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

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 nonenzymatic 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information related to nine genetic systems among the Waiãpi and two for the population of Manaus remained unpublished.This is unfortunate, since for population comparisons the reliability of the inferred relationships generally depends on the number of markers considered.In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 o N, 53 o W. 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(Gallois, , 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. (1978aTchen et al. ( ,b,c, 1981)), while Black et al. (1983) studied the Brazilian groups.
Manaus (geographical coordinates: 3 o 8'S; 60 o 2'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).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.