Language, Culture and Genes in Bantu: a Multidisciplinary ...20Der%20Veen/... · OMLL (Eurocores,...

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Language, Culture and Genes in Bantu: a MultidisciplinaryApproach of the Bantu-speaking Populations of Africa

(OMLL – 01-JA27: 01-B07/01-S08/01-V01)

The Case of Gabon

Lolke J. Van der Veen (Project Leader), Jaume Bertranpetit (Principal Investigator),David Comas, Lluis Quintana-Murci, Mark Stoneking (Principal Investigator)

IntroductionThis interdisciplinary EUROCORES OMLL project started at the beginning of 2002 as anextended version of a similar project on Bantu languages and genes. This latter officiallycommenced, under Lolke Van der Veen’s supervision, in July 2000 as part of the “Origine del’Homme, du Langage et des Langues” programme launched and funded by the FrenchCNRS1. The present Collaborative Research Project (CRP) consists of one main projectsubmitted by the research laboratory “Dynamique du Langage” (UMR 5596, Lyon, France)and two joint projects submitted respectively by research teams from Germany and Spain (seebelow). This detailed outline of the project wants to highlight the project’s goals and specificfeatures, and summarises the state of the art.

The LCGB project: objectives and approachThe ultimate goal of the “LCGB” project is to elaborate a solidly based multidisciplinarytheory of the origin and expansion of Bantu and the Bantu-speaking populations, on the basisof correlations between linguistic, biological and anthropological markers. Do historicallinguistics, population genetics, cultural anthropology and archaeology all tell the same story?Combining linguistics and population genetics still is a rather controversial issue, but applyingthis enlarged multidisciplinary approach to the study of Bantu expansion may positivelycontribute to the debate and offers promising perspectives for a new synthesis in this field ofinvestigation.

In the first time, existing solid linguistically based phylogenetic classifications are to becompared with biologically based classifications, which are in the process of beingconstructed. Therefore, at the start of the project, very well-defined linguistically baseddiachronic inferences were submitted to geneticists having accepted to collaborate with ourteam of linguists, for close examination. The OMLL programme has allowed the developmentof collaborations with geneticists from France (i.e. a team directed by Dr. Lluis Quintana-Murci, Institut Pasteur, Paris, CNRS URA 1961), from Spain (a team directed by Prof. Dr.Jaume Bertranpetit and Dr. David Comas, Unitat de Biologia Evoltiva, Facultat de Ciènciesde la Salut i de la Vida, Universitat Pompeu Fabra, Barcelona) and from Germany (a teamdirected by Prof. Dr. Mark Stoneking, Max Planck Institute, Leipzig), but also from Gabon(Dr. Lucas Sica, Centre International de Recherches Médicales Franceville) and the USA (Dr.Sarah Tishkoff, University of Maryland). The French team —directed by Prof. Dr. Lolke Vander Veen—, as well as the Spanish and German teams have been granted funding by theirrespective national funding agencies. Without this innovative interdisciplinary research

1 The main lines of this OHLL project were presented at the 32nd Annual Conference onAfrican Languages held in Berkeley, March 2001. Van der Veen & Hombert (forthcoming).

OMLL (Eurocores, ESF) — Language, Culture and Genes in Bantu:a Multidisciplinary Approach of the Bantu-speaking populations of Africa

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programme, it would have been impossible to develop such a variety of extensivecollaborations and to create networking activities between thematically related projects.

The combination of these different sources of information we are working with, willmost certainly reveal both similarities and discrepancies between the disciplinary approaches.Both cases (presence or absence of correlations) will of course be equally interesting. Theresults of this study will subsequently be compared with the results obtained so far (or still tobe obtained) in other fields of research such as Cultural Anthropology, Archaeology andHistory. In a wider perspective, the outcome of the “LCGB” project will allow a comparisonwith the results of similar OMLL projects on Berber (project leader: Prof. J.-M. Dugoujon),Eurasian (project leader: Dr. A. Sajantila), Himalayan (project leader: Dr. P. de Knijff) andIsland Melanesia (project leader: Prof. S. C. Levinson), also presented in the present volume.

Main specific features of LCGBOver the last few decades, population genetic studies of the African continent have especiallytried to establish parallels between genetic and linguistic classifications at a fairly general andthus (historically) deep level. These studies have mainly focussed on the four major linguisticphyla of this continent (i.e. NIGER-CONGO, NILO-SAHARAN, AFRO-ASIATIC and KHOI-SAN2).Several of these studies have made strong claims about the African prehistory on the basis ofspecific genetic markers. Both traditional and molecular markers reveal correlations betweenthese major linguistic phyla and genetic data (cf. Cavalli-Sforza & al. 1994; Excoffier & al.1987 and 1991; Hammer 1994; Soodyall & al. 1993 and 1996; Poloni & al. 1997; Watson &al. 1996 and 1997; Melton & al. 1997; Stoneking & al. 1997, Scozzari & al. 1994 and 1999;Spedini 1999; Salas & al. 2002, etc.).

At the start of the “LCGB” project, Bantu had mainly been studied as a whole. NIGER-CONGO populations in general and Bantu populations in particular showed to have the lowestlevel of internal genetic diversity. This is of course compatible with the hypothesis of a“recent” (i.e. 5 000 BP) and rapid (although gradual) expansion of these populations(Excoffier & al. 1987, and 1991 (traditional markers); Poloni & al. 1997 (Y-chromosomehaplotypes), Scozzari & al. 1999 (biallelic and microsatellite Y-chromosome poly-morphisms), Spedini & al. 1999 (suggesting an expansion through minor migrationmovements). However, virtually nothing was known about the internal relationships betweenBantu-speaking populations. Criteria used for sampling individuals were most of the timeimprecise. All available studies suffered from a lack of accuracy in linguistic labelling(undifferentiated lumping, errors related to narrow linguistic classification, etc.) as well asfrom a lack of representativity (e.g. central African Bantu is still very badly underrepresentednowadays, cf. Salas & al. 20023). Carefully checked ethnolinguistic data were rarely takeninto account and geneticists usually clung on to linguistic classifications that were more orless outdated (e.g. Greenberg’s 1963 classification). There clearly was a desperate need for amore accurate and more rigorous approach, and for a close collaboration between geneticistsand linguists. The “LCGB” project aims at developing such an approach in particular for thestudy of phenomena with a lesser time depth, by working out rigorous criteria for samplingand analysis, by submitting up-to-date linguistic classifications and hypotheses4, and by

2 For a detailed up-to-date presentation of these phyla and the languages they comprise, seeWilliamson & Blench (2000).3 For this study, data from only three central African Bantu languages were available, andonly two Pygmy languages.4 Considerable progress has been made in the field of African linguistics over the last threedecades. These advances cannot be ignored.


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drawing benefit from a large-scale multidisciplinary approach including also archaeologists,cultural anthropologists and historians (see also below).

It should be underlined here that the Bantu languages (a group of about 600 languagesand language varieties, being part of the much larger NIGER-CONGO phylum, and coveringabout one third of the African continent), along with the SINO-TIBETAN languages, are amongthe best-studied languages of the world, just after the INDO-EUROPEAN languages. Importantaspects of the proto-language (i.e. “Proto-Bantu”, a language variety or dialect continuumspoken most probably about 5 000 BP in the borderland between present-day Nigeria andCameroon) have been reconstructed (Meeussen 1967 and 1969; Guthrie 1967-71) and solidhistorical inferences have been made concerning the origin and expansion of Bantu. Althoughsome more or less competing theories exist as far as the role of the forest area concerns in theexpansion process (gradual penetration into the forest —following waterways and/ormountain ridges— or northern “migration” bypassing the forest5 or both6), and alsoconcerning some secondary convergence and/or expansion zones, a general consensus existsamong scholars about the main lines of the expansion as well as several much more local anddetailed phenomena. (See map 1 below, for an up-to-date overview.) The Bantu expansionprobably coincided with the end of the Neolithic Age and was at least at some stage related tothe diffusion of agriculture and iron metallurgy (Phillipson 1985, De Maret 1982, Nsuka-Nkutsi 1989, Clist 1995, Oslisly 1996, Holden 2002, Diamond & Bellwood 2003).Environmental, demographic, social and economic factors must have played an important rolein this gradual and wave-like expansion.

For the present project, three geographically strategic zones were chosen in order to testthe linguistically based hypotheses: the Gabon area (i.e. the enlarged “Ogooué-Ivindo-Ngounié”7 area), the Kenya-Tanzania area and the Angola-Namibia area. These three areasare known to have played an important role in the Bantu expansion and have also undergone,to a certain extent, convergence phenomena (linguistic homoplasy). The present chaptermainly concerns the first area for reasons that will become explicit hereafter.

Fieldwork —a crucial aspect of this kind of research— is presently ongoing. Samplecollection (7 ml of blood per individual) started in 2002 in the Gabon area, for two reasons:firstly, extensive linguistic fieldwork has been conducted over the last 20 years by our (DDL)research team and consequently our knowledge of the languages of this area as well as of thearea itself and of its inhabitants is solidly founded at present; secondly, on the basis ofdetailed linguistic studies, we now know that the central part of Gabon is a place wheredifferent “migration” paths have met and led to convergence phenomena (linguistichomoplasy). Several languages show clear signs of admixture. Some of the ethnic groupsspeaking these “mixed” languages have been retained for sampling and testing, e.g. the Viyaand Makina (Shiwa) groups (see below). Studies in comparative linguistics based onphonological, morphological and lexical markers, have allowed us to elaborate a fairlyaccurate picture of groupings as well as the main expansion patterns. (See maps 2 & 3 below.)The project also benefits from already existing tight scientific collaborations with severalGabonese institutions such as the “Laboratoire Universitaire de la Tradition Orale” (LUTO)and the “Laboratoire d’Anthropologie” (LABAN) of the Omar Bongo University ofLibreville.

5 Cf. Coupez & al. (1975) and also Phillipson (1985).6 Scholars nowadays more and more agree about the role played by the rain forest in theexpansion process(es). Cf. Nurse & Philippson (2003:164-167); also Vansina (1990 and1995).7 The three major waterways of the Gabon area.


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Although French is the national language and exerts increasing pressure on the locallanguages, some 50 Bantu language varieties are still spoken in Gabon nowadays. Thesedialects can be grouped into a dozen clusters (A30 (BUBE-BENGA), A70 (YAUNDE-FANG),A80 (MAKAA -NTEM), B10 (MYENE), B20 (KELE), B30 (TSOGO), B40 (SHIRA), B50 (NJABI),B60 (MBAAMA ), B70 (TEKE) and H10 (KONGO)8) and belong to three different, thoughadjacent, geolinguistic zones (A, B and H). The Bantu languages of Gabon all belong to whathas been recently called “Forest Bantu”, which is a part of “Western Bantu” opposed to“Westcentral Bantu” and “Eastern Bantu” (cf. Nurse & Philippson 2003, Bastin & al. 1999).For Bastin & al. (1999) zone A languages together with B10-B20-B30 constitute what theycall “Northwestern Bantu” opposed to the rest of the western Bantu languages. This impliesthe presence of a relatively important linguistic “barrier” within Gabon (see map 2). Apartfrom the Bakao Pygmies who live in the surroundings of Minvoul and speak an Ubangian (i.e.non-Bantu) language, all other Pygmy groups have adopted one or several of the languagesspoken by the Bantu populations in whose vicinity they live. Their lifestyle has become semi-nomadic in recent days.

On the basis of shared phonological, morphological, morphosyntactical and lexicalinnovations, one can establish at least 4 higher-level Bantu clusters: B10+B30 (MYENE-TSOGO), B40+H10 (SHIRA-Vili (the latter being part of KONGO)), B50+B60+B70 (NJABI-MBAAMA -TEKE), and the dialect cluster A75 (Fang). For a detailed discussion of the statusand complex history of the MYENE-TSOGO group, see Mouguiama-Daouda & Van der Veen(forthcoming). This cluster is probably an ancient branch of Western Bantu, but it alsopresents signs of homoplasy. This homoplasy is probably due to a prolonged period of contactthat occurred some time after the initial separation between the two groups. The cluster (andB30 in particular) furthermore shows some (ancient) affinities with languages belonging tozone A. The B20 group clearly appears as a floating group in most recent linguisticclassifications (cf. Bastin & al. 1999). It may cluster with the MYENE-TSOGO group, but alsowith other groups.

One can presently assume with a fair degree of certainty that several of the linguisticgroups of Gabon must have arrived from more northern regions (the MYENE-TSOGO groupand much later in history, the Fang A75), others from regions located to the east (the NJABI-MBAAMA -TEKE group) and still others from regions located to the south (the SHIRA-Viligroup) (cf. Map 3 below, for major expansion patterns and relative chronology). The well-defined hypotheses submitted to the geneticists are founded on these data and havedetermined the choice of the populations to be retained for sampling.

In addition to the multidisciplinary character of our project and the well-established workinghypotheses it is based on, one other important, and rather unique feature of our project to bestressed here, is the rigorous criteria used for sampling. The choice of the populations to besampled is based on very precise linguistic criteria and is not made randomly. Only maleindividuals are sampled (in order to obtain information from both the mitochondrial DNA andthe Y-chromosome variation!), 35 years of age or older in order to reduce the influence of(recent) admixture, and at least 50 individuals per population; and this for at least 2populations of each ethnolinguistic cluster, if possible. Both parents of the sampledindividuals should belong to the same ethnic group. Furthermore, the declared (ethnic) originof the individuals is systematically verified by means of an ethnolinguistic and

8 Malcolm Guthrie’s reference system (Guthrie 1948). Guthrie divided the Bantu territory upinto 15 geolinguistic zones (A, B, C, etc.). Tens refer to language groups and units to specificlanguages.


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anthropological questionnaire conceived to collect details concerning place of birth,lineage(s), ethnic origin of both parents, grandparents, etc., and language(s) spoken over atleast four generations (see photography, last page of this paper). A team of Gaboneseanthropologists (LUTO / LABAN) will enlarge our research by an extensive study of thepossible impact of kinship systems, mating patterns, matrilineal or patrilineal descent,patrilocality, polygamy, the role of local slavery in history, as well as the role of socialstratification, cultural and political organisation, economy, trade and technology within thedifferent ethnic groups. Blood sampling is performed only with the individual’s consent,exclusively in collaboration with the local Health Services, and of course in agreement withthe research permits granted by the Gabonese government. Blood samples are never takenfrom blood banks. This would of course be much simpler from a practical point of view, butwould make the verification of linguistic and ethnic origin extremely difficult, if notimpossible.

Procedures for data collectionSample and ethnolinguistic data collection is conducted in the field, not in Europe, in closecollaboration with Gabonese researchers working in the fields of linguistics, populationgenetics and cultural anthropology. This work is performed by several teams, simultaneouslyin different places of the country, all using the same procedure, during several periods of theyear. This requires necessarily a very well planned coordination and the use of efficientcommunication systems. Each team is composed of linguists and cultural anthropologists.

On arrival in the villages, local authorities (both civil and moral) are informed. Withtheir consent an information campaign is organised in the presence of the elders of the village.Individuals are sampled in accordance with the rigorous criteria described in the previoussection. During blood sampling sessions — usually organised in health centres—, consentingindividuals fill out a form and several questionnaires, a small quantity of blood is taken (7ml). Each individual receives in return a set of essential pharmaceutical products as well assome refreshment. The blood samples are maintained at a temperature of +4° C and sent bycar then by plane within a delay of 48 hours, to the “Centre International de RecherchesMédicales Franceville” (CIRMF) in Franceville, in the southeast of Gabon, where DNAextraction is performed. Blood samples can absolutely not be exported from Gabon as such. Asmall quantity of blood is being used by the CIRMF (Dr. Lucas Sica) for the study ofdrepanocytosis, a blood disease affecting a considerable number of Gabonese individuals.

Purified DNA samples (of excellent quality) are sent on a regular basis by the CIRMFto the Institut Pasteur in Paris for preservation, sequencing, typing and analysis. The analysisof the mitochondrial DNA (mtDNA) and the Y-chromosome variation is being performed inEuropean laboratories (see hereafter).

Ethnolinguistic anthropological data are processed in Lyon and recorded in a largedatabase for further analysis.


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State of the artThe Gabon area

Extensive fieldwork in the Gabon area has allowed to collect over 950 blood samples(representing almost exclusively male individuals, most of them at least 35 years of age) in 14different places spread all over the country9. The last of a total of four missions10 in the fieldwas organised in February 2004. This implies that we now have access to genetic data ofexcellent quality from 21 out of the 50 Gabonese populations11, whereas the initial objectivewas to sample only 6 populations12!

Molecular genetics has shown over the last decades that our genome reflects, not only theprocesses intrinsic to its molecular nature, such as recombination and mutation, but alsodemographic processes that have modelled its composition, such as migrations, admixture,population expansions and founder effects. This is of course crucial for the kind of researchwe intend to perform.

The molecular part of the present project will focus on the analysis of two genomicregions, the mitochondrial DNA (mtDNA) and the Y-chromosome, that present particularproperties suitable to capture the footprints of demographic effects in our genome. MtDNAanalysis is in process at the “Institut Pasteur” in Paris under the supervision of Dr. LluisQuintana-Murci, and the Y-chromosome analysis (joint project), which started in Barcelona inNovember 2003, is directed by Prof. Jaume Bertranpetit (Principal Investigator) andconducted by Dr. David Comas (Executive Researcher). These uniparental-inherited geneticmarkers will be used to reconstruct the population history of several populations in CentralAfrica and, in a broader context, of the African continent. Given that the mtDNA and the Y-chromosome do not experience recombination, a clear phylogeny of both molecules has beenestablished. Moreover, the lineages within these phylogenies are not distributed randomlyacross human populations: the clear geographic structure of the uniparentally-inheritedlineages allows us to reconstruct the phylogeography of the individuals analysed.

Y-chromosome analysis (team directed by Jaume Bertranpetit)The Y-chromosome determines the masculinity in our species due to the presence of a singlegene, SRY (sex-determining gene). The zygotes with an X-chromosome and a Y-chromosomeyield a male embryo, whereas the zygotes with two X-chromosomes form a female one.Therefore, males inherit their Y-chromosome from their fathers who received them from theirpaternal grandfathers, and so on. The lack of recombination along the major part of the Y-chromosome, allows us to trace back the history of the chromosome.

9 These places are Booué, Cap Estérias, Fougamou, Franceville, Lambaréné, Libreville, LaLopé, Lastoursville, Malibé, Minvoul, Mouila, Ntoum, Port-Gentil and Sindara.10 Field missions were conducted in 2002 (one mission), 2003 (two missions) and 2004 (onemission).11 As indicates above, the target was a minimum of 50 male individuals per population. Formost of the 21 ethnic groups this target has been achieved.12 List of populations for which both linguistic and genetic data is available (with the numberof individuals sampled): Benga (53), Fang (70), Makina-Shiwa (50), Bekwil (5), Galwa (51),Rungu (42), Kele (50), Kota (59), Mbangwe (6), Shake (52), Tsogo (66), Viya (38), Kande(8), Shira (53), Punu (52), Nzebi (Njabi) (63), Duma (49), Obamba (Mbaama) (54), Ndumu(44), Teke (56), Bakao Pygmies (39). The latter speak an Ubangian language (i.e. non-Bantu).(Populations for which the target has been achieved completely, have been underlined.)


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To-date, a large number of genetic markers have been analysed to define the Y-chromosome diversity within humans. There are some polymorphic insertions in thechromosome, such as some Alu elements; and an important number of repetitions in tandem,such as the STRs (short tandem repeats). Nonetheless, the most promising markers in the lastyears are SNPs (single nucleotide polymorphisms) that consist of a base substitution, in whichone base is exchanged for another.

The present Y-chromosome study of the expansion of Bantu languages and populationswill be focused on a total 18 STRs and around 50 SNPs that will be typed in the same sampleset used for the mtDNA. The information yielded by these markers will allow us to classifythe samples into haplogroups within the Y-chromosome phylogeny. The mutation rates ofboth classes of markers are substantially different: STRs have a mutation rate per locus andgeneration around 10-3-10-4, whereas SNPs have a slower mutation rate of around 10-8. Thisdifference has profound implications for the Y-chromosome analysis. Due to their slowermutation rate, SNPs are unique events that define the major lineages of the Y-chromosome.On the other hand, the STRs are highly variable and allow us to define the diversity foundwithin the major lineages defined by the SNPs. Thus, the combination of both kinds ofmarkers will provide a fine resolution of the Y-chromosome diversity.

The analysis of the STRs is performed by the amplification of the regions where themarkers lie and a subsequent determination of the number of repeats that each STR iscomposed of by a capilar electrophoresis. The determination of the 18 STRs is performed inthree multiplex reactions, therefore, each multiplex resolves the genotypes of six STRs. Thegenotyping of the SNPs is performed by a single reaction, a SNPlex reaction, in which theallelic state of around 50 SNPs is determined.

After the genotyping of both kinds of markers, the Y-chromosomes for each individualwill be classified into haplogroups, groups of lineages that have a common origin and aredefined by their SNP composition. After haplogroup classification, the diversity of eachhaplogroup will be assessed by the results given by the STRs. Since the phylogeography ofthe Y-chromosome is available, we will be able to trace back the origin of the Y-lineagesfound and compare the results to those found for the mtDNA. This will allow us to reconstructdemographic events in our population set.

MtDNA analysis (team directed by Lluis Quintana-Murci)Towards the end of the 1980s, the publication of the first paper on the implications of thestudy of mtDNA variation for understanding the origin of modern humans (Cann & al. 1987)opened a new era for the study of human genetic variation and evolution. MtDNApolymorphisms being the first DNA markers utilised for evolutionary purposes, the lastdecade has been characterized by a large number of studies on mtDNA variation, which haveoffered new perspectives on the understanding of the origin of modern humans. Due to thestrict maternal inheritance of this molecule, the geographic distribution of neutral mtDNAmutations reflects the prehistory of women, in an analogous manner to the Y-chromosome.Since mtDNA mutations accumulate sequentially along radiating female lineages, individualmtDNA lineages diverged as women migrated into the different parts of the world.

To define mtDNA variation in our population set, we perform two parallel approaches:sequence of the HVS-I region and genotyping of a number of coding-region SNPs in order toproperly identify haplogroups. The two sets of data present differences in mutations rates;SNPs in the coding-region are more stable and rarely present recurrent mutations. Conversely,HVS-I sequence variation is often affected by recurrent mutation (homoplasy), since thisregion presents a higher mutation. Thus, coding-region SNPs define the main lineages of thephylogenetic tree and HVS-I sequence variation is very useful to assess internal variation of


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the haplogroups (defined by the coding-region mutations). The internal haplogroup variationcan be translated into expansion times of the main lineages.

All samples are sequenced for HVS-I D-loop from the nucleotide position (np) 16023 tonp 16368. As to the coding-region, all the samples are typed using Victor technology, basedon fluorescence polarisation, for the SNP T3594C, which separates haplogroups L1-L2 fromL3, for T10810C, defining haplogroup L1, and for the RFLP +16390HaeIII, defininghaplogroup L2. In addition, we have used a hierarchical approach to subdivide L2 and L3samples into their internal derivative lineages. All L2 samples have been typed for+13803/+13958 HaeII (defining L2a), +4157AluI (L2b), -13803/-13958HaeII (L2c) and–3693MboI (L2d). L3 samples have been typed for +10086TaqI (L3b), -8618MboI (L3d) and+2352 (L3e). The samples not belonging to these three sub-haplogroups internal to L3, havebeen assigned to L3f, L3g or L3* according to the HVS-I motif as in Salas & al. (2002).Samples assigned to L3 that were not belonging to any of the African specific branches of L3,were also tested for +10871MnlI (N) and +10397AluI (M).

Statistical analysesMethods of phylogeny reconstruction, such as median joining networks, will be employed toestablish and refine the topology of the phylogeny of both molecules, and time-depths of themain characterising lineages will be estimated. The apportionment of the genetic variationbetween and within populations will be estimated by means of the Analysis of the MolecularVariance (AMOVA). Moreover, a spatial analysis of the molecular variance (SAMOVA) willalso be performed by presetting different numbers of population groups. This approachdefines groups of populations that are geographically homogeneous and maximizes theproportion of total genetic variance due to differences between groups. In order to establishthe genetic relationships between the populations analysed, an analysis of principalcomponents based on haplogroup frequencies will be performed. Some other descriptivestatistical indexes, such as the Tajima’s D and Fu’s Fs neutrality tests, will be performed,when relevant, to establish the demographic past (constant versus stationary size) of thedifferent populations.

Sex-biased population history?As stated above, mtDNA and Y-chromosome are unique for their lack of recombination atmeiosis and are inherited through only one sex. Thus, the study of both these two moleculesin a given population will give a complete panorama of their maternal and paternal genetichistory. However, there is a number of examples where both sets of data give distinctinformation and this has been interpreted as being due to the different female/male socialhabits along history, such as polygamy, patrilocal exogamy, etc. The historical fact thatwomen, at marriage, relocate(d) in most cases to join the male partner makes themconsequently more genetically mobile than men. Women in their movements have contributedto masking the original differences between populations and, consequently, they have“homogenized” their gene pool. On the other hand, Y-chromosome variants turn out to bemore geographically clustered than mtDNA as a consequence of less male historical mobility.Since men turn out to be less mobile, their gene pool has contributed to keeping thesedifferences that are still recognizable through Y-chromosome studies. This suggested thatwomen’s movements have been very important during the past leading to the conclusion thatfemales have had a much higher migration rate than males (Seielstad & al. 1998).Nevertheless, although patrilocality must explain the differences in the patterns between the Yand mtDNA, it operates on a local scale and is not thought to be responsible for thecontinental and world pattern of diversity. Thus, some details are still not clear and willcertainly be object of further studies. In this context, a careful comparison in our population


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dataset of mtDNA and Y-chromosome variation will help to examine the relative contributionof males and females in shaping the patterns of diversity of central Africa and, moregenerally, it will shed some light onto the proportion and extent to which males and femaleswere involved into the Bantu expansions from (western) central Africa southwards andeastwards.

Preliminary resultsThe results of the study of mtDNA variation obtained thus far allow to start dealing seriouslywith issues such as the origin of the prevailing haplotypes and the role of these geneticlineages in the Bantu expansion, the confirmation or invalidation of the linguistically basedgroupings13 —i.e. a rather solid “northern” cluster (B10+B30 —MYENE+TSOGO— withprobably at least a part of B20 —KELE) and two “southern” clusters14 (B40+part of H10 andB50-B60-B70) which certainly are related to each other from a comparative point of view, butmore loosely than B10 and B30—, the impact of genetic admixture (convergence), the statusof several specific endangered minority groups (e.g. the Viya and the Makina/Shiwapopulations) in the process of being assimilated by surrounding populations, and also theorigin of the Fang population.

The Viya for instance, speak a language that basically belongs to the TSOGO (B30)group but its lexicon and phonology have undergone a fairly strong influence from a language(B41) spoken by the neighbouring Shira population. As mentioned before, the TSOGO groupin its turn clusters with the MYENE group (B10), on the basis of a variety of structural traits.Whether this clustering (also claimed by local oral tradition) is due to shared heritage or toconvergence by prolonged contact is still a matter of debate, but see Mouguiama-Daouda &Van der Veen (forthcoming) for new perspectives. The preliminary results of the analysis ofmtDNA and Y-chromosome variation seem to confirm Viya’s affiliation with TSOGO, as wellas the MYENE+TSOGO cluster.

The Makina people speak Shiwa (A83), but their language and cultural tradition isnowadays yielding to the fast growing influence of the Fang. The latter, whose language(A75) belongs to the A70 (i.e. YAUNDE-FANG) cluster, claim an Egyptian (or Sudanic) origin.This rather surprising origin is claimed by a considerable number of members of thiscommunity since the publication of Rev. Trilles’ theory on this issue at the beginning of the20th century). The first results of the mtDNA analysis support the position of those who, onthe basis of linguistic and more general methodological considerations, reject this theory.However, Y-chromosome data as well as a more detailed study based on cautiousinterpretation is needed to clarify in a more substantial way the migration/expansion history ofthis population15. Whatever the final outcome of this issue may be, from a linguistic point ofview, there is absolutely no doubt that Fang is a Bantu language. This dialect cluster clearlyattests the defining characteristics of the YAUNDE-FANG group.

Last but certainly not least, this ongoing study also sheds important new light on thehistory of slave trade and the origin of African-Americans.

An accurate account of the results of the mtDNA analysis of these individuals andpopulations performed by Dr. Lluis Quintana-Murci and his collaborators is forthcoming. Thepreliminary results of the Y-chromosome analysis based on the same samples were presented

13 Taking into account the most up-to-date linguistic studies (published or unpublished).14 The “northern” vs. “southern” distinction is more based on the presumed historical origin ofthe language groups than on their present geographic distribution.15 It is a historical fact that groups of Fang reached the Estuary of Gabon during the secondhalf of the 19th century AD (1874).


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and discussed by Dr. David Comas during an interdisciplinary workshop held in November2004 at the Institut Pasteur in Paris. Both types of analyses are ongoing.

The other regions

So far, no fieldwork has been undertaken in Tanzania for administrative reasons (difficulty ofobtaining research permits), funding problems (see below) and because of importantoverlapping between the objectives of the present project and fieldwork already carried out byother researchers from the US not directly engaged in the project. The original objective wasto obtain samples for genetic analysis from the Mbugu and neighbouring groups in Tanzania,in order to test hypotheses about the origins of the Mbugu (see hereafter, “Some problematicissues”. It would of course not be scientifically nor ethically sensible to resample the groupswe are interested in.

In order to compensate this absence of onsite research in this other important region, aspecial two-day « East African Workshop » was organised in May 2003, in which participatedseveral geneticists (i.a. Dr. Sarah Tishkoff, University of Maryland; Dr. Lluis Quintana-Murci, Institut Pasteur; Dr. Richard Cordaux, Max Planck Institute, Leipzig) and ninelinguists (most of them specialists of the languages spoken in this area, i.a. Gérard Philippson(DDL, Lyon), Derek Nurse (Memorial University of Newfoundland), Maarten Mous(University of Leiden) and Gerrit Dimmendaal (University of Cologne)). The aim of thisworkshop was to define new research areas for fieldwork on the basis of specific linguisticcriteria and, especially, to develop close scientific collaborations between linguists andgeneticists having already been working in this area. Extensive blood sampling has alreadybeen carried out by Dr. Sarah Tishkoff and her team, but much remains to be done, especiallythe interpretation of the results. Linguistic hypotheses will prove essential. Some of theseconcern mixed languages (cf. joint project submitted by Maarten Mous and Mark Stoneking,Max Planck Institute, Leipzig. See also hereafter, « Some problematic issues ».) But given thepresent situation, a new strategy will have to be chosen. Instead of investing energy infieldwork (and previously in obtaining research permits), new workshops will be organised assoon as Dr. Sarah Tishkoff’s database will become available on the Web (i.e. towards the endof 2004). Linguists, geneticists, archaeologists, historians and cultural anthropologists will beable to discuss and compare the results from their respective fields, and commence preparinga multidisciplinary publication.

Extending our project to the third area (Namibia-Angola) has not been possible either, forpolitical reasons and lack of local contacts, although specialists of the languages spoken thereare ready to collaborate. Recently, however, genetic data from Angola has become available(Plaza & al. forthcoming), which can be integrated into the present study. This extension willbe extremely important for a fine-grained examination of the western Bantu expansion.Plaza’s study —which is a contribution of the Spanish team together with other geneticists ofrenown— clearly demonstrates that genetic flow took place between southeastern (datamainly from Mozambique) and southwestern Bantu populations, and thus corroborates thelinguistic classifications (eastern linguistic traits in southwestern languages). (Cf. Soodyall &al. 1996.) The eastern and western Bantu expansion routes were clearly not independent. Thisstudy, which is based on the analysis of the mtDNA variation of 44 Bantu-speakingindividuals from Angola (analysis of the two hypervariable segments —HVSI and HVSII—of the mtDNA control region of the 9-bp deletion in the COII/tRNALys intergenic region), alsoshows that there are no traces of Khoisan lineages in the extant Bantu-speaking population


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from Angola, which suggest that the Bantu expansion provoke the migration southwards ofthe Khoisan-speaking people without substantial admixture.

Since the phylogeny of the Y-chromosome lineages has been well established and thereis some data from other African samples, it will be possible to determine migration processes,admixture with neighbouring populations and differential modes of dispersal between menand women by comparison to the results yielded by the mtDNA analysis performed in thesame individuals. All these analyses will contribute to the reconstruction of the populationhistory of Bantu-speaking populations in its Western expansion route.

Some problematic issuesUnfortunately, the joint project concerning the Ma’a/Mbugu language community of Tanzania(a mixed language presumably due to Bantu-speaking women mixing with Cushitic-speakingmen) has not been retained for funding by the Dutch funding agency, in spite of its quality.However, this highly interesting hypothesis has since been maintained within the main projectfor testing, and will consequently be financed as much as possible from other available funds.Maarten Mous —a specialist of Nilotic as well as of Eastern Bantu— will closely collaboratewith Dr. Sarah Tishkoff for the study of this specific issue.

For some time there was uncertainty about funding for the joint projects submitted bySpain and Germany. It now has been clearly established that as far as both participatingcountries are concerned funding has been granted. The Spanish project (i.e. the Y-chromosome analysis of the Gabonese data) started in November 2003. The German project,which initially concerned the Tanzania area, will perhaps be focussing on the Bantupopulations of Zambia. This possible extension is presently being examined.

Short overview of presentations, publications & other realisationsThis section provides a concise overview of the main presentations and publications producedthus far, as well as of the other activities developed since the beginning of the OHLL andOMLL programmes.

Reports, presentations and publications

As part of the usual evaluation procedure, written scientific reports have been submitted on anannual basis to the French CNRS, with regard to both the French OHLL programme and theEuropean OMLL programme (2001, 2002, 2003, 2004). Oral presentations with slide supportswere solicited in order to summarise the results obtained and to enable exchanges betweenrelated projects. All the reports produced to-date may be consulted freely.

Ensuing the first field mission, the anthropologists of the LUTO/LABAN (Omar BongoUniversity, Libreville) submitted a preliminary written report to the linguists and geneticists,based on the genealogical data obtained during this mission, with details about lineagescovering a time span of four generations, as a preparation of an extensive study of localmating patterns. Constructive comments were made in order to render further fieldwork stillmore efficient.

Two important (oral) presentations should be mentioned here. During the first “EastAfrican Workshop” (May, 2003), Sarah Tishkoff (University of Maryland, USA) made adetailed presentation of the state of the art in the population genetics of Tanzania. This mainpresentation was followed by a series of presentations on several related linguistic topics byDerek Nurse (Memorial University of Newfoundland, Canada), Maarten Mous (University ofLeiden, the Netherlands), Gerrit Dimmendaal (Cologne University, Germany) and GérardPhilippson (DDL, Lyon, France). These presentations aimed at defining new domains ofinvestigation for East Africa. The other important presentation was part of a popularisation


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effort: Christophe Coupé and Lolke Van der Veen presented a paper entitled “Quand leslangues rencontrent les gènes : une histoire des populations du Gabon” (June 25th, 2004) at themonthly seminar of the “Institut des Sciences de l’Homme” (ISH, Lyon, France).

Francesca Luca of the Institut Pasteur (Paris) presented a detailed poster at the 2nd DNAPolymorphisms in Human Population International Symposium held in Paris in December2003. Her poster, prepared in close collaboration with Lluis Quintana-Murci, was entitled“MtDNA variation in Central Africa: a microevolutionary study in Bantu-speakingpopulations from Gabon”.

In collaboration with other geneticists, the Barcelona team prepared a highly interestingpublication on mtDNA variation in Bantu-speaking populations of Angola offering newinsights into the western Bantu expansion (Plaza & al. forthcoming). Their contribution hasbeen submitted to Human Genetics.

Two major contributions were made by Lolke Van der Veen to the description andclassification of the Bantu languages of Gabon, i.e. a bilingual dictionary of the endangeredViya language (Van der Veen 2002) and a chapter on the B30 language group published inThe Bantu Languages. This important reference book was published by Gérard Philippsonand Derek Nurse in 2003 (London, Routledge). Both contributions were initiated before thestart of the OMLL programme, but significantly improve the understanding of the languagegroups of Gabon.

Finally Patrick Mouguiama-Daouda (DDL, CNRS) and Lolke Van der Veen wrote achapter entitled “B10-B30: conglomérat phylogénétique ou produit d’une hybridation ?” for aFestschrift dedicated to two outstanding specialists of Bantu, viz. Claire Grégoire and YvonneBastin (MRAC, Tervuren, Belgium). This chapter is an important contribution to theunderstanding of the rather complex relationships between these two linguistic groups andtheir history.

Interdisciplinary workshops

In addition to the above mentioned “First East African Workshop” held in May 2003, severalother interdisciplinary workshops have been organised. During a workshop held October 31st

2003, Lluis Quintana-Murci gave a detailed presentation of the first official results concerningthe analysis of the mtDNA, based on 308 DNA samples, collected from seven different ethnicgroups of Gabon. Another workshop, on the state of the art of anthropological research inGabon, was organised March 26th 2004. Specialists as Louis Perrois, Philippe Laburthe-Tolraand Raymond Mayer participated in this important meeting. Its objective was to initiate theidentification of cultural markers that may be significant for the classification of the Bantu(and non Bantu) populations of the area.

The next workshop, in which participated Richard Oslisly (WCS, Libreville, Gabon),Bernard Clist, Raymond Lanfranchi and Bernard Peyrot, was organised April 23rd 2004, onthe state of the art of archaeological research in Gabon. The objective of this meeting was tostart examining possible parallels between the linguistic inferences about Bantu history andthe findings of archaeology in the Gabon area.

Finally, linguists and geneticists met in November 2004, in Paris, in order to present theongoing genetic analyses and discuss the new results. The first results regarding the Y-chromosome variation were presented by David Comas. Some interesting parallels have beendetected with the results of the mtDNA analysis, but also a certain number of strikingdifferences. However, these observations are too preliminary to be discussed here.


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Databases

An ethnolinguistic database integrating all data collected during the four field missions isnearing completion. It comprises information about the sampled individuals: the individual’slineage, the lineages of his parents, the individual’s date and place of birth, language(s)spoken by the individual and by his parents, etc.

Meanwhile, the anthropologists have started to produce an up-to-date account of theavailable multifactorial ethnological data for the sampled Gabonese populations. A first draftof this paper, providing specific information for a dozen of ethnic groups, has been submittedto the French geneticists in March 2004, and a much more detailed version is being prepared.

Additional linguistic fieldwork

More linguistic fieldwork was carried out during the summer of 2004 by Dr.PatrickMouguiama-Daouda (DDL, associated researcher) and Pascale Paulin (DDL, graduatestudent) in order to complete our linguistic database. Patrick Mouguiama-Daouda checkedand collected lexical data from B20, B30 and B60 languages, and Pascale Paulin studied theBaka (pygmy) language and culture in the extreme north of Gabon (viz. the Minvoul area).

Ongoing research and perspectivesWe shall conclude this detailed outline with a short overview of the activities in progress or tobe developed in the near future.

Further research on languages and ongoing analyses in genetics

The DNA samples as well as the ethnolinguistic data collected during the last researchmission in the Gabon area (February 2004, see above) are currently being processed. The aimof this (temporary) final mission was to obtain additional blood samples from 400 (male)individuals in order to enlarge sample sizes, and also to allow for finer anthropological andgenetic analyses taking as significant level the lineages rather than the ethnic groups (theconcept of ethnic group being rather vague and therefore much more difficult to define). Thisobjective has been achieved but due to haemolysis some samples have been lost.

In spite of increasing difficulties in exporting DNA samples from Gabon, we veryrecently (i.e. September and November 2004) managed to transfer the remaining samples ofpurified DNA to the Institut Pasteur in France. Since all samples from Gabon are presently inParis (a total of some 900 samples), deep diversity and phylogeographic analyses of centralAfrican mtDNA can be undertaken and the results of these studies will be integrated in theglobal African context in order to determine genetic barriers. The relative dating of thebranches will also have to be examined. These analyses will continue during the second halfof 2004 in order to refine the results previously obtained. The most interesting preliminaryresults of this ongoing analysis were presented at the OMLL Conference in Leipzig April 4-6th

2004.Y-chromosome analysis is also ongoing. The remaining samples (see above) will be

analysed during the second half of 2004 and the beginning of 2005. A first account of theprogress made in this analysis was presented by the Spanish team at the OMLL Conference inLeipzig, April 4-6th 2004. Some results have been presented during a workshop organised inNovember 2004 (see above).

Other genetic markers may be explored subsequently. The study of autosomal markersis one of the options that are currently being examined.


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Interpretation of the results of the genetic analyses

Only a very careful, critical and fine-grained comparison of the results of the genetic analyseswith linguistic, anthropological, archaeological, and available historical data can serve as abasis for the interpretation of the recently acquired data. Most obviously, this stage of theproject, initiated in March 2004, is the most crucial (and delicate!) of all. What kind ofdiachronic information (i.e. historical scenarios) can legitimately be inferred from thesynchronic genetic distances? What has been the impact of mating patterns (endogamous orexogamous strategies), birth rate, group internal social, cultural and political stratification,habitat, residence strategies, geographical proximity, and prolonged contact (trade, exchangeof technologies, etc.) between ethnic groups? What is the historical time depth of the probablyhighly complex observed genetic phenomena?

Close collaboration with cultural anthropologists and specialists of other fields will beessential here. For instance, a detailed atlas of the lineages attested in Gabon (being preparedby Prof. Raymond Mayer (LABAN, Libreville) as well as a solidly based inventory ofcultural markers will be indispensable tools for the task of interpreting the data.

Application of phylogenetically based methods to linguistic data

The limits of lexicostatistics as a possible tool of historical linguistics have been recognisedby the vast majority of linguists. Therefore, phylogenetically based methods, and in particularthe cladistic method (maximum parsimony, maximum likelihood) will experimentally beapplied to the classification of the languages of the Gabon area, taking into account theavailable phonological, morphological and lexical material, in close collaboration with severalspecialists of this field, with Mahé Ben Hamed (DDL, Lyon, and INSERM, Paris) as mainconsultant. These much more sophisticated methods will possibly also be useful for the studyof the anthropological traits. Collaborations are being developed with John Nerbonne of theUniversity of Groningen (RUG, the Netherlands), and also with Russell Gray and ClaireHolden (University of Auckland, New Zealand) in order to apply with their help differentnewly developed methods to the available lexical data, e.g. the Neighbor-Net method (Bryant& Moulton 2004), and elaborate more realistic classifications of the languages that are beingstudied. The Neighbor-Net method for instance, allows to reveal conflicting signals in thecomparative data and thus render more adequately their complexity. The conflicting signals inthe data are often due to convergence phenomena. This method may also provide usefulinformation about possible dialect chains.

Publications in preparation

Several publications are currently being prepared for journals of population genetics, humanbiology and/or anthropology of international renown, as well as new publications on thedescription and the internal classification of the Gabonese languages aiming to provide a stillmore accurate picture of the linguistic situation of this area (cf. Mouguiama-Daouda & Vander Veen forthcoming; also Mouguiama-Daouda in press, Mouguiama-Daouda & Hombertforthcoming).

The editing of a multidisciplinary book presenting the main results of our investigationsis one of the major objectives of the LCGB project. This publication will include chapters onlinguistics, genetics, anthropology, archaeology and history, as well as a critical comparisonof the results from these different approaches. It will also summarise the procedures we havefollowed during fieldwork (useful for further research of this kind).

Furthermore, a thorough reflection on ways of communicating the obtained results tothe local populations, often eager to learn more about their group’s history, is imperative. Thisethical aspect of the project should not be taken underestimated. The results of the project


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may easily be misinterpreted or misappropriated for ideological reasons, and can in such caseshave dramatic consequences!

Extension of data collection/research to other regions

Once the results for the Gabon area will be available, research will have to be extended toother regions, geographically adjacent or not. This can be done in two ways, the first of theseoptions being the most plausible one: (1) the comparison of our results with those obtained byteams presently working in neighbouring countries; (2) new fieldwork.

Two crucial studies offer highly interesting perspectives for comparison. One concernsthe genetic data of Bantu-speakers of Angola that has recently become available in apublication prepared by the Barcelona team in collaboration with Antonio Salas (Plaza & al.forthcoming). This issue is crucial for the study of the western Bantu expansion (see alsobelow). The available data mainly concern the male lineages (Y-chromosome). The otherstudy focuses on data being collected and analysed by a multidisciplinary research teamdirected by Alain Froment (IRD (ORSTOM), ERMES, Orleans), Evelyne Heyer (CNRS,Musée de l’Homme, Paris) and Serge Bahuchet (CNRS, Musée national d’Histoire naturelle,France) as for the Cameroon area. Comparing their results with the ones our project will yieldwill allow us, inter alia, to improve the study of the genetic affiliation of the Fang population(related to the Ewondo and other population groups of Cameroon) as also of the Baka (andother) Pygmies of Gabon (the Baka are probably related to the Aka population of the CentralAfrican Republic and most certainly to the Baka Pygmies living in Cameroon). A doctoralthesis was recently started by Pascale Paulin (DDL, Lyon) on this Pygmy population and itslanguage in Gabon, under the supervision of Lolke Van der Veen.

The constantly increasing genetic database concerning central Africa will also make itpossible to envisage in the near future the study of health related phenomena, e.g. the analysisof genes involved in the immunity system in Bantu-speaking farmers vs. in Pygmies (nomadicor semi-nomadic lifestyle), in order to examine possible cultural bases of health risk.

The option of new fieldwork is a priori less plausible. Fieldwork implies time-consumingpreparation, needs contact persons in the field, demands ideally some existing agreement andcollaboration between scientific organisms facilitating the administrative formalities, and ifpossible some political stability.

The extension of sample collecting activities to the Congo area seems compromised forthe time being, partly for safety reasons, partly because of a lack of representatives (i.e.potential contact persons) in this highly interesting region.

New possibilities may however exist for Tanzania. Excellent linguistic data has recentlybecome available for the Makonde, a rather isolated population, formerly practisingmatrilocality, living in the south-eastern part of Tanzania. This ethnic group moved into thearea coming from more southern regions, whereas the other Bantu-speaking populationsarrived from the west. A clear linguistic barrier exists between Makonde and the otherlanguages. This linguistically based hypothesis is worth being tested, but obtaining researchpermits may be difficult. Official authorities are becoming increasingly reticent in regard toblood sampling activities by foreigners. (linguist), and a number of medical researchers of theAnatomy Department of the Muhimbili Hospital University have shown real interest. Ifresearch permits will be attributed, fieldwork will probably be carried out by Mark Stonekingand his team, in close collaboration with Sophie Manus who is a specialists of this languagecommunity.

Whatever the outcome of this new opportunity may be, linguists, geneticists,archaeologists, historians and cultural anthropologists will have to get together in the nearfuture in order to compare the results obtained in their respective fields for the Kenia-


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Tanzania area. A series of workshops will be organised on the theme of the “East Africanlanguages, cultures and genes” when Dr. Sarah Tishkoff’s data will have become available onthe Internet for discussion. A “Second (Extended) East African Workshop” will probably beorganised during the first half of 2005.

ConclusionFor the time being, the main focus of the “LCGB” project will be on the western Bantuexpansion. Three important reasons justify this choice. Firstly, this region is the least studiedof sub-Saharan Africa from the genetic point of view. Sample collection and analysis fill inthe gaps. Secondly, this region presents a particular historical interest. The western Bantuexpansion is probably one of the oldest movements as part of the spread of Bantu and has inthe southwest (Angola-Namibia) undergone admixture as a result of exchanges withpopulations having arrived from the east (?southeast, ?northeast). In many cases, goodlinguistic documentation is available. And finally, our investigations have started in thisregion for the reasons indicated above and have given so far the best results. It is important tobuild on this newly acquired basis, and we trust that this study will improve our understandingof the nature of this expansion as well as of its underlying mechanisms.

More generally, the array of different though related scientific activities assembled herefor one specific purpose will allow us to considerably increase our knowledge and improveour understanding of the population genetics of Central and East Africa and also to verify towhat extent a multidisciplinary project as « Language, Culture and Genes in Bantu » maycontribute to the ongoing study of African history and prehistory.

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Bryant, D. & Moulton, V. (2004). Neighbor-Net: an agglomerative method for theconstruction of phylogenetic networks. Molecular Biology and Evolution 21(2):255-265.

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implications of the last four millennia. In Sutton, J. (ed.) ‘The growth of farmingcommunities in Africa from the equator southwards’. Azania (Nairobi). vols. 29-30:324-331.

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Salas, A. & al. (2002). The Making of the African mtDNA Landscape. American Journal ofHuman Genetics. November 2002. 71(5):1082-1111.

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Soodyall, H. & al. (1996). MtDNA control-region sequence variation suggests multipleindependent origins of an « Asian-specific » 9-bp deletion in sub-Saharan Africans.American Society of Human Genetics 58:595-608.

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Spurdle, A. B. & al. (1992). Y chromosome probe p49a detects complex PvuII haplotypes andmany new TaqI haplotypes in southern African populations. Am. J. Hum. Genet., 50:107-125.

Spurdle, A. B. & al. (1996). The origins of the Lemba « Black Jews » of southern Africa:evidence from p12F2 and other Y-chromosome markers. Am. J. Hum. Genet., 59:1126-1133.

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Trilles, H. (1912). Quinze ans au pays des Fang. Paris, Desclée.Van der Veen, L. J. & Hombert, J.-M. (forthcoming). On the origin and diffusion of Bantu: a

multidisciplinary approach. Proceedings of the 32nd Annual Conference on AfricanLanguages (ACAL 32). [March 2001 Berkeley.]

Vansina, J. (1990). Paths in the Rainforests: Toward a History of Political Tradition inEquatorial Africa. Madison, University of Wisconsin Press.

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Major (recent) publications of the Principal Investigators

Jaume Bertranpetit & David Comas (and associated researchers)Plaza S., Salas, A., Calafell, F., Corte-Real, F., Bertranpetit, J., Carracedo, A., Comas, D.

(forthcoming). Insights into the western Bantu dispersal: mtDNA lineage analysis inAngola. [Publication submitted to Human Genetics.]

Fadhlaoui-Zid, K., Plaza, S., Calafell, F., Ben Amor, M., Comas, D., Bennamar Elgaaied, A.Mitochondrial DNA heterogeneity in Tunisian Berbers. Ann Hum Genet (2004).

Comas, D., Plaza, S., Wells, R. S., Yuldaseva, N., Lao, O., Calafell, F., Bertranpetit, J.Admixture, migrations, and dispersals in Central Asia: evidence from maternal DNAlineages. Eur J Hum Genet (2004).

Plaza, S., Calafell, F., Helal, A., Bouzerna, N., Lefranc, G., Bertranpetit, J., Comas, D.Joining the Pillars of Hercules: mtDNA sequences show multidirectional gene flow in theWestern Mediterranean. Ann Hum Genet (2003). 67(4): 312-328.

Osier, M. V., Pakstis, A. J., Soodyall, H., Comas, D., Goldman, D., Odunsi, A., Okonofua, F.,Parnas, J., Schulz, L. O., Bertranpetit, J., Bonne-Tamir, B., Lu, R. B., Kidd, J. R., Kidd, K.K. A global perspective on genetic variation at the ADH genes reveals unusual patterns oflinkage disequilibrium and diversity. Am J Hum Genet. (2002). 71(1):84-99.


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Mark Stoneking (and associated researchers)Kayser, M., Kittler, R., Erler, A., Hedman, M., Lee, A.C., Mohyuddin, A., Mehdi, S. Q.,

Rosser, Z., Stoneking, M., Jobling, M. A., Sajantila, A., Tyler-Smith, C. (forthcoming). Acomprehensive survey of human Y-chromosomal microsatellites. American Journal ofHuman Genetics. [accepted]

Cordaux, R. & Stoneking, M. (2003). South Asia, the Andamanese and the genetic evidencefor an “early” human dispersal out of Africa. American Journal of Human Genetics72:1586-1590.

Kayser, M., Brauer, S., Schädlich, H., Prinz, M., Batzer, M. A., Zimmerman, P. A., Boatin, B.A., Stoneking, M. (2003). Y chromosome STR haplotypes and the genetic structure of U.S.populations of African, European, and Hispanic ancestry. Genome Research 13:624-634.

Pakendorf, B., Morar, B., Tarskaia, L. A., Kayser, M., Soodyall, H., Rodewald, A. andStoneking, M. (2002). Y-chromosomal evidence for a severe reduction in male populationsize of Yakuts. Human Genetics 110:198-200.

Stoneking, M. & al. (1997), Alu insertion polymorphisms and human evolution: evidence fora larger population size in Africa. Genome Research, 7:1061-1071. Cold Spring HarborLaboratory Press.

Lluis Quintana-Murci (and associated researchers)Quintana-Murci, L., Chaix, C., Wells, R. S., Behar, D. M., Sayar, H., Scozzari, R., Rengo, C.,

Al-Zahery, N., Semino, O., Santachiara-Benerecetti, A. S., Alfredo, Coppa, Qasim Ayub,Aisha Mohyuddin, Tyler-Smith, C., Mehdi, S. Q., Torroni, A., McElreavey, K. (2004).Where West meets East: The complex mtDNA landscape of the Southwest and CentralAsian corridor. American Journal of Human Genetics 74 : 827-845.

Quintana-Murci, L., Bigham, A., Rouba, H., Barakat, A., McElreavey, K., Hammer, M.(2004). Y-chromosomal STR haplotypes in Berber and Arabic-speaking populations fromMorocco. Forensic Sci Int 140(1):113-5.

Zei, G., Lisa, A., Fiorani, O., Magri, C., Quintana-Murci, L., Semino, O., Santachiara-Benerecetti, A. S. (2003). From surnames to the history of Y-chromosomes: the Sardinianpopulation as a paradigm. Eur J Hum Genet 11: 802-7.

Quintana-Murci, L., Veitia, R., Fellous, M., Semino, O., Poloni, E. S. Genetic structure ofMediterranean populations revealed by Y-chromosome haplotype analysis. (2003). Am JPhys Anthropol 121(2):157-71.

Manni, F., Leonardi, P., Barakat, A., Rouba, H., Heyer, E., Klintschar, M., McElreavey, K.,Quintana-Murci, L. (2002). Y-chromosome analysis reveals a genetic regional continuityin north-eastern Africa. Human Biology 74: 645-658.

Lolke Van der Veen (and associated researchers)Mouguiama-Daouda, P. & Van der Veen, L. J. (forthcoming). B10-B30 : conglomérat

phylogénétique ou produit d’une hybridation. [Accepted. To be published in 2005 in aFestschrift for Claire Grégoire and Yvonne Bastin.]

Van der Veen, L. J. (2003). The B30 languages. In Nurse, D. & Philippson, G. (eds.). TheBantu Languages. London, Routledge. 371-391.

Van der Veen, L. J. & Bodinga-bwa-Bodinga, S. (2002). Gedandedi sa geviya, dictionnairegeviya-français. Collection Langues et littératures de l’Afrique Noire (Philippson, G. ed.).XII. Louvain/Paris, Peeters Publishers. 569 pp.

Luca, F., Van der Veen, L., André, E., Mouguiama-Daouda, P., Sica, L., Hombert, J.-M.,Quintana-Murci, L. (2003). MtDNA variation in Central Africa: a microevolutionary studyin Bantu-speaking populations from Gabon. Poster presented by Francesca Luca of the


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Institut Pasteur (Paris) at the 2nd DNA Polymorphisms in Human Population InternationalSymposium held in Paris, December 2003.

Mouguiama-Daouda, P. (in press). Contribution de la linguistique à l’histoire du Gabon : laméthode comparative et son application au domaine bantou. Paris, Editions CNRS.[Accepted.]


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Map 1. Bantu Expansion: Presumed expansion routes.Zones retained for hypotheses testing.

Presumed major expansion routes, convergence zones, andsecondary expansion zones. The Bantu homeland is situated in theNorthwest (borderland Nigeria-Cameroon). From this place, theBantu populations gradually moved across the rainforest to the East(one major movement) and to the South (other major movement).Some of eastern populations continued their southward spread.Convergence between eastern and western populations occurred atdifferent times and places (cf. the Ambo population, R22 (Guthrie’sclassification). The different shades (colours) in the arrows havebeen used to differentiate major regional “migration” movements.

Map created with Bantu Map Maker (T. Schadeberg, LeidenUniversity, The Netherlands.)

?

•Ambo R22

±5000 BP±2500 BP

±3500 BP?

±1500 BP

+++


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Map 2. The Gabon area (enlarged “Ogooué-Ivindo-Ngounié” region):Major linguistic groups

1. = the B10+B30+?B20 (MYENE-TSOGO-?KELE) cluster (in green)2. = the B40-H12b (SHIRA-Vili) cluster (in red)3. = the B50-B60-B70 (NJABI-MBAAMA-TEKE) cluster (in brown)4. = A75 (Fang) (in violet)5. = Baka language (in black)

(E.g. Kele = language name, B10 = language group.)

GABON

References acc. to Guthrie (1948).

B30

B30B20

B10

B10

B30

B20

B20

B20

B10GalwaKele

Tsogo

Viya

Kota1

B50 B70

Mbaama

Teke

B60

Njabi 3

A75

A75A75A75

Fang

4

B40

B40 B40Punu

Shira

2

ViliH12b

A75

A75

A75

B40 B20B40

B20

B60

B50

B20

Bakao Pygmies

B20

Rungu

5


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Map 3. The Gabon area (enlarged “Ogooué-Ivindo-Ngounié” region).Its presumed major (gradual) expansion routes, the central convergence zone.

[Linguistically based inferences.]

1. = Main north-south Western expansion route2. = B10+B30 (MYENE-TSOGO) branch splitting off3. = B40+part of H10 (SHIRA-Vili) branch4. = B50+B60+B70 (NJABI-MBAMBA (OBAMBA)-TEKE) branch5. = A75 (Fang)

[Most probable chronological order.]

(Language group references according to Guthrie 1948.)

GABON

Reference numbers according to Maho (2003).

Main Convergence Zone

B10-B30 (B20?)Myene-Tsogo-(Kele?)

2

B40-H12aShira-Vili

3

B50?-B60-B70Njabi?-Mbete-Teke

4

A75Fang

51


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Field mission July-August 2003.Thierry Nzamba-Nzamba, cultural anthropology student, filling out the ethnolinguistic

questionnaire in the presence of the main representative of the Viya community.(Photograph: Gisèle Teil-Dautrey (DDL).)

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