Electrophoretic Variants in Three Amerindian Tribes: the Baniwa, Kanamari, and Central Pano of Western Brazil

Home , Baniwa, Macushi, Ticuna, Wapishana

Electrophoretic Variants in Three Amerindian Tribes: The Baniwa, Kanamari, and Central Pano of Western Brazil

HARVEY MOHRENWEISER,’ JAMES V. NEEL,‘ M. A. MESTRINER,‘ F. M. SALZANO,’ E. MI(:LIAZZA,4 A. L. SIMOES ’ AND C. M. YOSHIHARA ’ ’ Department of Human Genetics, Uniuerszty of Michigan Medical School, Ann Arbor, Michigan 48109; ‘Departmento de Genetrca, Faculdadr de Medicine, Uniuersidade de SiZn Paulo, Ribeirao Pretu, Srio Paulo, Brazil; Departmento de Genitica, Instltuto de Bincitkias, Uniuersidade Federal do Rlo Grande do Sul, 90000 Porto Alegre, R.S. Brazil and ‘Department of Anthropology, Uniuersity of Maryland, College Park, Maryland 20742

KEY WORDS Amerindian . Electrophoretic variant . Private polymorphism . Gene frequency

ABSTRACT Data are presented on electrophoretic variants of 25 polypeptides found in the blood serum and erythrocytes, in 812 individuals from three Amerindian tribes, the Pano, the Baniwa, and the Kanamari. Two “private polymorphisms” were encountered, of PEPB in the Pano and CAI1 in the Baniwa. A single example of a different PEPB variant was encountered in the Baniwa, and two possible examples of an unstable variant of HGB A2 in the Kanamari. In addition, the well-known A variant of ACP,, the Duarte variant of GALT, the 2 variant of Hp and the 2 variant of PGM, occurred in polymorphic proportions in all three tribes, and the TFDChl variant was present as a polymorphism in the Baniwa. These data have recently been incorporated into a treatment which concludes that the eight electrophoretically-defined “private polymorphisms” thus far encountered in Amerindian tribes can be explained by a mutation pressure of 0.7 x 10-5/locus/generation on the assump- tion of neutrality of the phenotypes in question (Neel and Thompson, ’78).

Most of the Amerindian tribes in close prox- the ultimate goal of assigning these groups imity of the great rivers of the Central their place in a nexus of Amerindian relation- Amazon Basin either disappeared shortly ships, and (3) by electrophoretic techniques, to after contacts with the Old World were initi- determine the frequency of genetic variants of ated, or, if they survived, underwent extensive a series of polypeptides, in a continuing effort admixture with neo-Brazilians. The central to define the genetic variation present at a position of these tribes in South American representative series of loci in Amerindians. geography renders them of especial interest In connection with this latter objective, one for attempts to reconstruct the movements of the salient findings in previous studies of and biological interrelationships of the tribes Amerindian tribes, by ourselves and others, of South America. In the summer of 1976 we has been the frequency of “private genetic were able to study representatives of four of polymorphisms,” apparently unique alleles these tribes (Central Pano, Kanamari, Bani- which within a single tribe or several adjacent wa, Ticuna) who, although relatively accul- tribes achieve gene frequencies well over 1% turated, are unusual in that they still have on the basis of a sample of some hundreds of undergone little or no admixture with neo- persons. Thus far in a systematic application Brazilians (< - 1%,Neel, ’78b; Gershowitz of electrophoretic techniques to an average of and Neel, ’78). The primary objectives were 25 proteins in ten tribes, we have encountered (1) a variety of medical studies, (2) typings six examples of this phenomenon: CRPLCAY with respect to 13 polymorphic systems, with (Salzano et al., ’72; Tanis et al., ’73; Ned,

AM. J. PHYS. ANTHROP. i1979) 50: 237-246 237 238 H. MOHRENWEISER ET AL.

'78b). ALBYAN2(Tanis et al., '74); ESAPMAC1 This report is restricted to the results of (Neel et al., '771, PEPAWAP1(Tanis et al., '73; surveying representatives of the Baniwa, Neel et al., '771, ALBMAK"(Tanis et al., '73; Kanamari, and Pano tribes, a total of 812 per- Neel et al., '771, and LDHEUA1(Tanis et al., sons, with respect to electrophoretic variants '77). Given the kind of knowledge of the breed- of 25 polypeptides. The most noteworthy finding structure and demography which we have ing is the demonstration of two more private developed on the basis of study of one unac- polymorphisms, one of peptidase B in the Pano culturated tribe, the Yanomama (Neel and and one of carbonic anhydrase I1 in the Weiss, '75; Neel, '78a1, it should on certain Baniwa. assumptions be possible to manipulate data of THE TRIBES this type to derive insights into the duration of the tribal genetic isolation and the muta- Baniwa tion rateshelective pressures consistent with At the time of first contact, the Rio Negro the findings (Neel and Thompson, '78). was lined with Arawak-speaking tribes from

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PERU . PAN0 1 ! 1 c a KANAMARI BOLIVIA ! 1 \, 65 60 Fig. 1 Map indicating present location of tribes and villages sampled. Further identification in text ELECTROPHORETIC VARIANTS IN AMERINDIAN TRIBES 239 its confluence with the Solimdes to form the linguistic groups. One, speaking Pano, is a Amazon, to its headwaters in what is now Col- subdivision of the Central Pano, whom we con- ombia. The Baniwa, who inhabit the area sider in the next section. One is a member of drained by the Iqana River, a tributary of the the Arawak-speaking tribes of the Upper Rio Negro in the extreme northwestern corner Purus River. The third group, the object of our of Brazil and adjacent Colombia, are one of the study, constitutes an independent tribe, very few of these tribes now extant. Classified whose language, Katukina, shows no close af- as "in permanent contact" (but not "inte- filiation with the other languages of the area; grated") by Gama Malcher in 1964, we would Greenberg ('60) classified the language as a now term them semi-acculturated. They are member of the Macro-Tucanovan family, the currently estimated to number some 1,500 other components of which are Ticuna and persons, distributed among some 16 villages, Tucano. They are a small tribe, estimated to in the area indicated in figure 1. They are or- number about 800 persons living in 19 villages ganized into 20 exogamous, patrilineal clans; located in the area shown in figure 1, with a the preferred marriage is with a paternal long history of contact with neo-Brazilians. cross-cousin. Ethnographic data will be found Our studies were carried out at a station of the in Goldman ('481, Galvao ('59), Oliviera and New Tribes Mission, at a location known as GalvSo ('731, and Noble ('62). This tribe was of Tres Unidos, on the Mamore Creek, a tribu- particular interest to us as a representative of tary of the Jurua River, some 25 air miles east the western neighbors of the Yanomama, of the town of Eirunepe. Approximate coordi- among whom a previously mentioned polymor- nates are 6"37'S, 69"32'W (cf. fig. 1). The sam- phism of serum albumin had been encoun- ple was obtained from representatives of three tered (Tanis et al., '74): would it have spread villages, totaling about 130 persons, who had to the Baniwa? The present collections were consolidated at this location following the es- carried out at a small station of the New tablishment of the Mission Station, in 1967. Tribes Mission on the Upper Iqana River, at The tribe, which has never been intensively latitude lo33'N,longitude 68"44'W (cf. fig. 1). studied ethnographically (characterized with The sample was obtained from representa- a single paragraph in Metraux's ('48) treat- tives from six villages who had gathered at the ment of the tribes of the Jurua-Purusbasin), station as a result of advance word of our mea- is now relatively acculturated, practicing sub- sles vaccination program. sistence agriculture. Kanamari Central Pano In the basins of the Jurua and Purus Rivers These Indians are located in the extreme in southwestern Amazonas State, Brazil, southwestern portion of Amazonas State and there are three different populations referred the western half of Acre State, Brazil, and in to as Kanamari. They belong to three different the adjacent portions of Peru. They are re-

TABLE 1

Data on the sampling of the Central Pano

Pano Brazilian Our subdivision place name I.ocation Comments designation

Cashinawa Cana Brava 8"7'S, 70"19'W New Tribes Station 40 A,B (villagessubdivided into Cana Brava and Pared&) Jaminawa Near Sete 8"17'S,71"34'W Village 6km down 35 B Estrelas Jurua River from Sete Estrelas Marubo Vida Nova 6'47'S, 72'8'W New Tribes Station 34 A-E Pano (Katukina) Sete Estrelas 8"17'S, 71"34'W New Tribes Station 35 A Morada Nova ' 8"9'S, 70'21'W Village 2 km up 41A Embira River from Feiji,

' Morada Nova IS In fact a "mixed" village recently established by Central Panu desinng proximity to the town of P'eijo. Of 60 adultti interviewed, 34 identified themselves 8s Katukina (Pano Proper), 10 as Cashinawa, 2 ns .Jaminawa. 5 as of mixed Pano ancestry. 1 as Ksshinawa/neo~Brdzilian,and 4 as Neo-Brazilian. None of the latter 5 was sampled. 240 H. MOHRENWEISER ET AL.

TABIL 2

Cognate density (percentages/ among fourgroups of the Central Pano

Croup

Mambo Pano Yaminawa

91 83 85 Kashinawa 87 84 90 ferred to by at least 15 different local names, each loosely considered a separate tribe. The approximate area they occupy is shown in figure 1. We sampled four different groups at five different locations, as shown in table 1. An analysis of cognate density on the basis of our modification of the Swadesh list of 200 items (Swadesh, ’55) revealed the situation shown in table 2. This degree of mutual in- telligibility is greater than that we encountered among the various subdivisions of the Yanomama (Spielman et al., ’741, and is a principal reason why we consider these various groups as all samples from a single tribe. The total number of Pano is about 18,000 (Kensinger, personal communication). The social structure of the various areas is, in general, similar. For instance, the Marubo (34A-E) are organized into nine matrilineal clans named after animals. Marriage is clan exogamous, preferentially with a cross-cousin. Residence is in communal houses containing representatives of two or more clans. Per- m w m tinent common features of their culture at NI-imd1wm time of first contact were slash-and-burnagri- d m culture depending heavily on sweet manioc and maize, the custom of each extended family occupying a single large house, and ritual endocannibalism and cremation of the dead. The principal sources of ethnographic data are Steward and Metraux (‘481, Steward and JzwNO Faron (’59) and Melatti and Melatti (’75). N Coincident with the pressures of the Rubber Room, a group of Cashinahua related to those we examined migrated to Peru, where they were studied by Johnston, Kensinger, and col- B leagues (cf. esp. Johnston et al., ’68, ’69). The Central Pano came into extensive contact with neo-Brazilians in the Seventeenth Century. Today the degree of acculturation of the various subgroups is quite variable. Gama Malcher in 1964 classified most groups as “in- tegrated’ but several still only had “sporadic contact.” Since then all groups have established permanent contacts with missions or ELECTROPHORETIC VARIANTS IN AMERINDIAN TRIBES 241 government posts. Among the groups we stud- tration of neo-Brazilians into these tribal ied, the Marubo are still culturally intact and areas, some intermarriages with non-Indians relatively unacculturated but the other have occurred. Although it was expedient to groups are now living much as the Brazilian of sample them in the field, the neo-Brazilians the interior. contracting these marriages and their off- METHODOLOGY spring have been excluded from the tabula- Blood samples were collected in Becton- tions. After these exclusions, the number of Dickinson vacutainers containing 2 ml of persons examined, each for all 25 of the sys- acid-citrate-dextrose anticoagulant and tems listed above, were as follows: Baniwa, chilled as soon as possible, usually within 12 377; Kanamari, 100; and Pano, 335 (Cashi- hours of collection. The samples were shipped nawa, '88; Jaminawa, 48; Marubo, 112; Pano, with ice to Ann Arbor where the plasma and 87). A total of 812 persons was examined, re- washed packed cells were stored at -80°C or sulting in 40,600 locus determinations. Except in liquid N, until typing. Samples were nor- for the findings noted in table 3 (common poly- mally processed for storage within seven days morphisms) and table 4 (private polymorphisms and rare variants) all individuals were of of collection. standard phenotypes. The findings will be de- The following erythrocyte proteins were scribed under three headings: studied in this laboratory: acid phosphatase-1 (ACP,), adenosine deaminase (ADA), adeny- Common polymorphisms late kinase-1 (AK), carbonic anhydrase-I Well known polymorphisms of five loci (CAI), carbonic anhydrase I1 (CAII), galac- (ACP, GALT, HP, PGM,, and TFI were tose-1-phosphate uridyltransferase (GALT), detected in various of the tribes. Phenotype hemoglobin A, (Hb, and Hb&, hemoglobin A, and gene frequencies for these polymorphisms (HB,, and HB&, isocitrate dehydrogenase are presented in table 3. The following points (ICD,) lactate dehydrogenase (LDHn and are of interest: (1)The Pano villages exhibited LDHB), malate dehydrogenase (MUHs), nu- the range of intratribal variation with respect cleoside phosphorylase (NP), peptidase A to the ACP, GALT, HP, and PGM, polymor- (PEPA) peptidase B (PEPB), phosphoglu- phisms which is characteristic of Amerin- comutase-1 (PGMI), phosphoglucomutase-2 dians, although the Jaminawa (35 B) are (PGM2), phosphogluconate dehydrogenase unusual in lacking three of these four poly- (PGD) , phosphoglucose isomerase (PHI) and morphisms. (2) The tribal values with respect triosephosphate isomerase (TPI). Four serum to these four polymorphisms were within the proteins, albumin (ALB), ceruloplasmin (CP), range characterizing Amerindians, with the haptoglobin (HP) and transferrin (TF), were exception of the Duarte allele of GALT, which also studied. In addition, esterases (A and D) in all three tribes was less common than has were examined in the laboratory of Mestriner; been the case in most tribes studied to date the results will be reported elsewhere. All of (Neel et al., '77; Tanis et al., '77). (3) A TF the electrophoretic methods were as previous- variant with mobility similar to the Dc~,vari- ly described (Tanis et al., '73; Neel et al., '76, ant when examined by either starch or poly- '77). acrylamide gel electrophoresis (fig. 2a) was RESULTS encountered in one of the three tribes, the In recent years, with the continuing pene- Baniwa, with an allelic frequency of 0.016.

TABLE 4

Systems in which only private variants were identified

Tribe/protein CA I1 HGBI; PEP B

2 BAN 1 Variants -~37BAN 1/+ -0 lBANl/+ Baniwa Total typed 377 377 377 Variants ___0 2KAN1/+(?) -0 Kanamari Totaltyped 100 100 100 Variants lCAII2/+ -0 16PAN 1/+ Pano Total typed 335 335 335 242 H. MOHRENWEISER ET AL.

The Dc~,variant has been reported in several 11 HI, 11 YZ and 15 QR-see fig. 2, Tanis et Amerindian tribes (Arends and Gallango, '64; al., '74 for village locations) located on the Tanis et al., '77). southwest edge of the Yanomama territory Private polymorphisms were screened for the presence of the CAI1 BAN-1 variant. No affected individuals were Previously undescribed variants of CAII identified. The Dc~,variant of Tf was also not and PEPB occurred in polymorphic frequency detected in the Yanomama. Thus the evidence in the Baniwa and Pano tribes, respectively. continues to accumulate for a relative paucity The CAII variant in the Baniwa was identified of intertribal migration. electrophoretically by a slightly slower migra- The second variant in polymorphic proportion than the normal CAII enzyme (CAII 1).It tions, in the Pano, was an electrophoretic vari- also stained less intensely than CAII 1 when ant of PEPB with a mobility slightly slower the fluorescein diacetate substrate was used than the previously reported PEPB 2. This be- (fig. 2c). The mobility and staining intensity havior was identical in the tris-maleate buffer were also different from the CAII 2 variant system of Lewis and Harris ('67) (fig. 2b) and which most commonly occurs in Negroes (fig. the tris-phosphate system of Harris and 2c). Although the variant enzyme had dif- Hopkinson ('76). The substrate specificity of ferent staining characteristics than normally the variant enzyme was identical to the nor- observed for CA it was inhibited by acetazol- mal B isoenzyme, i.e., the normal and variant amide (Diamox). This inhibitor is specific for enzyme stained with equal intensity when CA and has no effect on esterase staining Leu-Gly-Gly,Phe-Leu, Phe-Tyr and Leu-Leu- (Tashian and Carter, '76). Thus the possibility Leu were utilized as substrate. No bands mi- that the activity was due to a new esterase grating with the electrophoretic mobility of variant was excluded. The reduced staining normal or variant PEPB were noted when sub- intensity of the variant band relative to the strates specific for PEPA, PEPC or PEPD normal band was still observed after the he- were utilized. The electrophoretic pattern was molysate was extracted with CHCl ,-MEOH to unaffected when the sample was incubated remove the hemoglobin (procedure of Tashian in the presence of 5 mM dithiothreitol. No and Carter, '76). The CA in the CHU,-MEOH variation was noted in the electrophoretic extracted sample can be localized following mobility of PEPA, PEPC or PEPD in the af- starch gel electrophoresis with the protein fected individuals. An illustrative pedigree stain, nigrosin. The intensities of the two CAII is presented in figure 3a. The pattern is of bands (normal and variant) were similar after codominant inheritance of an allele we des- nigrosin staining, indicating similar amounts ignate PEPB 2PAN-'. The frequency of the of protein (fig. 2d). This would indicate that PEPB ZPAN-l allele in the Pano was 0.023 the specific activity of the CAII BAN-1 vari- although the distribution among the four ant is less than the specific activity of either groups was not uniform. The variant was not CAII 1or CAII 2. Additional characterization detected in the Cashinawa (40A,B) or in the of the variant enzyme continues. Pano (Katukina) located at Morada Nova The variant was detected in 39 of the 377 (41A). Only 2 of 48 individuals from the Jami- Baniwa; two of the 39 exhibited no CAII activ- nawa (35B) were affected; an allele frequency ity in the normal position and are presumed of 0.021. The allele frequency was 0.045 in the homozygotes. Thus the frequency of the Marubo (34A-E)and 0.048 in the Sete Estrelas CAZIBAN-lallele is 0.054. The pedigree of an il- Pano (35A). lustrative family is shown in figure 3b. As noted in the description of the tribes, in Rare variants view of the proximity of the Baniwa to the Three rare variants were encountered, in- Yanomama (cf. figs. 1 and 2 in Nee1 et al., '721, volving three different systems. The findings we were interested in the question of whether are given in table 4; a brief characterization of theALBYAN-'polymorphism would be encoun- each follows. tered in the Baniwa. It was not. The discovery of the CAII BAN-1 polymorphism in the Ba- HGB niwa obviously created a new opportunity to A suspected HGBA, variant was detected in seek evidence for genetic exchange between the Kanamari. On the original starch gel, the these two tribes. Accordingly, some 194 indi- staining intensity of the variant band was viduals from four Yanomama villages (11 G, equal to the staining intensity of the band in ELECTROPHORETIC VARIANTS IN AMERINDIAN TRIBES 243

Fig. 2 Electrophoretic pattern of several variants. a Transferrin: Wells 1 and 8 type C, wells 2 and 7 CDch, from Piaroa, wells 3 and 8 CDChi from Guaymi, wells 4 and 10 CDchi standard, wells 5, 9 and 11 samples from 3 different Baniwa individuals identified as CDch,, well 6 CDcua. b Peptidase B: Wells 1 and 8 type 1, wells 2, 4 and 6 type 1-PAN-1,wells 3 and 7 type 1-2, well 5 type 1-RAN-1. c Carbonic Anhydrase I1 - Activity stain: Wells 1 and 6 type 1-2, well 2 type BAN-1, wells 3 and 4 type 1, well 5 type 1-BAN-1. d Carbonic Anhydrase - protein stain: Well 1 CAII type 1-BAN-1, well 2 type 1, well 3 purified CAII type 1, well 4 purified CAI type 1. Samples in wells 1 and 2 were not purified, thus CAI and some residual hemoglobin are also present and stained. 244 H. MOHRENWETSER ET AL. the A, position; this latter in turn was approx- for the Baniwa (Gershowitz, personal com-

imately one half of normal. No alteration in munication), this variant is not felt~~~ to ~~ have- banding pattern or staining intensity was ob- been introduced and the allele has been desig- served in the HbA, region. from which we nated PEPB‘ BAN-1, infer that this is a 6-chain variant. This obser- vation could not be repeated. On cellulose ace- CAII tate electrophoresis, freshly prepared hemoly- A CAII variant which on direct comparison sates from packed cells stored in NLreproduci- exhibited an electrophoretic mobility identi- bly exhibited in addition to the normal HGBA, cal with the CAII 2 variant which occurs in band, a pair of abnormal hemoglobin bands polymorphic frequencies in Negroes was cathodal to the normal position of HGBA,; as detected in one individual in the Panoan the hemolysate ages these disappear and a Jaminawa. The ABO system reveals no evi- precipitate is seen at the origin. One of the dence of non-Indian admixture with the tribe, three children of the propositus regularly but the Gm groups suggest a 0.003 Negro com- exhibited similar findings with cellulose ace- ponent (Gershowitz and Neel, ’78).According- tate electrophoresis, but no abnormality has ly, this was not designated as a new variant. been seen with starch gel electrophoresis. The presence of an unstable variant of HGBA, is DISCUSSION suspected but clearly unproved. Unfortunate- The most noteworthy finding in this report ly, circumstances preclude obtaining the con- is the demonstration of two more electropho- firmation sample that a finding such as this retic “private polymorphisms” in Amerindian requires. tribes, involving PEPB in the Pano and CAII in the Baniwa. This brings to eight the num- PEPB ber of such polymorphisms encountered in a A PEPB variant was detected in a single in- study of 13 tribes with respect to an average of dividual in the Baniwa (fig. 2d). The variant 25 proteins (N.B.-the Kanamari were not in- had an electrophoretic mobility identical in cluded in the summary of Neel, ’78b). In addi- both the tris maleate system (Lewis and Har- tion, we encountered a single example of a var- ris, ’67) and tris/phosphate system (Harris iant PEPB in the Baniwa, and two possible and Hopkinson, ’76) with a variant seen in examples of an unstable variant of HGB ALin this laboratory in Caucasian populations, the Kanamari; in view of the lack of evidence which variant we assume to be the PEPB 2 of non-Indian admixture for these two tribes, variant described by Lewis and Harris (’67). we view the two variants as autochthonous. Because the ABO and Gm genetic systems The occurrence of this number of private reveal no evidence of non-Indian admixture polymorphisms in Amerindian tribes natural-

a) PEP6 2 PAN-1 b) CA II. BAN-1 ELECTROPHORETIC VARlANTS IN AMERINDIAN TRIBES 245 ly leads to speculation concerning their main- and Development Administration (now De- tenance by selective forces. Elsewhere we partment of Energy); by Grant NSF-DEB-76- have demonstrated that in a situation where 20591 from the National Science Foundation; there is a marked degree of tribal isolation, and by the Conselho Nacional de Desenvolvi- with the passage of time a high proportion of mento Cientifico e Tecnologico (Programa In- the neutral variants which are not lost by tegrado de Genetica). We thank the Instituto chance may be expected to assume polymor- Nacional des Pesquisas da Amaz6nia and the phic proportions (Thompson and Neel, '78). In Fundacao Nacional do Indio for the necessary the situation as currently defined for South clearances. We also acknowledge with ap- American Indians, the findings are consistent preciation the superb logistic support of the with neutrality of the mutant-bearing pheno- Research Vessel Alpha Helix of the National types and a mutation rate for electrophoretic Science Foundation. Other participanh in the variants of 0.7 X 10-5/locus/generation (Neel field work, whose assistance we gratefully and Thompson, '78). This estimate of the acknowledge, were Doctors Dale Lawrence, mutation rate may be biased downwards by Peter Smouse, Richard Spielman, W. J. Oliver, failure to detect all the private polymor- and James V. Neel, Jr., and Mrs. James V. phisms in the tribes under consideration. For Neel. instance, despite the fact that we have LITERATURE CITED sampled the Pano in four different locations, Arends, T., and M. L. Gallango 1964 Tranvferrins in the Western Pano are untouched, and our Venezuelan Indians: High frequency of a slow-moving sample is only 1.9%of the estimated 18,000 variant. Science, 143: 367-368. Galvao, E. 1959 Aculturaqio Indigena no Rio Negro. In: Pano. Be that as it may, the findings to date Boletin do Museu Paraense Emilio Goeldi, Nova Serie, can be explained by mutation pressure con- Antropologia, No. 7. Belem, Brazil. sistent with current estimates of mutation Gama Malcher, J. M. 1964 Indios. Conselho Nacional de rates in man, without the necessity to postu- Proteqao aos Indios. Publicaqao No. 1. Ministerio da Agricultura, Rio de Janeiro, p. 245. late the action of positive selection. Should Gershowitz, H., and J. V. Neel 1978 The immunoglobulin the variants on the average be to some extent allotypes (Gm and Inv) of twelve Indian tribes of Central deleterious, then a higher mutation rate is and South America. Am. J. Phys. Anthrop.. 49: 289-302. necessary to account for the existence of these Goldman, I. 1948 Tribes of Uaupes-Caqueta region. In: Handbook of South American Indians 3. Smithsonian In- polymorphisms. stitution, Washington, D.C.. pp. 763-798. The second noteworthy finding of this paper Greenberg, J. H. 1960 The general classification of Central is its supplementation of the previous evi- and South American languages. In: Men and Cultures, dence, based on the restricted distribution of Selected Papers of the Fifth International Congress of Anthropological and Ethnological Sciences, Wiladel- the private polymorphisms, for a relative lack phia, 1956. University of Pennsylvania Press, Philadel- of gene flow between the Indian tribes of the phia, pp. 791-794. Amazon and Orinoco basins. The available Harris, H., and D. A. Hopkinson 1976 Handbook of Enzyme evidence suggests that in pre-Columbian Electrophoresis in Human Genetics. North-Holland Pub- lishing Co., Amsterdam. times, the Yanomama and Baniwa appear to Johnston, F. E., R. L. Jantz, K. M. Kensinger, G. F. Walker, have been separated by the Arawak-speaking F. H. Allen, Jr. and M. E. Walker 1968 Red cell blood Bare, now reduced to a few remnant villages groups of the Peruvia Cashinahua. Hum. Biol., 40: on or near the Rio Negro (Loukotka, '68). This 508-516. Johnston, F. E., K. M. Kensinger, R. .L. Jantz and G. F. tribe has yet to be studied for variants of the Walker 1969 The population structure of the Peruvia systems surveyed in this paper. Pending such Cashinahua: Demographic, genetic and cultural inter- studies, which would greatly improve the relationships. Hum. Biol., 41: 29-41. ability to make detailed statements, we point Lewis, W. H. P., and H. Harris 1967 Human red cell pep- tidases. Nature, 215: 351-355. out that intertribal movements in that area Loukotka, C. 1968 Clawification of South American have not been sufficient to introduce the Indian languages. Ref. Series Vol. 7. University of Cali- CA IIBANor the TFDChlvariants of the Baniwa fornia Latin American Center, Los Angeles, 451 pp. into the Yanomama, nor theALBYANvariant Melatti, D. M., and J. C. Melatti 1975 Relatorio sobre 0s Indios Marubo. In: Skrie Antropologia Social, 13. Fun- of the Yanomama into the Baniwa. Thus the dacso Universidade de Brasilia, Brasil, p. 162. case is strengthened for regarding the tribe as Metraux, A. 1948 Tribes of the Jura-Purus Basins. In: the basic unit of Amerindian populations. Handbook of South American Indians. Vol. 3. Smithson- ian Institution, Bulletin 143. GPO. Washington, D.C., pp. ACKNOWLEDGMENTS 657-712. Neel, J. V. 1978a The population structure of an This work was supported by Contract EY- Amerindian tribe, the Yanomama. Ann. Rev. Genetics, in 76-C-02-2828 from the U.S. Energy Research press. 246 H. MOHRENWEISER ET AL.

__ 1978b Rare variants. private polymorphisms, 1972 Serum proteins, hemoglobins and erythrocyte en. and locus heterozygosity in Amerindian populations. Am. zymes of Brazilian Cayapo Indians. Hum. Biol.. 44: J. Hum. Genet., in press. 443-458. Neel, J. V., T. Arends, C. Brewer, N. Chagnon, H. Spielman, R. S.,E. C. Migliazza and J. V. Neel 1974 Re- Gershowitz. M.Layrisse, 2. Layrisse. J.MacCluer, E. Mi- gional linguistic and genetic differences among Yano- gliazza, W. Oliver, F. Salzano, R. Spielman, R. Ward and mama Indians. Science, 184: 637-644. L. Weitkamp 1972 Studies on the Yanomama Indians. Steward, J. H., and L. C. Faron 1959 Native Peoples of Proceedings, IV Int. Cong. Hum. Genet., Paris. 1971. Ex- South America. McGraw-Hill, New York, pp. 555-595. cerpta Medica, Amsterdam, pp. 96-111. Steward, J. H.. and A. Metraux 1948 Tribes of the Peruvian Neel, J. V., R. E. Ferrell and R. A. Conrad 1976 The fre- and Ecuadorian Montana. In: Handbook of South Ameri- quency of “rare” protein varients in Marshall Islanders can Indians. Vol. 3. Smithsonian Institution, Bulletin and other Micronesians. Am. J. Hum. Genet..28: 262-269. 143. GPO, Washington, D.C., pp. 535-656. Neel, J. V., R. J. Tanis, E. C. Migliazza, R. S. Spielman, Swadesh, M. 1955 Towards greater accuracy in lex- F. Salzano, W. J. Oliver. M. Morrow and S. Bacbofer iostatic dating. Int. J. Am. Linguistics, 21: 121-137. 1977 Genetic studies of the Macushi and Wapishana In- Tanis, R. J., R. E. Ferrell, J.V. Neel and M. Morrow 1974 dians. I. Rare genetic variants and a “private polymor- Alhumin Yanomama-2, a ‘private’ polymorphism of phism” of esterase A. Hum. Genet., 36: 81.107. serum albumin. Ann. Hum. Genet. (London),38: 179-190. Neel, J. V., and E. A. Thompson 1978 Founder effect and Tanis. R. J., J. V. Neel and R. T. deArauz 1977 Two more the number of private polymorphisms observed in “private” polymorphisms of Amerindian tribes: LDHB Amerindian tribes. Proc. Nat. Acad. Sci. (U.S.A.), 75: GUA 1 and ACPl BCUA1 in the Guaymi of Panama. Am. 1904-1908. J. Hum. Genet., 29: 419-430. Neel, J.V.. andK. M. Weiss 1975 The geneticstructureof a Tanis, R. J.. J. V. Neel, H. Dovey and M. Morrow 1973 The tribal population, the Yanomama Indians. XII. Biodem- genetic structure of a tribal population, the Yanomama ographic studies. Am. J. Phys. Anthrop., 42: 25-52. Indians. IX. Gene frequencies for 18 serum protein and Noble, G. K., Jr. 1962 Proto-Arawakan and its de- erythrocyte enzyme systems in the Yanomama and five scendants. In: IJAL 1963-1964, Part 11. Bloomington, neighboring tribes; nine new variants. Am. J. Hum. Indiana, Research Center in Anthropological Folklore Genet., 25: 655-676. and Linguistics. Tashian, R. E.. and N. D. Carter 1976 Biochemical genetics Oliveira, A. E., and E. Galvao 1973 A situaqao atual dos of carbonic anhydrase. Adv. Hum. Genet., 7: 1-56. Baniwa (Alto rio Negro)-1971. In: Museu Goeldi no Ano Thompson, E. A,, and J. V. Nee1 1978 The probability of do Sesquicentenario. Publicaqoes Avulsas, 20: 27-40. founder effect in a tribal population. Proc. Nat. Acad. Sci. Salzano, F. M., J.V. Neel. L. R.Weitkamp and J. P. Woodall (U.S.A.): 75: 1442-1445.