• mtDNA;
  • admixture;
  • Reunion Island;
  • phylogeography;
  • Indian Diaspora;
  • ethnic groups


  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information

Reunion Island is a French territory located in the western Indian Ocean. The genetic pattern of the Reunionese population has been shaped by contributions from highly contrasting regions of the world. Over the last 350 years, several migration waves and cultural and socio-economic factors have led to the emergence of six main ethnic groups in Reunion. India is one of the principal regions that contributed to the setting up of the Reunionese population. Diversity, demographic and admixture analyses were performed on mtDNA variation of the Reunionese of Indian ancestry, including the Malbar and Zarab ethnic groups, in order to question their history. Using a phylogeographical approach, we generated and analysed quantitative data on the contribution of the Indian parental populations. Furthermore, we showed that the settlement of Reunion Island by Indians did not involve a founder effect, except in the very beginning of the Reunionese settlement (at the end of the 17th century). The accuracy of our results was optimised by a re-evaluation of the classification of the Southern Asian mtDNA haplogroups. Finally, by comparing our results to a previous study dealing with the Reunionese population, we highlighted how ethno-historical data are critical for reconstructing the complex history of multiethnic populations.


  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information

Reunion Island is a French overseas ‘département’ located in the western Indian Ocean, off the eastern coast of Madagascar (Fig. 1). Reunion was first settled in 1663, when France decided to make this island a supply point on the Road to the Indies. The Reunionese population is currently over 780,000 inhabitants (Institut National de la Statistique et des Etudes Economiques; INSEE, and constitutes a multiethnic and multicultural society. Through a process of creolisation, the structure of the Reunionese population was affected by admixtures between groups originating from highly contrasting parts of the world: Sub-Saharan Africa, Madagascar, India, western Europe and south-eastern China (Barat et al., 1986; Scherer, 1994). Although creolisation in Reunion is of synthetic type (i.e., leading to the construction of one single identity; here, the Reunionese identity), the Reunionese population keeps a certain degree of ethnocultural segmentation (Médéa, 2002, 2003). As a result of the different immigration waves and the cultural and socio-economic interactions, six main ethnic groups emerged in Reunion Island: the Créoles Blancs (of European major ancestry), the Kaf (of African and Malagasy major ancestry), the Malbar (of Indian major ancestry), the Créoles Métis (of mixed ancestry), the Shinwa (of South-eastern Chinese ancestry) and the Zarab (of Gujarati ancestry). These ethnic groups are defined here according to the ‘nomenclature’ used by Reunionese themselves (Médéa, 2002, 2004).


Figure 1. Topographic map and geographic location of Reunion Island.

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India is one of the main regions that contributed to the setting up of the Reunionese population. The first wave of Indian immigration started at the end of the 17th century and continued during the 18th century. This migration involved mainly both Indian slaves (Filliot, 1974; Lacpatia, 1982) and Indian free workers who were hired as masons, carpenters, and weavers or for military or police purposes (Gerbeau, 1986a, 1991; Govindin, 1994). Nevertheless, the flows from India during the 17th and 18th centuries were not very important, notably because the preferred sources of Reunionese slaves were Sub-Saharan Africa and Madagascar (Filliot, 1974). At the dawn of the 19th century, only 3% (∼1,700 individuals) of the Reunionese servile population was of Indian ancestry (Lacpatia, 1982; Gerbeau, 1986b). Slave trade was prohibited in 1801, just as Reunion Island needed more and more workers to develop the sugar cane culture (Defos du Rau, 1960). The Reunionese planters had to search for an alternative source of workforce, and a new type of labour was established in the first half of the 19th century, the Engagisme. From 1828, but more importantly after 1848 (when slavery was abolished in the French colonies), Reunion attracted a huge number of workers who had freely—at least formally—signed a working contract: indentured workers. About 120,000 Indian indentured workers were successively recorded by the Reunionese Immigration Department (Lacpatia, 1982; Gerbeau, 1986a). Although discouraged by the Reunionese planters, the indentured workers were theoretically allowed to go back to India at the end of their 5 years long working contract. At the time that the influx of Indian indentured workers ended, in 1885, about 45,000 individuals of Indian origin or ancestry (one quarter of the whole Reunionese population at this time) stayed and settled down in Reunion Island (Dupon, 1974; Marimoutou-Oberlé, 1999). Finally, from the end of the 19th century, and until the middle of the 20th century, other free immigration took place from the state of Gujarat. These immigrants were of Muslim ancestry, and their religion prevented extensive marital exchanges with the other Reunionese ethnic groups (Scherer, 1994; Ismaël-Daoudjee, 2002).

In Reunion Island, the Indian Diaspora is represented by the Malbar (18.0% of the current Reunionese population; Médéa, 2004) and the Zarab (4.3% of the current Reunionese population; Médéa, 2004) ethnic groups. Whereas the Zarab population history is quite well-known (e.g.: Nemo, 1983; Delval, 1988; Ismaël-Daoudjee, 2002), records concerning the migration pattern of Indian slaves, indentured workers and their offspring are far less detailed. In fact, historical studies suggest five different regions for the Indian parental populations: (i) western India (including current Gujarat, Goa and Maharashtra), known for its contribution of slaves in the 17th and the 18th centuries (Filliot, 1974); (ii) south-western India (Karnataka and Kerala), (iii) south-eastern India (Andhra Pradesh and Tamil Nadu), and (iv) northern Indian (Bangladesh, West Bengal, Bihar, Jharkhand, Chhattisgarh, Madhya Pradesh and Uttar Pradesh) all of which provided both slaves in the 18th century (Barassin, 1957; Gerbeau, 1978) and indentured workers in the 19th century (Dupon, 1974; Lacpatia, 1982); and (v) Orissa from which indentured workers were shipped to Reunion Island in the 19th century (Dupon, 1974; Lacpatia, 1982; Fig. 2). Nevertheless, the quantitative contribution of each of these regions, involved in the Indian slave trade and the Engagisme, is scarcely known. Moreover, the settlement of Reunion Island remains to date poorly investigated through the means of DNA markers (but see Berniell-Lee et al., 2008).We intend to contribute to fill this gap by dissecting the mitochondrial genetic pool of the Reunionese of Indian descent.


Figure 2. Main Indian geographic areas (concerning the Reunionese immigration) and their haplogroup frequencies. Geographical abbreviations: EI, Eastern India; NI, Northern India; WI, Western India; SWI, South-western India; SEI, South-eastern India; ORI, Orissa. Clusters abbreviations: Other EA, HGs of assumed Eastern Asian origin (A, B, C, D, E, F, G, M7, M8, M12, R9, R22 and Z); Other WE, HGs of assumed western Eurasian origin (HV2, I, J, K, N1a, N1b, R0a, T, U1, U2e, U3, U5, W and X); Other M/N/R IS, other HGs M, N and R of assumed Indian origin (M20a, M20b, M26, M30b, M30c1, M30d1, M30e, M33a1, M34a, M34b, M36a, M37b, M39b, M40a, M41, N5/M30, N5a, N5b and R2); R30/R31, includes R30a1, R30b1, R30c, R30d, R31a and R31b; U2 IS, U2 clusters of assumed Indian origin (U2*, U2a, U2b, U2c).

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Using a phylogeographical approach, mtDNA proved to be very informative in deciphering the contribution of different African regions to the gene pool of current American populations stemming from the Atlantic slave trade (Salas et al., 2004, 2005). Such studies were based on the highly resolved phylogeny of the Sub-Saharan mitochondrial haplogroups (HGs) (e.g. Salas et al., 2002; Kivisild et al., 2004; Behar et al., 2008). Concerning Southern Asia, the classification of mitochondrial HGs is not as successfully completed and is still in progress. In order to dissect the Indian Diaspora's history in Reunion Island, we propose here, as an essential preamble, a re-evaluation of the Southern Asian mitochondrial HGs. Our phylogeographic results have allowed us to decipher the admixture processes involved during the setting up of the Malbar group, and more generally the admixture processes involving Indian immigrants. Two different types of admixture can be distinguished: (i) inter-Indian admixture, which involved Indians from different geographical regions, languages, castes and religions (Lacpatia, 1987; Marimoutou-Oberlé, 1999); and (ii) extra-Indian admixture, which involved Indians and individuals from the other Reunionese population components (mainly originating from south-eastern Africa and Madagascar: Dupon, 1974; Fuma, 2002). Concerning inter-Indian admixtures, our study has shown that the areas delimited by historical studies as involved in Indian migrations for Reunion Island must be revised.

Additionally, the colonisation of an island often involves few individuals and leads to a founder effect. This founder effect usually induces a decrease in genetic diversity and/or important and random fluctuations in allele frequencies (Holgate, 1966; Nei et al., 1975; Chakraborty & Nei, 1977). Such a founder effect was recently hypothesised concerning the mitochondrial gene pool of the Indian indentured workers who arrived in Reunion in the 19th century (Berniell-Lee et al., 2008). We therefore explored the Reunionese mtDNA pool of Indian ancestry in order to test the impact of a possible founder effect on their genetic variation.

Materials and Methods

  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information

Sampling Protocol and Criteria

This study was approved by the French Commission Nationale de l’Informatique et des Libertés. More than 650 Reunionese contributed their DNA to this study. All donors were volunteers and signed an informed consent. Additionally, all volunteers were interviewed and filled in a questionnaire in which they provided ethnic and genealogical information about their ascendants. Jugal epithelial cell samples were collected and DNA was extracted as described elsewhere (Dubut et al., 2009). We subsequently investigated each genealogy to ensure the unrelatedness of volunteers for at least three generations. The ethnic affiliation of the earliest known uterine (strictly maternal) ancestor (most often the maternal grand-mother) was chosen for the mitochondrial ethnic affiliation.

Among the sampled mitochondrial chromosomes, 143 belonged to the Malbar (sample MB) and 42 to the Zarab (sample ZB). After being typed for mitochondrial variation and assigned to HGs (see below), the mitochondrial chromosomes from the other Reunionese ethnic groups were examined in order to extract haplotypes (HTs) of assumed southern Asian ancestry, labelled ‘Other’ or ‘Other #’ (n = 76) in Table S5. Among the non-Malbar and non-Zarab HTs, three HTs exhibited a relatively high frequency (labelled ‘Other #’ in Table S5). We conducted further genealogical investigations demonstrating that their frequency was conditioned by a previously well-characterised founder event (e.g. Barassin, 1960) involving Indian women who arrived in Reunion Island during the 17th century, whose name and surname are known. ‘Other #’ HTs were discarded from admixture and diversity analyses since it has been shown that they disrupt the estimation of several parameters. In particular, the frequencies of these HTs enhance the frequency of some HGs and disrupt the accuracy of the admixture coefficients (Dubut, 2008). By keeping only ‘Other’ HTs and Malbar HTs of assumed southern Asian ancestry, a third sample, called RIN, was constituted. RIN (n = 151; see Table S1) includes: (i) HTs that revealed belonging to HGs of southern Asian ancestry; and (ii) HTs belonging to HGs shared by western Eurasians and southern Asians (specifically: I, W and X). HTs from the Zarab group were excluded from RIN because the geographical origin of this group is known (Gujarat) and because Islam, a strong identity factor, prevented maternal gene flows from the Zarab to the other Reunionese ethnic groups (Scherer, 1994). The sample RIN is expected to allow the estimation of the contributions of the different Indian regions, since it takes into account both the Indian genetic variation that is predominantly present in the extant Malbar group and the genetic variation that passed into the non-Malbar groups by admixture over the Reunionese history.

Population Comparisons and HVS-1 Database

Following results from historians (Defos du Rau, 1960; Dupon, 1974; Filliot, 1974; Lacpatia, 1982), we compiled data from source populations in order to investigate the admixture processes that may have shaped the mitochondrial gene pool of the Reunionese of Indian ancestry (represented by samples MB, ZB and RIN). These data include HVS-1 sequences available in the literature from populations in the Indian sub-continent (including India, Sri Lanka and Bangladesh; n = 3883), South-eastern Africa (n = 416), Madagascar (n = 37), South-eastern China (n = 213) and France (n = 437). Information about these populations is reported in Appendix I. Concerning the Indian populations, the data were first clustered by Indian states, and then by broader geographical regions delineated according to the historical sources dealing with the Indian immigration to Reunion Island (Fig. 2). Eastern India was not recorded as a source population by historians. We included this region to test the robustness of Salas et al.'s (2004) method within the Southern Asia framework.

mtDNA Analyses and Haplogroup Determination

Hypervariable segments 1 and 2 (HVS-1 and HVS-2) of the control region, along with part of the neighbouring coding region, were simultaneously amplified by PCR using primers L15832 (Dubut et al., 2004) and H408 (Vigilant et al., 1991). The segment of the coding region between primers L10170 and H10660 also was amplified as described in Yao et al. (2002). Before sequencing, the two PCR products were co-purified using the QIAquick PCR Purification Kit (QIAGEN, Hilden, Germany). The control region was sequenced using primers L15832, L16200 (light chain, nps 16194-16217, relative to the revised Cambridge reference sequence; Andrews et al., 1999) and HV2S (Macaulay et al., 1999). For sequences containing a homopolymeric cytosine stretch from positions 16182, 16183 or 16184 to 16193 (usually associated with length heteroplasmy), additional sequencing was carried out using primer H16350 (heavy chain, nps 16327-16351). The segment 10148-10659 was sequenced using primer L10170. Our protocol permitted the unambiguous achievement of the control region from position 16024 to 00393, and of the coding regions from position 15882 to 16023 and from 10192 to 10659. These coding regions are known to contain several phylogenetically relevant sites that are helpful in assigning a HT to a HG (e.g. Yao et al., 2002). Sequencing was carried out on an ABI PRISM® 3100 Genetic Analyser using the BigDye® Terminator v1.1 Cycle Sequencing Kit (Applied Biosystems, Foster City, CA, USA). Since traces were of excellent quality and unambiguous, the obtention of only one strand was sufficient. Nevertheless, doubtful segments, segments presenting apparent incongruent association of polymorphisms, indels, and heteroplasmies were re-amplified and re-sequenced, as recommended (Bandelt et al., 2001; Yao et al., 2002). Moreover, several coding region nucleotide sites were typed in order to allocate more precisely each HT to a HG. Nucleotide sites 3594, 10873, and 12705 were systematically typed using allele-specific PCR (ASPCR) optimised as previously described (Dubut et al., 2004; primers and PCR protocols in Table S1). The number of repeats of the 9bp motif from the COII/tRNALys intergenic region was also systematically typed as described in Yao et al. (2002). Further molecular analyses were conducted in order to determine more precisely the mtDNA HG of each sample (Table S3). Additionally, some nucleotide sites were typed in order to test the monophyly of clusters previously characterised by the molecular analyses described above (Table 1). These sites were chosen according to the coding regions’ polymorphisms encountered on published complete mitochondrial genome sequences. Furthermore, each HVS-1 sequence from Eurasian populations used for comparison was classified into a mitochondrial haplogroup following the determination key reported in Appendix II. Except in case of other specification (see Appendix II), the classification was done according to (i) Metspalu et al. (2004), Palanichamy et al. (2004), Quintana-Murci et al. (2004) and Thangaraj et al. (2006) for sequences from the Indian sub-continent, (ii) Richards et al. (2000) for western Eurasian sequences, and (iii) Kong et al. (2006) for eastern Asian sequences. Additionally, the proposal of classification integrates a re-evaluation of the human mitochondrial phylogeny based on HVS-1 motifs and the additional genotyping of the coding regions described above. The HVS-1 sequences of Sub-Saharan Africa were classified into haplogroups according to Salas et al. (2004) and Kivisild et al. (2004).

Table 1.  Coding regions sites genotyped to investigate monophyly of some specific HVS-1, HVS-2 (nps 16023-00300) and coding regions (nps 15882-16023 plus 10148-10659) motifs
HG of the investigated HTsHVS-1motif (minus 16000)HVS-2 motifPolymorphisms of the coding regionsGenBank ID of the involved complete mtDNA sequences (Ref ID; Reference)Investigated nucleotide sites
  1. Notes: All the other coding region sites which were investigated to assign HTs to HG are listed in Table S3. The results from the coding region typing are reported in Table S5.

(R53; Palanichamy et al., 2004)8251
(T26; Palanichamy et al., 2004) 
(R43; Palanichamy et al., 2004) 
(S11; Palanichamy et al., 2004) 
(R188, R58; Sun et al., 2006)10750
M*223-290-9bp delAY9223008623
15924(C4; Sun et al., 2006)8674
(R81; Sun et al., 2006)3652
(R44; Sun et al., 2006) 
(R61; Sun et al., 2006)15287
(C48; Sun et al., 2006) 
(T13; Sun et al., 2006)10906
(A24, T72; Sun et al., 2006) 
(T69, R110; Sun et al., 2006) 
(T71; Sun et al., 2006)2442

Data Analysis

When reported, HG frequencies were associated with a bayesian 0.95 Credible Region (0.95CR; Richards et al., 2000) calculated using the program SAMPLING kindly provided by Vincent Macaulay. Network 4.5 (Fluxus Technology Ltd, Suffolk, UK, was used to compute median-joining networks (Bandelt et al., 1999) for newly characterised HGs based on HVS-1 polymorphisms. The network construction involved both sequences present in our database (Table S4) and additional data from Thangaraj et al. (2005a), Barnabas et al. (2006), and Kumar et al. (2006) (these additional data were not used for the other statistical analyses). For each cluster, the Time of the Most Recent Common Ancestor (TMRCA) was estimated using the ρ statistics methodology as described in Forster et al. (1996) and Saillard et al. (2000a) using Network 4.5. Admixture coefficients were estimated by using the method described by Salas et al. (2004) with a program provided by Vincent Macaulay. Several combinations of parental populations were tested. The standard error generated by this method depends on the phylogenetic resolution (Salas et al., 2005). A further characterisation of the Indian mitochondrial gene pool was thus an essential prerequisite.

Then, the genetic diversity of Malbar, Zarab and parental populations was investigated. We notably expected to be able to detect the impact of a possible founder effect or genetic drift on the Reunionese mitochondrial gene pool. Mitochondrial DNA sequences from parental populations that contained unresolved nucleotides were discarded from the following analyses. Moreover, mtDNA heteroplasmies were not considered in the statistical analyses. According to (i) the maximum parsimony principle, (ii) the known human mitochondrial genome phylogeny and (iii) the phylogenetic information available in our data set, we set the heteroplasmic nucleotide sites to one-step-before ancestral state. In order to compare a large number of populations, we analysed HVS-1 between sites 16090–16365, excluding nps 16182 and 16183 (for the same approach, see Trejaut et al., 2005).

In order to detect a possible founder effect that may have affected the mtDNA pool of the Reunionese population of Indian ancestry, we conducted diversity and demographic analyses. The following diversity estimators were calculated using Arlequin 3.0 (Excoffier et al., 2005): the number of distinct haplotypes (k); the number of polymorphic sites (S); the gene diversity (H; Nei, 1987); the nucleotide diversity (η; Nei, 1987); and the mean number of pairwise differences (θπ; Tajima, 1983). We used R 2.1.0© (R Development Core Team, 2005) to bootstrap individuals within samples and to estimate the mean and 95% Confidence Interval (95%CI) of θk. θk was defined by Ewens (1972) as the solution to inline image, where E(k) is replaced by k, and computed using the Maximum Likelihood equation. The demography of the Reunionese population was explored by estimating its impact on the genetic pattern using neutrality tests and tests of the pairwise distribution. In fact, these tests are commonly interpreted as tests of demographic equilibrium for neutral genetic systems (e.g. Rogers & Harpending, 1992; Tajima, 1993), and revealed useful in case of historical (recent) settlements (e.g. Helgason et al., 2003, for Iceland; Chaix et al., 2004, for the Vlax Roma). Moreover, it has been shown in the population of Quebec that, when the source population is at mutation-drift disequilibrium and experiences demographic expansion, a founder effect in the newly founded population creates an allele frequency distribution identical to that expected for a population at mutation-drift equilibrium (Heyer et al., 2001). Comparing the results of neutrality tests and tests of the pairwise distribution between the source populations and their daughter populations, therefore, allows for the detection of a possible founder effect in the daughter population. We investigated the pairwise distribution (Slatkin & Hudson, 1991; Rogers & Harpending, 1992) within samples by performing two validity tests of the stepwise expansion model based on the pairwise distributions using the bootstrap approach implemented in Arlequin 3.0 (Excoffier et al., 2005): the raggedness index (RI; Harpending, 1994) and the SSD statistic (Schneider & Excoffier, 1999). Furthermore, we performed two neutrality tests: Tajima's (1989) D statistic (DT; using DnaSP 4.0, Rozas et al., 2003), and Fu's (1997) neutrality test (Fs; using Arlequin 3.0).


  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information

Phylogeographical preamble

Using (i) more than 3,800 Southern Asian HVS-1 sequences, (ii) information stemming from published complete mitochondrial genomes, and by (iii) conducting a specific and oriented genotyping of Reunionese mtDNA haplotypes (Tables S4, S5), we were able to characterise and/or redefine 42 mtDNA HGs or sub-HGs (Appendix II, Fig. S1, Table S6). Forty-one of them are associated with a diagnostic HVS-1 motif (Appendix II). These clusters include sub-clusters of HG M (M4a1, M4c, M5a1, M5a2a, M5a2b, M5b, M5b1, M6a1, M6a2, M19, M19a, M20a, M20b, M20b1, M26, M30c1, M30d1, M30e, M33a1 and M39b), HG N (N5a and N5b) and HG R (HV10, HV11, R5a2b, R5a2b1, R5a2c, R6a1a, R6a2, R6b, R30a1, R30b1, R30b2, R30c, R30d, R31b, T2b1, U2b1, U2b2, U2c1 and U2c2). In order to further characterise these newly defined clusters, we reported their TMRCA based on transitions on segment 16090-16365 from the D-Loop (Table S6). As for M44, its characterisation is based on coding region polymorphisms alone (transitions at nps 8623 and 15924, and one 9bp repetition within the COII/tRNALys intergenic region), according to complete sequence C4 from Sun et al. (2006) and three distinct Malbar HTs (JC106, YO020 and VA003). HGs HV10, HV11, M19, M20, M26 and M44 are characterised for the first time in this study.

The Reunionese HGs

The mitochondrial variation observed in MB, ZB and RIN samples are summarised Table S5. Three southern Asian clusters (M5a*, M6a1 and U2a) are especially frequent within our non-Malbar and non-Zarab sample (14.7%, 0.95CR = 8.4-24.4%; 10.7%, 0.95CR = 5.6-19.7%; and 37.3%, 0.95CR = 27.2-48.7% respectively), whereas they are far less frequent among the Malbar and Zarab sample (for M5a*: MB, 2.8%, 0.95CR = 1.1-7.0%; ZB, 2.4%, 0.95CR = 0.6-12.3%; for M6a1: MB, 2.8%, 0.95CR = 1.1-7.0%; ZB, 0.0%, 0.95CR = 0.0-6.7%; and for U2a: MB, 0.0%, 0.95CR = 0.0-2.1%; ZB, 0.0%, 0.95CR = 0.0-6.7%; Table S5). Moreover, the variation of each of these clusters within the non-Malbar and non-Zarab sample is almost resumed by only one HT. Additional genealogical investigations dealing with the uterine line of descent of the individuals presenting these HTs has indicated that the high frequency of these clusters was conditioned by a founder event that occurred at the end of the 17th century (data not shown). This founder event involved Indian women indentified by name and surname, and was previously well-characterised by historical studies (Barassin, 1960) and genealogical compilations (Ricquebourg, 2001).

Several HGs and HVS-1 motifs found among the non-Malbar and Malbar Reunionese reveal the impact of South-eastern India (see Tables S4, S5 and references therein): (i) the basal HVS-1 motif of HG HV10 (304-311) is particularly common in Andhra Pradesh; (ii) the HVS-1 motif 093-184-223-311-362 of M6b appears Tamil Nadu-specific; (iii) the basal HVS-1 motif of M36a (193-223) is especially encountered in Andhra Pradesh and Tamil Nadu (but also in Sri Lanka); (iv) N5a is rare in India but found in Andhra Pradesh (and Karnataka); (v) the T2b1 HVS-1 motif 126-275-294-296-325 was only detected in Andhra Pradesh and Tamil Nadu; and finally, (vi) two Reunionese HTs (GY101 and ZQ083) classified as R* exhibit a HVS-1 motif (189-319-362) were only found in Tamil Nadu. On the contrary, M19, quite frequent in Orissa, is absent from the Reunionese mitochondrial gene pool. Concerning the Zarab, the mitochondrial variation is exclusively comprised of clusters of Southern Asian origin (HV11, M2, M3, M4’30, M5, M20, M38, M39, N5, R5, R8, R30, U2b, U2c and U7; Table S5). This observation provides further evidence that Indian Muslim communities mainly stem from the conversion of Hindu people and not from Muslim population flows from the Middle East and/or Central Asia between 711 A.D. and the start of the 14th century (Agrawal et al., 2005; Terreros et al., 2007).

Admixture Analyses

The estimated admixture coefficients computed from RIN, MB and ZB are reported in Table 2. For admixture analyses, the RIN sample was used to estimate the contributions of the different Indian regions, since RIN encompasses all the sampled Indian mitochondrial variation in Reunion: the HTs from India that are still present in the Malbar, and the HTs from India that passed into other ethnic groups over the years. Several combinations of Indian source populations were investigated, in order to estimate the contributions of southern India and of its sub-regions. Indeed, historical studies indicated that southern India is the main contributing region for Reunion, and that south-eastern India contributed more than south-western India. Moreover, within south-eastern India, Tamil Nadu is usually considered as the principal origin of Indian emigrants compared to Andhra Pradesh (Dupon, 1974; Lacpatia, 1982; Singhvi et al., 2001). The MB sample permitted the quantification of admixtures between Malbar and the other Reunionese components (South-eastern Africa, Madagascar, France and South-eastern China) before the collapse of social barriers in the second part of the 20th century. For the admixture analyses, the ZB sample can be considered as a control since our sampling criteria ensured that we discarded all the Zarab mtDNA of non-Gujarati ancestry from ZB. Analysis of the Zarab sample therefore allowed investigation of the performance of the admixture method in the case of a non-admixed sample.

Table 2. Admixture coefficients for the Reunionese population samples.
Sample ID1nParental populations combinationAdmixture coefficients ± SD for parental populations2
RIN151A0.702 ± 0.072----0.129 ± 0.0630.036 ± 0.0310.123 ± 0.0600.010 ± 0.010
B-0.634 ± 0.088--0.084 ± 0.0570.116 ± 0.0610.037 ± 0.0310.114 ± 0.0580.010 ± 0.010
C--0.520 ± 0.0930.127 ± 0.0540.080 ± 0.0550.114 ± 0.0610.037 ± 0.0310.111 ± 0.0570.010 ± 0.010
Sample ID1nAdmixture coefficients ± SD for parental populations3
  1. Notes: 1MB, Malbar; ZB, Zarab; RIN, Reunionese with haplotypes of Indian ancestry; 2SI, Southern India (SEI+SWI); SEI, South-eastern India (ANP+TN); TN, Tamil Nadu; ANP, Andhra Pradesh; SWI, South-western India; WI, Western India; ORI, Orissa; NI, Northern India; EI, Eastern India; 3SEA, South-eastern Africa; MD, Madagascar; IND, India (SI+WI+NI+ORI); FR, France; SEC, South-eastern China.

MB1430.076 ± 0.0220.070 ± 0.0210.810 ± 0.0360.029 ± 0.0210.015 ± 0.011
ZB420.021 ± 0.0210.021 ± 0.0210.872 ± 0.0590.061 ± 0.0480.024 ± 0.024

Southern India, and especially its eastern part, is known by historians to have been the major source of indentured workers during the 19th century. The high admixture coefficients associated with Southern India (0.721 ± 0.066; source population combination A, Table 2) and with South-eastern India (0.630 ± 0.081; combination B) confirm the pre-eminence of these regions as sources of the Reunionese Indian Engagisme. Nevertheless, the contribution of South-western India turned out to be marginal (∼9%), while the contribution of South-eastern India encompassed nearly two thirds of the entire Indian contribution. Interestingly, within South-eastern India, it was revealed that Andhra Pradesh contributed far more immigrants than Tamil Nadu (0.462 ± 0.088 vs. 0.175 ± 0.066; combination C). Northern India exhibits admixture coefficients which indicate that it was involved in the settlement of Reunion in a minor but non-negligible way (∼15%). On the contrary, admixture analysis revealed a very limited contribution by Orissa and Western India (∼5-6%). As expected, Eastern India exhibited a negligible admixture coefficient (∼1%). Moreover, the different combinations of the Southern source populations affected the values and standard error of the admixture coefficients associated with the other Indian source populations only marginally. This suggests that Salas et al.'s (2004) method fits the Indian mtDNA context, though the phylogeographical resolution of Indian populations is not so complete.

About 90% of the Zarab variation is attributed to their unique parental population (India). Moreover, in respect of their values and standard errors, the coefficients associated with the other parental population appear to be negligible. Most of the Malbar mitochondrial variation (>80%) is also of Indian origin. Nevertheless, our results showed that they significantly admixed with Malagasy and African components of the Reunionese population. Indeed, these two components represent about 15% of the entire Malbar mitochondrial gene pool. However, Malbar did not admix with Reunionese components of French or South-eastern Chinese ancestry, admixture coefficients being within 2 SDs of 0.

Diversity and Demographic Analyses

The results from the analyses carried out in order to evaluate the genetic diversity and demography of the parental populations and of the Reunionese mitochondrial gene pool of Indian ancestry are summarised in Table 3. Both Indian population samples (SEI and GUJ), as well as French (FR) and South-eastern Chinese (SEC) samples, exhibit relatively high diversity and show signs of population expansion in respect of the neutrality tests and the unimodality tests (RI and SSD). On the contrary, populations that are known to have experienced one or more founder effects [South-eastern African (SEA) and Madagascar (MD); Pereira et al., 2001; Richards et al., 2004; Hurles et al., 2005] have lower diversity with respect to θk parameter (95% CI intervals of θk are lower and do not straddle those of the other source populations). Additionally, the DT statistic is not significantly negative for SEA, and all demographic parameters indicate that MD is at demographic (mutation-drift) equilibrium. Interestingly, SEA and MD do not exhibit noticeably lower H and θπ than the other comparison populations. In fact, H and θπ were shown to be poorly affected by the decrease in effective population size, and consequently, by genetic drift or founder effect (Helgason et al., 2003). As for the Reunionese samples, MB and RIN exhibit diversity and demographic indices that are within the range of the Indian comparison populations (and of FR and SEC samples): MB and RIN present no signs of diversity erosion, genetic drift or founder effect. On the contrary, the ZB sample exhibits reduced diversity, especially when comparing its θk to θk associated to the GUJ sample; the θk value of the ZB falls into the range of SEA (which is known to have experienced genetic drift).

Table 3.  Genetic diversity and demographic statistics for the Reunionese and comparison population samples
Sample ID1nkSH ± SDη ± SDθπ ± SDθk[95% CI]DTFsRISSD
  1. Notes: 1MB, Malbar; ZB, Zarab; RIN, Reunionese with haplotypes of assumed Indian ancestry; SEI, South-eastern India; GUJ, Gujarat; SEA, South-eastern Africa; MD, Madagascar; SEC, South-eastern China; FR, France; 2The sequence range of the Malagasy data is 16090-16362 (see Hurles et al., 2005). As a proxy, we assumed that nps 16363-16365 were monomorphic in this sample; * p-value<0.05; ** p-value<0.02.

MB143101930.9919 ± 0.00220.0217 ± 0.01155.99 ± 3.18154.78[106.57-228.40]−2.03**−25.04**0.00930.0005
ZB4228490.9803 ± 0.00890.0221 ± 0.01196.13 ± 3.3037.66[19.12-77.47]−1.62*−15.25**0.01210.0034
RIN149100850.9907 ± 0.00240.0187 ± 0.01015.15 ± 2.51134.15[93.82-193.99]−2.06**−25.27**0.01060.0002
SEI14435171650.9914 ± 0.00070.0196 ± 0.01055.51 ± 2.62332.24[295.76-372.84]−2.08**−24.43**0.00930.0003
GUJ9179800.9961 ± 0.00250.0214 ± 0.01145.90 ± 2.84305.69[167.48-614.45]−2.04**−25.23**0.01070.0002
SEA416133770.9604 ± 0.00450.0293 ± 0.01518.07 ± 4.1667.32[53.84-84.25]−0.88−24.15**0.00370.0024
MD23714310.8844 ± 0.03140.0172 ± 0.00964.76 ± 2.387.93[3.68-16.37]−1.10−1.850.0441**0.0188**
SEC2131541070.9950 ± 0.00120.0248 ± 0.01306.85 ± 3.24251.59[184.52-347.16]−1.91**−24.65**0.13300.0012
FR4372161090.9672 ± 0.00550.0138 ± 0.00773.82 ± 2.13169.20[138.10-207.11]−2.25**−25.31**0.01430.0004


  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information

The re-evaluation of the Indian mitochondrial phylogeny that we performed allowed the classification or reclassification of more than 20% (837/3883) of the southern Asian HTs present in our database. This phylogenetic re-evaluation was an essential prerequisite for optimising the accuracy of the estimation of the admixture coefficients. Indeed, the standard deviation of the admixture coefficients depends on the phylogeographical resolution of the mitochondrial gene pool under study (Salas et al., 2005). Moreover, our results contribute towards improving the phylogeographical resolution of the Southern Asia mtDNA.

People of Indian ancestry from the Mascarenes archipelago, where Reunion Island is located, form a non-negligible part of the worldwide Indian Diaspora (e.g. Leclerc, 2006). The Reunionese Indian Diaspora stems from different migration waves mainly motivated by the need for manpower induced by the plantation economy. The indenture system (Tinker, 1993; Lal, 2006) brought approximately 120,000 Indians to Reunion Island during the 19th century. By taking into account historical sources, which delineated the geographical origins of the Reunion Island population, we estimated the contribution of each of the parental regions. Most of the Reunionese mitochondrial gene pool of Indian ancestry (∼70%) is of Southern Indian origin (excluding the Zarab). More precisely, South-eastern India seems to have been the main region of recruitment, totalling more than 85% of the Southern India contribution. In this case, our results agree with the semi-quantitative historical investigations (e.g. Lacpatia, 1982). Our study also highlights Andhra Pradesh as the principal South-eastern region that provided workforce to Reunion. As a matter of fact, approximately 3/4 of the South-eastern Indian contribution originated from Andhra Pradesh, versus only 1/4 from Tamil Nadu. In this case, our results strongly suggest that historical studies (e.g. Dupon, 1974; Lacpatia, 1982), which pinpointed Tamil Nadu as the principal region involved in indenture recruitment, largely underestimated the volume of workers from Andhra Pradesh that were shipped to Reunion Island. Therefore, our study suggests that the South-eastern Indian recruitment area defined by historians (e.g. Dupon, 1974) must be revised and enlarged to include the entire current Andhra Pradesh State. In a similar way, we were able to show that the contribution from the region currently covered by Orissa remained very small. Indeed, the admixture coefficients for this parental population appeared quite negligible (<0.065; Table 2), and no M19 HTs were detected in Reunion Island. The indentured workers who embarked at Yanam (northern Andhra Pradesh) seem to have been enrolled in the current Southern districts of Orissa and North-eastern districts of Andhra Pradesh (Dupon, 1974; Lacpatia, 1982). Our study shows that the Orissan districts were in fact far less concerned by the indenture system than previously estimated, and that Yanam exported indenture workers mainly from northern Andhra Pradesh. The Malbar group is the main recipient of India-to-Reunion migrations that occurred from the end of the 17th century until the end of the 19th century: about 80% of the Malbar mtDNA pool is of Indian ancestry. The identity of this Reunionese group has so far been principally built on a vast common geographic origin: the Indian sub-continent (Barat, 1979). The Indian immigrants were of very diverse geographical and social origin. As far as the diverse social origins of the indentured workers are concerned, most of them were Dalits (Untouchables) or from tribal populations, but some indentured workers were also Shudras or even Brahmins (Dupon, 1974; Lacpatia, 1982; Benoist, 1988). Some Indian indentured workers were also Muslims, but they renounced their religion when in contact with the Hindu majority (Barat, 1989; Marimoutou-Oberlé, 1999; Govindin, 1994). The requirement to live in the same work camps, combined with a high female deficit among the Indian indentured workers (from 1 Indian woman for 55 Indian men in 1848 to 1:4 in 1883; Lacpatia, 1982), brought the various Indian geographical and social groups close together (Marimoutou-Oberlé, 1999). In Reunion Island, social status, and specifically caste, was no longer a relevant factor for marriages (Marimoutou, 1989). Consequently, children with parents of different Indian geographic and/or social origin were quite numerous (Lacpatia, 1987). All these factors, which led to extensive intra-Indian admixtures, tended to shape a single identity for the Malbar ethnic group (Barat, 1979). Nevertheless, this ‘Indian’ identity did not exclude extra-Indian admixture. We notably showed that a limited but non-negligible part of the Malbar mitochondrial genome (nearly 15%) is of Sub-Saharan and Malagasy ancestry. The introduction of non-Indian HTs into the Malbar group is partly explained by the recurrent female deficit in the 19th century. This sex bias led some Indian men to marry women of non-Indian origin (e.g. Dupon, 1974; Fuma, 2002). Sex-biased migration therefore appears as a key factor in the setting up of the Malbar ethnic group. As for the Muslim Gujarati migrants, they freely migrated to Reunion Island, and were not part of the indenture system context. The first Gujarati migrants also faced a female deficit, and some of them married Creole women (Ève, 1987; Ismaël-Daoudjee, 2002). Nonetheless, Islam rapidly established marital barriers around the Zarab ethnic group (Scherer, 1994). Moreover, our sampling criteria and protocol ensured that we discarded all the HTs inherited from well-known non-Gujarati ancestors (as well as non-Malbar ancestors for the MB sample), and therefore studied a non-admixed sample of the Zarab group.

Then, we detected three HTs of Southern Asian origin with high frequency in our ‘Other’ sample. Genealogical investigations allowed us to trace back each HT to its initial introduction in Reunion Island. All three HTs were introduced in Reunion Island by three well-identified Indian women in 1679 and 1689, and the current frequency of these HTs was conditioned by a founder effect that occurred between the end of the 17th century and 1720. During this time period, the effective size of the female population remained very low: only 34 immigrant women (including the three identified Indian women) had had offspring (Barassin, 1960; Ricquebourg, 2001). The Reunionese mitochondrial gene pool of Indian ancestry thus experienced a founder effect at the very beginning of the settlement of Reunion Island. Nevertheless, these three HTs have low or nil frequency within either Malbar or Zarab. Therefore, this early founder event may have affected the diversity of a few ethnic groups, but had no impact on the genetic diversity of the current Reunionese ethnic group of claimed Indian ancestry. By means of diversity and demographic parameters, we further explored the mtDNA of the Reunionese of Indian ancestry in order to detect whether other founder events had affected their genome variation. The samples from South-eastern India and Gujarat exhibited clear signs of population expansion, and all three Reunionese samples of Indian ancestry (RIN, MB and ZB) show clear signs of population expansion as well. Therefore, no founder effect has affected the Reunionese genetic pool of Indian ancestry except during the very start of the settlement of Reunion Island. Nevertheless, Zarab exhibit a low θk value, within the range of populations that are known to have experienced founder events (namely South-eastern Africa; Pereira et al., 2001; Richards et al., 2004). Genealogical investigations and sampling criteria however allowed us to exclude any potential genetic drift that occurred in Reunion Island either after the Gujarati people arrived or at the time of settlement (see also Dubut et al., 2009). Alternatively, it is very likely that our Zarab sample is representative of the Muslim Gujarati parental population from which these are known to have originated (Baruch and Surat districts; Ismaël-Daoudjee, 2002). In fact, the parental region of the Zarab, i.e. Gujarat, is little-represented to date (91 individuals: Kivisild et al., 1999; Metspalu et al., 2004; Quintana-Murci et al., 2004); moreover, this Gujarati limited sample does not include Muslims. And yet, the Indian genetic pool is very complex and can be envisaged as the juxtaposition of numerous endogamous population pockets (Chaubey et al., 2007).

Finally, our results generally contrast with those obtained by Berniell-Lee et al. (2008), which have recently analysed the mtDNA variation of 41 Reunionese individuals. They assumed the Reunionese population is homogenous with respect to the intensity of the admixtures. They notably concluded that the mtDNA Reunionese genetic pool as a whole was deeply affected by Indian migration waves and by genetic drift (caused by a founder effect involving Indian women) during the 19th century. First of all, Berniell-Lee et al. (2008) overestimated the Indian contribution by misclassifying HTs with HVS-1 motif 086-148-223-259-278-319 (24.4%, 0.95CR = 13.9-39.5%, in their Reunionese sample). In fact, these HTs belong to M46 (Hill et al., 2007), a HG quite common in Madagascar (see data from Hurles et al., 2005) and not to M2 (a typical Southern Asian HG; Kivisild et al., 2003). Secondly, none of the three Reunionese samples we investigated (RIN, MB and ZB) experienced a founder effect or genetic drift inside Reunion Island. These contradictions highlight the fact that the Reunionese cannot be considered a homogenous population, and the importance of taking into account its ethnic stratification. Admittedly, the social and cultural barriers preventing extensive mixing between groups became more and more obsolete during the last third of the 20th century (Labache, 1999). Consequently, ethnic boundaries in Reunion Island can appear as quite penetrable nowadays, although they remained hard to cross during most of the history of Reunion Island (e.g. Benoist & Gerbeau, 1993). Differences between Berniell-Lee et al.'s (2008) study and ours underline that ethno-historical parameters (here, the ethnic group) must be taken into account, in order to maximise the sample variability and to reliably reconstruct the biological history of a socio-culturally stratified population. In fact, as was recently shown by empirical studies (Chaix et al., 2007; Marchani et al., 2008), it has been emphasized that these are cultural and social factors shaping the genetic diversity, rather than genetics acting on cultural and social structures (e.g. Benoist, 1966; Macbeth, 1993, 1997; Marks, 1995; Cavalli-Sforza, 1997). In order to study a multiethnic population, introducing an ethno-historical questionnaire during the sampling period is critical to achieving an accurate reflection of its colonisation history and subsequent admixtures. Following this approach, we have deciphered the Indian counterpart of the history of the Reunionese population. Nevertheless, when our results and those of Berniell-Lee et al. (2008) are considered together, they suggest very different demographic histories depending on each ethnic group. Consequently, further analyses, including other ethnic groups and other DNA markers (such as Y-chromosome polymorphisms), would be helpful in dissecting the complex history of the stratified Reunionese population, and will be very welcome for further investigating sex-biased admixture in human populations. Furthermore, to our knowledge, no genetic information concerning the Indian Diaspora is available to date. By exploring the mtDNA inheritance of ethnic groups (Malbar and Zarab) stemming from the Indian Diaspora, our study contributes data to fill this gap and provides information about the genetic diversity of this worldwide human population component.


  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information

Warm thanks go to all volunteers who contributed their DNA to this research, and to all those who helped us during the sample collection, especially Alexandre Vienne. We thank André Gilles and two anonymous reviewers for very helpful and valuable comments on this manuscript. We are indebted to Mait Metspalu for further information about his Indian database, and to Partha P. Majumder for providing HVS-1 sequences from Basu et al. (2003). We are grateful to Helen Conté-McArley and Thomas Blanc for their help with the English editing. This work was supported by the Conseil Régional de La Réunion. M.-D. Thionville was supported by the Conseil Général de La Réunion.


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  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information
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  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information


Table Appendix I.  List of comparison populations for HVS-1 sequences: location and sample size
Geographical regionsState or ProvincePopulation codeSizeReferences
South-eastern AfricaMozambiqueSEA416Pereira et al., 2001; Salas et al., 2002
Madagascar MD37Hurles et al., 2005
France FR437Rousselet & Mangin, 1998; Richards et al., 2000; Calì et al., 2001; Dubut et al., 2004
South-eastern China SEC213 
Fujian-54Wen et al., 2004
Guangdong-135Kivisild et al., 2002; Yao et al., 2002; Wen et al., 2005
Hong-Kong-24Betty et al., 1996
India -3721 
Andhra PradeshANP1016Bamshad et al., 1998; Thangaraj et al., 1999; Cordaux et al., 2003; Kivisild et al., 1999, 2003; Wooding et al., 2004; Sharma et al., 2005; Thanseem el al., 2006
Arunachal PradeshARP26Cordaux et al., 2003
AssamASS58Cordaux et al., 2003
West BengalWEB285Kivisild et al., 1999; Roychoudhury et al., 2000; Basu et al., 2003; Metspalu et al., 2004
BiharBIH45Kivisild et al., 1999; Basu et al., 2003; Sharma et al., 2005
GujaratGUJ91Kivisild et al., 1999; Metspalu et al., 2004; Quintana-Murci et al., 2004
Himachal PradeshHIM37Metspalu et al., 2004
Jammu & KashmirKSH19Kivisild et al., 1999
KarnatakaKAR201Mountain et al., 1995; Cordaux et al., 2003
KeralaKER230Mountain et al., 1995; Cordaux et al., 2003; Metspalu et al., 2004
Madhya PradeshMAD82Roychoudhury et al., 2000; Basu et al., 2003
MaharashtraMAH221Kivisild et al., 1999; Basu et al., 2003; Baig et al., 2004; Metspalu et al., 2004
ManipurMAN9Basu et al., 2003
MizoramMIZ14Basu et al., 2003
NagalandNAG43Cordaux et al., 2003
OrissaORI153Kivisild et al., 1999; Basu et al., 2003; Sahoo & Kashyap, 2006
PunjabPJB362Kivisild et al., 1999; Kaur et al., 2002; Basu et al., 2003; Cordaux et al., 2003; Metspalu et al., 2004; Sharma et al., 2005
RajastanRAJ36Kivisild et al., 1999; Metspalu et al., 2004
Tamil NaduTN427Kivisild et al., 1999; Roychoudhury et al., 2000; Basu et al., 2003; Cordaux et al., 2003; Metspalu et al., 2004; Sharma et al., 2005; Watkins et al., 2008
TripuraTRP134Roychoudhury et al., 2000; Basu et al., 2003; Cordaux et al., 2003
Uttar PradeshUTP232Kivisild et al., 1999; Basu et al., 2003; Metspalu et al., 2004; Sharma et al., 2005
Bangladesh BAN30Kivisild et al., 1999; Cordaux et al., 2003
Sri Lanka SRL132Metspalu et al., 2004
Table Appendix II.  Key for haplogroup classification from HVS-1 motif in Eurasian context
HaplogroupHVS-1 motif (minus 16000)1,2Reference3
  1. Notes: 1Unless specified, numbers indicate transitions; 2within parentheses, nucleotide positions whose availability depends on publications; 3unless specified, haplogroup classification from Richards et al. (2000), Metspalu et al. (2004), Palanichamy et al. (2004), Quintana-Murci et al. (2004), Kong et al. (2006) and Thangaraj et al. (2006).

B4c1b3140-189-217-274-335Hill et al., 2007
HV10304-311this study
HV11 or U4356this study
M2a1223-270-274-319-352Kumar et al., 2008
M2a1a223-270-319-352Kumar et al., 2008
M2a1a1223-243-270-319-352Kumar et al., 2008
M2a2223-240C-274-311-319Kumar et al., 2008
M2a3a223-265C-274-319Kumar et al., 2008
M2b2223-274-295-319-320Kumar et al., 2008
M4a1145-176-223-261-311this study
M4c184-223-311this study
M5 or M10129-223-311 
M5a1129-223-291this study
M5a2a129-223-265Cthis study
M5a2b129-144A-223this study
M5b(048)-129-223this study
M5b1(048)-129-218-223this study
M6a*223-231-362Thangaraj et al., 2008
M6a1223-231-356-362this study
M6a2223-231-288-356-362this study
M6b184-223-311-362Thangaraj et al., 2008
M19189-223-300this study
M19a189-194-300this study
M20a111-192-223this study
M20b051-223-316this study
M20b1051-189-223-316this study
M26223-275-327Athis study
M30b192-223-278Sun et al., 2006
M30c1166d-223this study
M30d1179d-223this study
M30e223-368this study
M33a1169-172-223this study
M39b166-223-311Thanseem et al., 2006; this study
M43093-311-362 or 129-213-249-293CThangaraj et al., 2008
M45a129-209-223Hill et al., 2007
M46223-278Hill et al., 2007
N5 or M30111-223-311 
N5a051-092-111-223-311this study
N5b111-144-223-311this study
N9a6257A-261-292Hill et al., 2007
N9a6a257A-261-292-294Hill et al., 2007
R5a/R5a1266-304Chaubey et al., 2008
R5a2266-304-356Chaubey et al., 2008
R5a2a266-304-311-356Chaubey et al., 2008
R5a2b266-304-325-356this study
R5a2b1266-304-309-325-356this study
R5a2c256-266-304-356this study
R6a1a129-266-318-320-362this study
R6a2129-213-362this study
R6b227-245-266-278-362this study
R7b260-261-311-319-362Chaubey et al., 2008
R9c157-304Wen et al., 2004
R21168-295-304Hill et al., 2007
R22249-288Hill et al., 2007
R23256-290-(465)Hill et al., 2007
R30a1172-278this study
R30b1129-298-299this study
R30b2292-(497)this study
R30c126-181-209this study
R30d192-291-390this study
R31a172-304-362Chaubey et al., 2008
R31b051-189-218this study
T2b1126-294-296-325this study
U2b1051-209-239this study
U2b2051-168this study
U2c1051-179-234-240Cthis study
U2c2051-234-247this study
U5b1bc189-270Achilli et al., 2005

Supporting Information

  1. Top of page
  2. Summary
  3. Introduction
  4. Materials and Methods
  5. Results
  6. Discussion
  7. Acknowledgments
  8. References
  9. Appendix
  10. Supporting Information

Table S1 Primers and protocols for ASPCR genotyping of the mitochondrial coding regions.

Table S2 PCR protocols for amplification of the mitochondrial genome.

Table S3 Nucleotide positions of the mtDNA coding regions typed by sequencing.

Table S4 Indian HVS-1 database used for admixture and phylogeographical analyses.

Table S5 mtDNA polymorphisms in Malbar and Zarab groups and for the haplotypes of assumed Indian ancestry in the other Reunionese ethnic groups.

Table S6 mtDNA characteristics and TMRCA for mitochondrial haplogroups defined in this study.

Figure S1 Median-Joining networks and revised phylogeny of (A) R5a, (B) R6a and R6b, (C) M5, (D) M19 and (E) M20a and M20b lineages.

Please note:Wiley-Blackwell are not responsible for the content or functionality of any supporting materials supplied by the authors. Any queries (other than missing material) should be directed to the corresponding author for the article.

AHG_519_sm_TableS1.pdf30KSupporting info item
AHG_519_sm_TableS2.pdf23KSupporting info item
AHG_519_sm_TableS3.pdf120KSupporting info item
AHG_519_sm_TableS4.xls611KSupporting info item
AHG_519_sm_TableS5.pdf39KSupporting info item
AHG_519_sm_TableS6.pdf93KSupporting info item
AHG_519_sm_FigureS1.pdf217KSupporting info item

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