Monomorphic and polymorphic enzyme genetic markers of the Waiãpi Indians of Amapá and of inhabitants of Manaus, Amazonas

Mestriner, Moacyr A.; Salzano, Francisco M.

doi:10.1590/S1415-47571998000300003

Abstracts

One hundred nineteen Waiãpi Indians and 348 inhabitants of Manaus, Amazonas, were tested using nine enzyme genetic systems in the first population and two in the second. The Waiãpi showed some marked differences compared to South American Indian averages in ACP*A (1% vs. 14 ± 9%), GLO1*1 (9% vs. 30 ± 14%) and ESD*1 (99.6% vs. 70 ± 17%), while ESD*1 and CA2*1 prevalences in the Manaus population were within the range previously found in the northern region of Brazil (ESD*1: 85% vs. 80-94%; CA2*1: 98% vs. 89-99.8%). The almost identical frequencies obtained for these two markers in this population, when the sample was subdivided according to skin color, suggest that such morphological classifications have little value in this region. Genetic distances between the Waiãpi and other tribes, obtained using these and other genetic markers, showed good parallelism with the geographical distances which separate these Indians from the other groups considered.

Um total de 119 índios Waiãpi e 348 habitantes de Manaus, Amazonas, foram testados usando nove sistemas genéticos enzimáticos na primeira população e dois na segunda. Os Waiãpi mostraram algumas diferenças marcantes quanto às médias dos indígenas sul-americanos em ACP*A (1% vs. 14 ± 9%), GLO1*1 (9% vs. 30 ± 14%) e ESD*1 (99,6% vs. 70 ± 17%), enquanto as prevalências de ESD*1 e CA2*1 na população de Manaus mostraram-se dentro do intervalo encontrado previamente na região norte do Brasil (ESD*1: 85% vs. 80-94%; CA2*1: 98% vs. 89-99.8%). As freqüências quase idênticas obtidas para esses dois marcadores nesta população, quando a amostra foi subdividida de acordo com a cor da pele, sugerem que tais classificações morfológicas têm pouco valor nesta região. Distâncias genéticas entre os Waiãpi e outras tribos, obtidas usando esses e outros marcadores genéticos, mostraram bom paralelismo com as distâncias geográficas que separam esses índios dos outros grupos considerados.

SHORT COMMUNICATION

Monomorphic and polymorphic enzyme genetic markers of the Waiãpi Indians of Amapá and of inhabitants of Manaus, Amazonas^* * This paper is dedicated to our colleague Dr. Maria José de Melo e Freitas, who collected the Manaus sample and collaborated with us in many projects. Her untimely death is mourned by all who had the privilege of sharing with her the joys of life.

Moacyr A. Mestriner¹ and Francisco M. Salzano²

¹Departamento de Genética, Faculdade de Medicina, Universidade de São Paulo, Campus Universitário, 14049-900 Ribeirão Preto, SP, Brasil.

²Departamento de Genética, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, Caixa Postal 15053, 91501-970 Porto Alegre, RS, Brasil. Fax: 051 339-2011. E-mail: salzano@if1.if.ufrgs.br Send correspondence to F.M.S.

ABSTRACT

One hundred nineteen Waiãpi Indians and 348 inhabitants of Manaus, Amazonas, were tested using nine enzyme genetic systems in the first population and two in the second. The Waiãpi showed some marked differences compared to South American Indian averages in ACP*A (1% vs. 14 ± 9%), GLO1*1 (9% vs. 30 ± 14%) and ESD*1 (99.6% vs. 70 ± 17%), while ESD*1 and CA2*1 prevalences in the Manaus population were within the range previously found in the northern region of Brazil (ESD*1: 85% vs. 80-94%; CA2*1: 98% vs. 89-99.8%). The almost identical frequencies obtained for these two markers in this population, when the sample was subdivided according to skin color, suggest that such morphological classifications have little value in this region. Genetic distances between the Waiãpi and other tribes, obtained using these and other genetic markers, showed good parallelism with the geographical distances which separate these Indians from the other groups considered.

INTRODUCTION

Development of meaningful evolutionary histories for mankind depends on the painstaking accumulation of genetic data. We have previously studied the Waiãpi Indians of Amapá for 20 blood groups, enzymatic and non-enzymatic blood proteins, as well as HLA (Black et al., 1983), and the population of Manaus (tested for seven such systems; Santos et al., 1983). However, information related to nine genetic systems among the Waiãpi and two for the population of Manaus remained unpublished. This is unfortunate, since for population comparisons the reliability of the inferred relationships generally depends on the number of markers considered. In addition, some markers such as the carbonic anhydrase 2 alleles can provide useful evidence about interethnic admixture, a process that undoubtedly influenced the gene pools of Brazilian groups. This note, therefore, puts on record the indicated data.

SUBJECTS AND METHODS

The Waiãpi (Oyampi, Wayampi, Wayãpy) Indians inhabit an area that was specifically delimited for them, around geographical coordinates 1^oN, 53^oW. Their present population is estimated as approximately 500 individuals, with an equivalent number of members of this tribe living in French Guiana (Ricardo, 1996). They speak a Tupi language, and contacts of a more permanent nature with non-Indians started in 1973. Their land, however, is practically intact and free of intruders. In the 580 thousand hectares of tropical forest they have at their disposal, they hunt and gather, practice rudimentary agriculture, and recently have engaged in small-scale gold extraction. This latter activity is subject to much controversy (Gallois, 1996, 1997). Aspects of the Waiãpi schooling process, especially the integration of the Indians' objectives with a need to adapt to the surrounding non-Indian society, have been dealt with by Kahn (1996). Genetic studies of the French Guiana population were performed by Tchen et al. (1978a,b,c, 1981), while Black et al. (1983) studied the Brazilian groups.

Manaus (geographical coordinates: 3^o8'S; 60^o2'W) is the capital of the Brazilian State of Amazonas. Founded in 1669, its demographic evolution involved the contribution of Indians, Europeans (mainly Portuguese), and African-derived individuals. Present population is around one million persons. The sample reported here is composed of control subjects who were studied for a comparison with malaria patients (Santos et al., 1983).

Bloods were collected in 1978-1980 (Manaus) and 1980-1981 (Waiãpi) with anti-coagulant, refrigerated shortly afterwards, and transported in this condition to Porto Alegre (in the case of the Indian samples via Belém) and then to Ribeirão Preto. The typing procedures are given in Harris and Hopkinson (1976), and were performed in 1984.

RESULTS AND DISCUSSION

The Waiãpi, similar to most other South American Indians (Salzano and Callegari-Jacques, 1988) were found to be monomorphic for esterase A, carbonic anhydrase 2, and peptidases A, B, and C (Table I). The frequency of GPT*1 (40%) was similar to the average obtained for Amazonian Indians in general (18 groups, 50 ± 14%; Callegari-Jacques et al., 1994). Three other markers, however, showed quite different frequencies (ACP*A: 1% vs. 14 ± 9%, 69 groups; GLO1*1: 9% vs. 30 ± 14%, 39 groups; ESD*1: 99.6% vs. 70 ± 17%, 61 groups). All averages refer to South American Indians in general, as reported in Santos et al. (in press). Tchen et al. (1978b) examined five of the systems studied here in the Waiãpi of French Guiana. As in the present study, no variation was found for peptidases A and B. However, for three of the markers, they found values which were more similar to the general averages (46% for GPT*1, 14% for ACP*A and 88% for ESD*1).

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Santos et al. (in press) compared the Waiãpi with 15 other Tupi-speaking populations, using 21 genetic systems (ABO, ACP, AK, ALB, CA2, CHE1, CP, ESD, Duffy, G6PD, GLO, HBA, HBA2, HP, P, PGD, PGM1, PGM2, PEPA, PEPB, TF). They showed a closest relationship with the Araweté, followed by the Urubu-Kaapor, and more distantly the Gavião. The two first links make sense, since the three populations do not live very far away from each other, but the Gavião are more distant, being located in the southeastern portion of the Amazon Region. This distance is greater than those which separate the Waiãpi from other Tupi tribes considered in the comparison. Callegari-Jacques and Salzano (1989) performed another, more restricted, within-Tupi comparison (eight populations, 12 systems), and in this case the Waiãpi clustered with their neighbors, the Emerillon (not included in the 1998 study). In another analysis involving 27 Amazonian Indian tribes (Salzano et al., 1991), the Waiãpi clustered with the Wayana, Apalai and Tiriyó (all Carib tribes living in their vicinity), but also with the Pano, who live far away.

The values obtained in the total Manaus sample (85% for ESD*1; 98% for CA2*1; Table I) are within the range obtained in northern Brazil (ESD*1: 80-94, eight populations; CA2*1: 89-99.8%, six populations; Schüler et al., 1982; Schneider et al., 1987; Santos et al., 1987; Bortolini et al., 1992; Guerreiro et al., 1993; Ribeiro-dos-Santos et al., 1995; Santos et al., 1996; Bortolini et al., in press). The CA2*1 results do not strictly correlate with the amount of African admixture estimated for these communities (see Santos and Guerreiro, 1995; Bortolini et al., 1995; Bortolini et al., in press), although, as expected, high CA2*1 frequencies are generally associated with low African admixture (Óbidos: 10% and CA2*1: 99%; Manaus: 12% and 98%; Parintins: 13% and 99.8%), the opposite also being true (Trombetas: 56% and 89%). The almost identical frequencies observed for ESD*1 and CA2*1 in the three subgroups classified according to color in Manaus in our study (Table I) indicate that such groupings are meaningless in this population.

ACKNOWLEDGMENTS

We thank the Fundação Nacional do Índio (FUNAI) for permission to study the Waiãpi Indians and for logistical assistance. The Indian leaders and subjects of the investigation were adequately informed about the aims of the study and gave their approval, which is also gratefully acknowledged. Prof. Francis L. Black collected the Waiãpi samples, and Denise A. Conte helped in the laboratory determinations of the Manaus material. Financial aid was provided by Programa de Apoio a Núcleos de Excelência (PRONEX), Financiadora de Estudos e Projetos (FINEP), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), and Fundação de Amparo à Pesquisa do Estado do Rio Grande do Sul (FAPERGS). Publication supported by FAPESP.

RESUMO

Um total de 119 índios Waiãpi e 348 habitantes de Manaus, Amazonas, foram testados usando nove sistemas genéticos enzimáticos na primeira população e dois na segunda. Os Waiãpi mostraram algumas diferenças marcantes quanto às médias dos indígenas sul-americanos em ACP*A (1% vs. 14 ± 9%), GLO1*1 (9% vs. 30 ± 14%) e ESD*1 (99,6% vs. 70 ± 17%), enquanto as prevalências de ESD*1 e CA2*1 na população de Manaus mostraram-se dentro do intervalo encontrado previamente na região norte do Brasil (ESD*1: 85% vs. 80-94%; CA2*1: 98% vs. 89-99.8%). As freqüências quase idênticas obtidas para esses dois marcadores nesta população, quando a amostra foi subdividida de acordo com a cor da pele, sugerem que tais classificações morfológicas têm pouco valor nesta região. Distâncias genéticas entre os Waiãpi e outras tribos, obtidas usando esses e outros marcadores genéticos, mostraram bom paralelismo com as distâncias geográficas que separam esses índios dos outros grupos considerados.

(Received January 5, 1998)

Black, F.L., Salzano, F.M., Berman, L.L., Gabbay, Y., Weimer, T.A., Franco, M.H.L.P. and Pandey, J.P. (1983). Failure of linguistic relationships to predict genetic distances between the Waiăpi and other tribes of lower Amazonia. Am. J. Phys. Anthropol. 60: 327-335.
Bortolini, M.C., Weimer, T.A., Franco, M.H.L.P., Salzano, F.M., Layrisse, Z., Schneider, H., Schneider, M.P.C. and Harada, M.L. (1992). Genetic studies in three South American black populations. Gene Geogr. 6: 1-16.
Bortolini, M.C., Weimer, T.A., Salzano, F.M., Callegari-Jacques, S.M., Schneider, H., Layrisse, Z. and Bonatto, S.L. (1995). Evolutionary relationships between black South American and African populations. Hum. Biol. 67: 547-559.
Bortolini, M.C., Silva-Junior, W.A., Weimer, T.A., Zago, M.A., Castro de Guerra, D., Schneider, M.P.C., Layrisse, Z., Mendez Castellano, H. and Salzano, F.M. Protein and hypervariable tandem repeat diversity in eight African derived South American populations: Inferred relationships do not coincide. Hum. Biol. (in press).
Callegari-Jacques, S.M. and Salzano, F.M. (1989). Genetic variation within two linguistic Amerindian groups: Relationship to geography and population size. Am. J. Phys. Anthropol. 79: 313-320.
Callegari-Jacques, S.M., Salzano, F.M., Weimer, T.A., Hutz, M.H., Black, F.L., Santos, S.E.B., Guerreiro, J.F., Mestriner, M.A. and Pandey, J.P. (1994). Further blood genetic studies on Amazonian diversity ^_ data from four Indian groups. Ann. Hum. Biol. 21: 465-481.
Gallois, D.T. (1996). Controle territorial e diversificaçăo do extrativismo na área indígena Waiăpi. In: Povos Indígenas no Brasil, 1991/1995 (Ricardo, C.A., ed). Instituto Socioambiental, Săo Paulo, pp. 263-271.
Gallois, D.T. (1997). Campanha contra o projeto de autonomia dos índios Waiăpi, Amapá. Bol. Assoc. Bras. Antropol. 28: 11-12.
Guerreiro, J.F., Ribeiro-dos Santos, A.K.C., Santos, E.J.M., Cayres, I.M.V. and Santos, S.E.B. (1993). Genetic structure and demography of the human population of Óbidos, in the Brazilian Amazon. Rev. Bras. Genet. 16: 1075-1084.
Harris, H. and Hopkinson, D.A. (1976). Handbook of Enzyme Electrophoresis in Human Genetics. North-Holland, Amsterdam.
Kahn, M. (1996). A escolarizaçăo Waiăpi. In: Povos Indígenas no Brasil, 1991/1995 (Ricardo, C.A., ed.). Instituto Socioambiental, Săo Paulo, pp. 272-274.
Ribeiro-dos-Santos, A.K.C., Santos, E.J.M., Guerreiro, J.F. and Santos, S.E.B. (1995). Demographic and genetic structure of the population of Castanhal, in the Amazon Region of Brazil. Braz. J. Genet. 18: 469-474.
Ricardo, C.A. (1996). Povos Indígenas no Brasil, 1991/1995. Instituto Socioambiental, Săo Paulo.
Salzano, F.M. and Callegari-Jacques, S.M. (1988). South American Indians. A Case Study in Evolution. Clarendon Press, Oxford.
Salzano, F.M., Black, F.L., Callegari-Jacques, S.M., Santos, S.E.B., Weimer, T.A., Mestriner, M.A., Kubo, R.R., Pandey, J.P. and Hutz, M.H. (1991). Blood genetic systems in four Amazonian tribes. Am. J. Phys. Anthropol. 85: 51-60.
Santos, S.E.B. and Guerreiro, J.F. (1995). The indigenous contribution to the formation of the population of the Brazilian Amazon region. Rev. Bras. Genet. 18: 311-315.
Santos, S.E.B., Salzano, F.M., Franco, M.H.L.P. and Freitas, M.J.M. (1983). Mobility, genetic markers, susceptibility to malaria and race admixture in Manaus, Brazil. J. Hum. Evol. 12: 373-381.
Santos, S.E.B., Guerreiro, J.F., Salzano, F.M., Weimer, T.A., Hutz, M.H. and Franco, M.H.L.P. (1987). Mobility, blood genetic traits and race mixture in the Amazonian population of Oriximiná. Rev. Bras. Genet. 10: 745-759.
Santos, E.J.M., Ribeiro-dos-Santos, A.K.C., Guerreiro, J.F., Aguiar, G.F.S. and Santos, S.E.B. (1996). Migration and ethnic change in an admixed population from the Amazon region (Santarém, Pará). Braz. J. Genet. 19: 511-515.
Santos, S.E.B., Ribeiro-dos-Santos, A.K.C., Guerreiro, J.F., Weimer, T.A., Callegari-Jacques, S.M., Mestriner, M.A., Franco, M.H.L.P., Hutz, M.H. and Salzano, F.M. New protein genetic studies in six Amazonian Indian populations. Ann. Hum. Biol. (in press).
Schneider, H., Guerreiro, J.F., Santos, S.E.B., Weimer, T.A., Schneider, M.P.C. and Salzano, F.M. (1987). Isolate breakdown in Amazonia _ the Blacks of the Trombetas River. Rev. Bras. Genet. 10: 565-574.
Schüler, L., Salzano, F.M., Franco, M.H.L.P., Freitas, M.J.M., Mestriner, M.A. and Simőes, A.L. (1982). Demographic and blood genetic characteristics in an Amazonian population. J. Hum. Evol. 11: 549-558.
Tchen, P., Bois, E., Feingold, N., Grenand, F. and Degos, L. (1978a). Histocompatibility antigens in two American Indian tribes of French Guiana. Tissue Antigens 11: 315-319.
Tchen, P., Bois, E., Séger, J., Grenand, P., Feingold, N. and Feingold, J. (1978b). A genetic study of two French Guiana Amerindian populations. I. Serum proteins and red cell enzymes. Hum. Genet. 45: 305-315.
Tchen, P., Séger, J., Bois, E., Grenand, F., Fribourg-Blanc, A. and Feingold, N. (1978c). A genetic study of two French Guiana Amerindian populations. II. Rare electrophoretic variants. Hum. Genet. 45: 317-326.
Tchen, P., Bois, E., Lanset, S. and Feingold, N. (1981). Blood group antigens in the Emerillon, Wayampi and Wajana Amerindians of French Guiana. Hum. Hered. 31: 47-53.

*

This paper is dedicated to our colleague Dr. Maria José de Melo e Freitas, who collected the Manaus sample and collaborated with us in many projects. Her untimely death is mourned by all who had the privilege of sharing with her the joys of life.

Publication Dates

Publication in this collection
23 Feb 1999
Date of issue
Sept 1998

History

Received
05 Jan 1998

This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License.

[1] Black, F.L., Salzano, F.M., Berman, L.L., Gabbay, Y., Weimer, T.A., Franco, M.H.L.P. and Pandey, J.P. (1983). Failure of linguistic relationships to predict genetic distances between the Waiăpi and other tribes of lower Amazonia. Am. J. Phys. Anthropol. 60: 327-335.

[2] Bortolini, M.C., Weimer, T.A., Franco, M.H.L.P., Salzano, F.M., Layrisse, Z., Schneider, H., Schneider, M.P.C. and Harada, M.L. (1992). Genetic studies in three South American black populations. Gene Geogr. 6: 1-16.

[3] Bortolini, M.C., Weimer, T.A., Salzano, F.M., Callegari-Jacques, S.M., Schneider, H., Layrisse, Z. and Bonatto, S.L. (1995). Evolutionary relationships between black South American and African populations. Hum. Biol. 67: 547-559.

[4] Bortolini, M.C., Silva-Junior, W.A., Weimer, T.A., Zago, M.A., Castro de Guerra, D., Schneider, M.P.C., Layrisse, Z., Mendez Castellano, H. and Salzano, F.M. Protein and hypervariable tandem repeat diversity in eight African derived South American populations: Inferred relationships do not coincide. Hum. Biol. (in press).

[5] Callegari-Jacques, S.M. and Salzano, F.M. (1989). Genetic variation within two linguistic Amerindian groups: Relationship to geography and population size. Am. J. Phys. Anthropol. 79: 313-320.

[6] Callegari-Jacques, S.M., Salzano, F.M., Weimer, T.A., Hutz, M.H., Black, F.L., Santos, S.E.B., Guerreiro, J.F., Mestriner, M.A. and Pandey, J.P. (1994). Further blood genetic studies on Amazonian diversity ^_ data from four Indian groups. Ann. Hum. Biol. 21: 465-481.

[7] Gallois, D.T. (1996). Controle territorial e diversificaçăo do extrativismo na área indígena Waiăpi. In: Povos Indígenas no Brasil, 1991/1995 (Ricardo, C.A., ed). Instituto Socioambiental, Săo Paulo, pp. 263-271.

[8] Gallois, D.T. (1997). Campanha contra o projeto de autonomia dos índios Waiăpi, Amapá. Bol. Assoc. Bras. Antropol. 28: 11-12.

[9] Guerreiro, J.F., Ribeiro-dos Santos, A.K.C., Santos, E.J.M., Cayres, I.M.V. and Santos, S.E.B. (1993). Genetic structure and demography of the human population of Óbidos, in the Brazilian Amazon. Rev. Bras. Genet. 16: 1075-1084.

[10] Harris, H. and Hopkinson, D.A. (1976). Handbook of Enzyme Electrophoresis in Human Genetics. North-Holland, Amsterdam.

[11] Kahn, M. (1996). A escolarizaçăo Waiăpi. In: Povos Indígenas no Brasil, 1991/1995 (Ricardo, C.A., ed.). Instituto Socioambiental, Săo Paulo, pp. 272-274.

[12] Ribeiro-dos-Santos, A.K.C., Santos, E.J.M., Guerreiro, J.F. and Santos, S.E.B. (1995). Demographic and genetic structure of the population of Castanhal, in the Amazon Region of Brazil. Braz. J. Genet. 18: 469-474.

[13] Ricardo, C.A. (1996). Povos Indígenas no Brasil, 1991/1995. Instituto Socioambiental, Săo Paulo.

[14] Salzano, F.M. and Callegari-Jacques, S.M. (1988). South American Indians. A Case Study in Evolution. Clarendon Press, Oxford.

[15] Salzano, F.M., Black, F.L., Callegari-Jacques, S.M., Santos, S.E.B., Weimer, T.A., Mestriner, M.A., Kubo, R.R., Pandey, J.P. and Hutz, M.H. (1991). Blood genetic systems in four Amazonian tribes. Am. J. Phys. Anthropol. 85: 51-60.

[16] Santos, S.E.B. and Guerreiro, J.F. (1995). The indigenous contribution to the formation of the population of the Brazilian Amazon region. Rev. Bras. Genet. 18: 311-315.

[17] Santos, S.E.B., Salzano, F.M., Franco, M.H.L.P. and Freitas, M.J.M. (1983). Mobility, genetic markers, susceptibility to malaria and race admixture in Manaus, Brazil. J. Hum. Evol. 12: 373-381.

[18] Santos, S.E.B., Guerreiro, J.F., Salzano, F.M., Weimer, T.A., Hutz, M.H. and Franco, M.H.L.P. (1987). Mobility, blood genetic traits and race mixture in the Amazonian population of Oriximiná. Rev. Bras. Genet. 10: 745-759.

[19] Santos, E.J.M., Ribeiro-dos-Santos, A.K.C., Guerreiro, J.F., Aguiar, G.F.S. and Santos, S.E.B. (1996). Migration and ethnic change in an admixed population from the Amazon region (Santarém, Pará). Braz. J. Genet. 19: 511-515.

[20] Santos, S.E.B., Ribeiro-dos-Santos, A.K.C., Guerreiro, J.F., Weimer, T.A., Callegari-Jacques, S.M., Mestriner, M.A., Franco, M.H.L.P., Hutz, M.H. and Salzano, F.M. New protein genetic studies in six Amazonian Indian populations. Ann. Hum. Biol. (in press).

[21] Schneider, H., Guerreiro, J.F., Santos, S.E.B., Weimer, T.A., Schneider, M.P.C. and Salzano, F.M. (1987). Isolate breakdown in Amazonia _ the Blacks of the Trombetas River. Rev. Bras. Genet. 10: 565-574.

[22] Schüler, L., Salzano, F.M., Franco, M.H.L.P., Freitas, M.J.M., Mestriner, M.A. and Simőes, A.L. (1982). Demographic and blood genetic characteristics in an Amazonian population. J. Hum. Evol. 11: 549-558.

[23] Tchen, P., Bois, E., Feingold, N., Grenand, F. and Degos, L. (1978a). Histocompatibility antigens in two American Indian tribes of French Guiana. Tissue Antigens 11: 315-319.

[24] Tchen, P., Bois, E., Séger, J., Grenand, P., Feingold, N. and Feingold, J. (1978b). A genetic study of two French Guiana Amerindian populations. I. Serum proteins and red cell enzymes. Hum. Genet. 45: 305-315.

[25] Tchen, P., Séger, J., Bois, E., Grenand, F., Fribourg-Blanc, A. and Feingold, N. (1978c). A genetic study of two French Guiana Amerindian populations. II. Rare electrophoretic variants. Hum. Genet. 45: 317-326.

[26] Tchen, P., Bois, E., Lanset, S. and Feingold, N. (1981). Blood group antigens in the Emerillon, Wayampi and Wajana Amerindians of French Guiana. Hum. Hered. 31: 47-53.