Profile of West Africa Guineans: Towards a Better Understanding of the Senegambia Region , 2004, Annals of human genetics.

[65]  C. Capelli,et al.  Variation of female and male lineages in sub-Saharan populations: the importance of sociocultural factors. , 2004, Molecular biology and evolution.

[66]  M. Hasegawa,et al.  Toward a more accurate time scale for the human mitochondrial DNA tree , 1993, Journal of Molecular Evolution.

[67]  J. Wakeley Substitution rate variation among sites in hypervariable region 1 of human mitochondrial DNA , 1993, Journal of Molecular Evolution.

[68]  Giovanni Destro-Bisol,et al.  The effective mutation rate at Y chromosome short tandem repeats, with application to human population-divergence time. , 2004, American journal of human genetics.

[69]  V. Pascali,et al.  The Analysis of Variation of mtDNA Hypervariable Region 1 Suggests That Eastern and Western Pygmies Diverged before the Bantu Expansion , 2004, The American Naturalist.

[70]  R. J. Herrera,et al.  The Levant versus the Horn of Africa: evidence for bidirectional corridors of human migrations. , 2004, American journal of human genetics.

[71]  A. Torroni,et al.  Phylogeographic analysis of haplogroup E3b (E-M215) y chromosomes reveals multiple migratory events within and out of Africa. , 2004, American journal of human genetics.

[72]  R. Nielsen,et al.  Multilocus Methods for Estimating Population Sizes, Migration Rates and Divergence Time, With Applications to the Divergence of Drosophila pseudoobscura and D. persimilis , 2004, Genetics.

[73]  Sonia Sharama,et al.  Grid Computing , 2004, Lecture Notes in Computer Science.

[74]  Alfredo Coppa,et al.  The African diaspora: mitochondrial DNA and the Atlantic slave trade. , 2004, American journal of human genetics.

[75]  Giovanni Destro-Bisol,et al.  Contrasting patterns of Y chromosome and mtDNA variation in Africa: evidence for sex-biased demographic processes , 2005, European Journal of Human Genetics.

[76]  S. Tishkoff,et al.  African human diversity, origins and migrations. , 2006, Current opinion in genetics & development.

[77]  Daniel S. Myers,et al.  Grid Services Base Library: A high-level, procedural application programming interface for writing Globus-based Grid services , 2007, Future Gener. Comput. Syst..

[78]  Holly M. Mortensen,et al.  Whole-mtDNA genome sequence analysis of ancient African lineages. , 2007, Molecular biology and evolution.

[79]  Broome,et al.  Literature cited , 1924, A Guide to the Carnivores of Central America.