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Demography and genetics of the Sateré-Mawé and their bearing on the differentiation of the tupi tribes of South America
F. M. Salzano Tania A. Weimer Maria H. L. P. Franco Mara H. Hutz Departamento de Gendtica, instituto de Bioci&cias, Universidade Federal do Rio Grande do Sul, 90000 Porto Alegre, RS, Brazil
M. A. Mestriner A. L. Sim6es Departamento de Gendtica, Faculdade de Medicina, Universidade de Sdo Paulo, 14100 Ribeirdo Preto, SP, Brazil
Demography and Genetics of the Sater6-Maw6 and their Bearing on the Differentiation of the Tupi Tribes of South America Information concerning age and sex distribution, inter-racial marriages, migratory movements, fertility and mortality was obtained in one community of the Sater6-Maw6 Indians of the Brazilian Amazon. In addition, 29 genetic systems were studied from the blood and two from the saliva of 170 persons. A high level of exogamy was found, but low marital distances (average 56 km). The Sater&Maw6 are characterized by high frequencies ofL M~, Le, PGM I and GLO l, but low prevalences ofL Ns and ESD I. The data obtained in this group were compared with those of six other Tupi tribes, and the pattern of geographical variation of 18 alleles was analyzed. The population structure of the community studied suggested ample opportunity for genetic differentiation, and the genetic data confirmed this prediction. Some regularities in the distribution of several alleles were observed; in general the southern Tupi groups are more different from the Sater&Maw~ than the Amazonian ones.
Maria J. de Melo e Freitas Instituto de Ci&cias Bioldgicas, Universidade Federal do Amazonas, 69000 Manaus, AM, Brazil
Received 18January 1985 and accepted 21 February 1985 Keywords: Brazilian Indians,
migration patterns, polymorphisms, language and genetic diversification. Journal of Human Evolution (1985) 14, 647-655
1. I n t r o d u c t i o n W h e n the Portuguese discovered Brazil in 1500 they found, scattered t h r o u g h o u t the littoral areas, I n d i a n s s p e a k i n g l a n g u a g e s that were classified afterwards as belonging to one of the more w i d e s p r e a d South A m e r i c a n stocks: the T u p i - G u a r a n i . Pericot y G a r c i a (1962) asserts t h a t the nucleus of dispersion of people who spoke l a n g u a g e s of this stock should u n q u e s t i o n a b l y be p l a c e d between the P a r a g u a y a n d Paranfi rivers, from which, by the X i n g u river a n d other ways, they r e a c h e d the A m a z o n , the G u i a n a s a n d the A t l a n t i c coast (Figure l). Such a wide dispersion gives the o p p o r t u n i t y for a new a p p r o a c h to the p r o b l e m of the r e l a t i o n s h i p between l a n g u a g e a n d genetic diversity ( S p i e l m a n et al., 1974; Salzano el al., 1977; Black et al., 1983; Salzano et al., 1985). Therefore, when we recently s t u d i e d m e m b e r s of a T u p i tribe, the S a t e r & M a w ~ , we decided to evaluate the results o b t a i n e d trying to a n s w e r the following questions: (a) I n what w a y can the p r e s e n t p o p u l a t i o n s t r u c t u r e a n d d e m o g r a p h i c p a t t e r n of this tribe give clues a b o u t the factors that influenced the m i g r a t i o n of T u p i groups?; a n d (b) w h a t is the degree of genetic diversification p r e s e n t a m o n g T u p i - s p e a k i n g populations? C a n g r a d i e n t s or clines be distinguished? T h e results of this i n q u i r y are r e p o r t e d herein. 0047-2484/85/070647 + 09 $03.00/0
9 1985 Academic Press Inc. (London) Limited
648
F . M . SALZANO
ETAL.
E~~ri lon ai ~ pi \
~o.~.~Guarani Nucleus of
e~s,o~j~.~ ~ e r
] /
=G~oyoki
Figure 1. Map of South America showing the geographical location of the tribes considered, the probable nucleus of dispersion of the Tupi groups, and some of the rivers that provided the ways for their dispersion.
2. Subjects and Methods The Sater~-Maw~ (synonyms: Mau6, Mahu6, Mauh6, Maguaz, Magu6, Mau-au; Sater~ is the name of their main subdivision) have traditionally lived between the Tapajoz and Madeira rivers, close to the Amazon (Figure 1). They were found there by the Jesuits in the 17th century, and in that same region were partially grouped in missions, submitted to slavery or killed, depending on their attitude towards the church and the colonizers. But they never gave up, and despite three and half centuries of contact they still maintain today their cultural and biological identity. Their total population is presently estimated at 2500,
649
STUDIES ON SOUTH AMERICAN INDIANS
Table I
Demographic characteristics of the population studied
Age and sex distribution Age interval Group or characteristic
0-14
Males Females Total Percentage Sex ratio
100 100 200 54 100
15-30
31 and more
Age not determined*
Total
Estimated average age X + S.D.
44 53 97 26 83
38 38 76 20 100
166 99 265 -168
348 290 638 -120
19 + 16 19 _+ 18 19 + 17 ---
Interracial marriages Type of marriage Saterfi-Maw6 x Saterfi-Maw6 Sater6-Maw6 x Mestizo Sater6-Maw~ x Neo Brazilian Mestizo X Mestizo Mestizo x Neo Brazilian Total
Number
Percentage
49 8 2 2 1 62
79 13 3 3 2 --
Number
Percentage
4 5 47 56 44 56
7 9 84
Marriage and migratory movements Type of marriage Both partners born in Marau One partner from another population Both partners from outside Total number of marriages Number of populations involved Average marital distance (km)
Fertility and mortality Sample All families Completed families
Opportunity for selection+
No. femMes
No. live births
Mean + S.D.
Premature deaths (%) t
60 12
274 65
4.6 -+ 2"5 5.4 _+ 2.5
20
Im
If
If/ps
I
0"25
0"20
0"25
0'50
* Persons referred in the interviews as living in the village, but who were not seen. ~"Before age 15. ++Im = pd/ps where pd = premature deaths and ps = proportion surviving or 1 - p d . I f = Vf/Yc2 where V f = variance in offspring number in completed sibships and Yc= mean number of live births per woman who was past reproductive age. I = Im + If/ps = Index of Opportunity for Selection (Crow, 1958).
650
v.M.
SALZANO E T A L .
scattered over 33 communities (Pericot y Garcia, 1962; Malcher, 1964; Ribeiro, 1970; Ricardo el al., 1984). Ethnographic information about them can be found in Pereira (1954) and Leacock (1964). Details about their language are given by G r a h a m & G r a h a m (1978). Bellizzi (1957, 1958) has studied their dental health and anthropometric variation. Demographic information, blood and saliva samples for the present study were obtained in July 1982 at the Indian Post Marau, located at the margins of the Marau river. Access to the Post can only be made by water, and it took about seven hours to reach it from the town of Maufis. After collection the samples were kept under refrigeration and transported to M a u l s and Manaus. Blood group determinations and the preparation of aliquots were performed at the latter city, and subsequently the material was carried to Porto Alegre and RibeirS~o Preto, for further testing. The typing procedures have been described or referenced in Harris & Hopkinson (1976) and Salzano et al. (1980).
3. Results The demographic data arc summarized in Table 1. The estimated avcragc agc of thc population is low (19 years, independent of sex), with 54% of the individuals from whom age could be obtained falling in the range 0-14 years. The sex ratio shows some deviations that could be due to male emigration in the range 15-30 years and to preferential recalling of males outside the studied group. As expected from the history of the group, part of the marriages (21%) involved persons with acknowledged non-Indian ancestry. To avoid complications in the analysis of the genetic data, these individuals were not considered in the tabulation of the results. The migration data present a pattern that seems to be characteristic of Amazonian Indians: high levels of exogamy (only 7% of marriages in which both partners had been born in Marau), large number of places (44) involved in the exogamic unions, but relatively low marital distances (average: 56 km). Fertility can be regarded as moderate (five live-born children per woman who was past reproductive age), as well as mortality (20% of deaths before age 15). As a result, the Index of Opportunity for Selection also shows a moderate value (0'50). Table 2 presents phenotype and gene frequencies for 31 genetic systems studied among the Sater~-Maw~, and in Table 3 the results on 18 of the variable alleles are compared with those from six other Tupi tribes, as well as with South American Indians in general. The occurrence of two Kell positives, one Lu (a+), and six carriers of Gm3;5 or Gmt; 5 among the Sater6-Maw~ (Table 2) indicates that despite our efforts in trying to eliminate admixed individuals, some non-Indian genes did occur in the sample. This problem will be considered again in the Discussion section. The Saterfi-Mawfi present high frequencies of L Ms (0'76, range in South American Indians, RSAI: 0'19-0"84), Le (0-76, the highest found thus far in Indians of this continent), PGM~ (0-96; RSAI: 0"55-1"00), and G L O 1 (0'36; RSAI: 0"17-0.30); but low prevalences of L Ns (0'03; RSAI: 0"00-0'41) and E S D 1 (0"62; RSAI: 0"43-1"00). On the other hand, the Tupi tribes as a whole can be characterized as having low values o f R z and A C P A, and moderate frequencies of L Ms, Rl,,]k ~ and Gml,2; 21. Since the Sater~-Maw~ occupy a somewhat intermediate geographical position considering the other Tupi tribes tested (Figure 1), it could be asked whether significant clines or gradients of frequencies occur in the distribution of these 18 loci. First, it should be pointed out that if we examined the Sater~-Maw~ frequencies and those of the other Tupi tribes, there is no constant increase of the differences with geographical distance. But some
651
STUDIES ON SOUTH AMERICAN INDIANS Table 2
Phenotype and gene frequencies for 31 genetic systems studied among the Sater~-Maw~
System
No. tested
Phenotypes found
Numbers found
Gene or haplotype
Frequency
ABO MNSs
170 105
1.00 0"16 0"76 0.05 0"03
170
pt
0-55
Rh
170
R1 R2 Rz R0
0.61 0.24 0-12 0.03
Kell
170
k
0.99
Duffy
160
b~
0"67
Kidd
170
Jk ~
0"36
Lutheran
125
Lug
<0"01
Diego
170
Di n
O"14
ABH Secretion
165
Se
0'83
Le~ Secretion
164
Le
0"73
Hemoglobin
170
170 29 59 1 !2 3 1 136 34 27 60 t1 58 5 7 2 2 168 I43 17 100 70 1 124 45 125 160 5 152 12 170 170 79 91 170
I0 L Ms L Ms L Ns L N~
P
O MSs Ms MNS MNSs MNs NSs Pt P2 CDEe CDe CcDE CcDEe CcDe oDE cDEe K+ Ka+ aa+ aa+ aa+ aSec. Non-sec. Sec. Non-sec. A A~ B B A
Hb~ A Hb~a2
1"00 1"00
Gd~
1.00
PGDA
1"00
1-1 2-1 1-1 1-1 1-1 A AB B A 1-1 2 1 2-2 1-1 2-1 2-2 1-1 1- 1 1 1 1-1
156 14 170 170 170 4 40 126 168 60 89 20 26 69 74 169 168 t68 168
PGMI ~
0"96
PGM21 AK I ADAL
1"00 1.00 t'00
A CP A
O"14
ESAA~_~ ESD~
1"00 0"62
GLO l
0"36
CA~ l PEPA ~ PEPB l PEPC 1
1"00 1.00 t.00
Glucose-6-Phosphate Dehydrogenase Phosphogluconate dehydrogenase Phosphoglucomutasel
Males: 79 Females: 91 170
Phosphoglucomutas% Adenylate kinase Adenosine deaminase Acid phosphatase
170 170 170 170
170
Esterase A~ .~ Esterase D
168 169
Glyoxalase I
169
Carbonic Anhydrase 2 Peptidase A Peptidase B Peptidase C
169 168 168 168
1"00
652
F. M. SALZANO E T A L .
Table 2 continued No. tested
Phenotypes found
Numbers found
Gene or haplotype
Frequency
Haptoglobin
170
0"71
170 170 170 80
79 72 12 7 170 170 170 33 41 4 1 1
Hp I
Transferrin Ceruloplasmin Albumin Gm*
1-1 2-1 2-2 0 C B A 1;21 1,2;21 1;5,21 1,2;5,21 1,3;5,21
System
Tff
1"00
Cp B Al A
1.00 1.00
1;21 1,2;21 3;5 1;5
0"65 0-31 0.01 0-03
* D a t a reported by H a m e l et al. (1984).
Variability on 18 alleles among seven Brazilian Tupi tribes compared with that observed among South American Indians in general
Table 3
South A m e r i c a n Indians'~
T u p i tribes*
Allele L Ms L M~ L Ns L Ns pl R1 R2 Rz R~ Fy~ arka Di a Le PGM~ 1 ACP a ESD t Hp ~ Gm t,'2;21
No. of rare alleles++ No. of indiv. studied
SaterfiMaw~ 0"16 0'76 0"05 0"03 0"55 0"61 0"24 0"12 0"03 0"67 0-36 0.14 0"73 0.96 0.14 0-62 0"71 0.31
WaiS, pi 0"38 0'31 0"02 0"29 0"24 0"50 0"49 0"01 0"00 0"79 -0'08 -0.91 <0.01 1.00 0-51 0"33
ParakanS. E m e r i l l o n
Sirion6
Guayaki
Guarani
Average
Range
--
0"00-0'72 0"19-0'84 0'00-0'42 0.00-0.41 0'09-1 '00 0"09-0'96 0'03-0'73 0"00-0'33 0"00-0'23 0"24-1'00 0.11-1.00 0.00-0.44 0.34-0.63 0.55-1.00 0.01 0'37 0-43-1"00 0"20.0'83 0-00.0"53
0"19 0'81 0"00 0"00 0"68 0"71 0"08 0"00 0'16 1"00 0.55 0"39 -0'98 0.11 0'36 0'78 --
0"12 0"35 0-41 0"12 0"29 0'60 0'30 0'09 0"01 0'72 0"25 0.41 0'28 1.00 0.11 0.70 0'56 0-21
0'22 0"09 0'00 0"69 0"54 0'31 0"69 0"00 0"00 0'81 0"57 0.02 0.62 0-55 0.01 -0.34 0-16
0"13 0"59 0"00 0"28 1"00 0"61 0'29 0"00 0'10 1"00 0.63 0"00 1.00 --0.22 0.31
-0'50 0.22
0'22 0"50 0'06 0"22 0'45 0'57 0'34 0'05 0'02 0'69 0.40 0.12 0'51 0.85 0.11 0.73 0"62 0.25
--0"46 0"72 0"12 0"07 0"09 0-41 -0"23 ---
0
1
4
1
0
0
0
--
--
170
135
100
64
108
62
34
--
--
* D a t a reported by Fernet et al. (1964), Salzano (1964), Salzano & Steinberg (1965), Salzano & Sutton (1965), M a t s o n et al. (1966, 1968), Brown et al. (1974), Quiliei (1975), T c h e n et al. (1978, 1981), Black et al. (1980, 1983), H a m e l et al. (1984) and in the present c o m m u n i c a t i o n . "~ Samples w i t h at least 50 individuals. ++T h e rare alleles are the following (frequencies in parentheses): Waifipi: TJ Dchi (0'07). Parakangt: ry (0.05), PGM.2 I1 (0"01), CP ACAY 1 (0'07), TJ TM (<0"01). Emerillon: PGM26 (0.04).
S T U D I E S ON
SOUTH
AMERICAN
INDIANS
653
regularities can be discerned; always considering the Sater~-Maw6 as point of reference, they can be summarized as follows: (a) The frequencies ofR z and A C P a decline and those from f y a increase (with one exception) in all directions; (b) The Parakan~ sometimes present the opposite picture, when compared with the differences found at the north and south of the Saterfi-Mawfi. Thus, the Parakan~ prevalences of L Ms and H p 1 are higher and those o f L Ns and E S D 1 lower than those of the Sater~-Maw~, while the contrary is true for the other tribes; (c) Considering the differences as a whole, and disregarding the Guarani because there are no data for them in relation to many loci, it could be seen that there are less differences between the Sater6-Maw~ and the three other Amazonian tribes (Parakanfi, Emerillon, Wai~tpi) than between them and the two southern tribes (Guayaki and Sirion6).
4. D i s c u s s i o n
Before we discuss the questions posed in the introduction, it is important to consider the problem of the race admixture present among the Sater~-Maw6. As was mentioned in the previous section, alleles at the Kell, Lutheran and Gm systems indicate the occurrence of non-Indian genes among them. How significant are they? Fortunately, as the number of polymorphic loci amenable to study increases,' the possibility of correctly answering this question also increases. Fifteen of the systems investigated (ABO, Rh, Kell, Lutheran, Hemoglobin, G6PD, PGD, PGM2, AK, ADA, CA2, Hp, Tf, Cp and Gm) have specific markers that can detect non-Indian alleles if they were present. Considering them together, and using the method of Szathmary & Reed (1978) we estimate that the minimum and maximum amount of admixture present among the Sater~-Mawd are 1% and 5%, respectively. This value is in good agreement with the average calculated using the Gm markers only, which is of 4% (Hamel et al., 1984). Given the other factors that can influence gene frequency estimates (non-random samples, deterioration of reagents and/or blood samples, typing and transcription mistakes, for instance) we consider that this amount of admixture is not high enough to vitiate our comparisons. If the patterns of dispersion observed among the Sater~-Maw~ are at least an indication of how the Tupi groups moved in the past, they suggest that these migrations might have occurred at a slow pace. But a movement of 50 km per generation is compatible with the colonization of the French Guiana region, located some 2000 km from the postulated center of dispersion, in 800 years (taking 20 years as the period of one generation: 2000/50 = 40 • 20 = 800). Another question is the amount of tribal intermixture that might occur after these groups have settled at different places. Among the Sater~-Maw~ not a single instance of intertribal marriage has been observed, though the situation might be different among other Tupi communities. We conclude that the population structure of these groups provides ample opportunity for genetic differentiation. The genetic data reviewed here confirm this prediction. Although constant regular trends were not found (and would also be difficult to expect, given the conditions prevalent in our species), the southern Tupi groups differ more from the Sater~-Maw6 than the Amazonian ones; and despite the noted "average" pattern o f T u p i gene frequencies, wide departures can be observed among them. Thus, considering the 18 loci listed in Table 3, the mean of the differences between the extreme values of the Tupi tribes was as high as 45%.
654
F . M . SALZANO E T A L .
T h a n k s are d u e to t h e F u n d a ~ o N a c i o n a l do I n d i o ( F u n a i ) w h i c h g a v e p e r m i s s i o n to s t u d y the I n d i a n s a n d p r o v i d e d o t h e r assistance; a n d to G i r l e y V. S i m 6 e s a n d A l t a i r F. Santos, for h e l p in t h e field. F i n a n c i a l s u p p o r t was p r o v i d e d b y the C o n s e l h o N a c i o n a l de D e s e n v o l v i m e n t o Cient~fico e T e c h n o l d g i c o ( P r o g r a m a I n t e g r a d o de G e n ~ t i c a e T r d p i c o U m i d o ) , F u n d a ~ f i o d e A m p a r o ~ P e s q u i s a do E s t a d o do R i o G r a n d e do Sul, a n d special f u n d s f r o m the F e d e r a l U n i v e r s i t i e s o f R i o G r a n d e d o Sul a n d A m a z o n a s .
References Bellizzi, A. M. (t957). Sobre a incid~ncia de les6es gengivodent~.rias entre os indios Mau6s. Revista Brasileira de Odontologia 15, 22-27. Bellizzi, A. M. (1958). Pesquisas antropom6tricas nos indios Maw6, Karajfi e Kayapo. Free Docent Thesis, Escola de Medicina e Cirurgia, Rio de Janeiro. Black, F. L., Salzano, F. M., Berman, L. L., Gabbay, Y., Weimer, T. A., Franco, M. H. L. P. & Pandey, J. P. (1983). Failure of linguistic relationships to predict genetic distances between the Wai~tpi and other tribes of Lower Amazonia. American Journal of Physical Anthropology 60, 327-335. Black, F. L., Salzano, F. M., Layrisse, Z., Franco, M. H. L. P., Harris, N. S. & Weimer, T. A. (1980). Restriction and persistence of polymorphisms of HLA and other blood genetic traits in the Parakan~t Indians of Brazil. American Journat of Physical Anthropology 52, 119-132. Brown, S. M., Gajdusek, D. C., Leyshon, W. C., Steinberg, A. G., Brown, K. S. & Curtain, C. C. (1974). Genetic studies in Paraguay: blood group, red cell, and serum genetic patterns of the Guayaki and Ayore Indians, Mennonite settlers, and seven other Indian tribes of the Paraguayan Chaco. American Journal of Physical Anthropology 41, 3 t 7-343. Crow, J. F. (1958). Some possibilities for measuring selection intensities in man. Human Biology 30, 1-13. Fernet, P., Larrouy, G. & Rufti6, J. (1964). Etude h6motypologique des populations indiennes de Guyane Frangaise. II. Les groupes s6riques du syst~me Gm. Bulletins et Mgmoirs de la Societd d'Anthropologie de Paris 7, 119-123. Graham, A. & Graham, S. (1978). Assinalamento fonol6gico das unidades gramaticais em Sater6. Arquivos de Anatomia e Antropologia (Rio de Janeiro ) 3, 218-231. Hamel, M. L. H., Salzano, F. M. & Freitas, M.,I. M. (1984). The Gm polymorphism and racial admixture in six Amazonian populations. Journal of Human Evolution 13, 517-529. Harris, H. & Hopkinson, D. A. (1976). Handbook of Enzyme Electrophoresis in Human Genetics. Amsterdam: North-Holland. Leacock, S. (t964). Economic life of the Mau6 Indians. Boletim do Museu Paraense Emilio Goeldi, Antropologia 19, 1-30. Malcher, J. M. G. (1964). ]ndios, Grau de Integrar na Comunidade Nacional, Grupo Lingiilstico, Localizafdo. Rio de Janeiro: Conselho Nacional de Prote~fio aos Indios. Matson, G. A., Sutton, H. E., Swanson, J. & Robinson, A. (1968). Distribution of blood groups among Indians in South America. VI. In Paraguay. American Journal of Physical Anthropology 99, 81-98. Matson, G. A , Swanson, J. & Robinson, A. (1966). Distribution of hereditary blood groups among Indians in South America. 1II. In Bolivia. American Journal of Physical Anthropology 25, 13-33. Pericot y Garcia, L. (1962). Amgrica tnd~gena. Barcelona: Salvat. Pereira, N. (1954). Os Indios Mauds. Rio de Janeiro: Organizag~to Sim6es. Quilici, j. c. (1975). Structure hfimotypologique des populations indiennes cn Am6rique du Sud. Ph.D. Thesis, Toulouse. Ribeiro, D. (1970). Os ]ndios e a Civilizaf6o. Rio de Janeiro: Civiliza~o Brasileira. Ricardo, C. A., Gallois, D., Ricardo, F. P. & Carelti, V. (t984). Aconteceu. Povos Indlgenas no Brasil/83. Sho Paulo: Centro Ecum~nico de Documentagho e Inf0rma~o. Salzano, F. M. (1964). Blood groups of Indians from Santa Catarina, Brazil. American Journal of Physical Anthropology 22, 91-t06. Salzano, F. M., Callegari-Jacques, S. M., Franco, M. H. L, P., Hutz, M. H., Weimer, T. A., Silva, R. S. & da Rocha, F.J. (1980). The Caingang revisited: blood genetics and anthropometry. American Journal of Physical Anthropology 53, 513-524. Salzano, F. M., Gershowitz, H., Mohrenweiser, H., Neel, J. V., Smouse, P. E., Mestriner, M. A., Weimer, T. A., Franco, M. H. L. P., Sim6es, A. L., Constans, J., Oliveira, A. E. & Freitas, M . J . M . (1985). Gene flow across triba~ barriers and its effect among the Amazonian Igana River Indians (submitted for pubIicadon). Salzano, F. M., Neel, J. V., Gershowitz, H. & Migliazza, E. C. (1977). Intra and intertribal gernetic variation within a linguistic group: the Ge-speaking Indians of Brazil. American Journal of Physical Anthropology 47, 337-347.
STUDIES ON SOUTH AMERICAN INDIANS
655
Salzano, F. M. & Steinberg, A. G. (1965). The Gm and Inv groups of Indians from Santa Catarina, Brazil. American Journal of Human Genetics 17, 273-279. Salzano, F. M. & Sutton, H. E. (1965). Haptoglobin and transferrin types of Indians from Santa Catarina, Brazil. AmericanJournal of Human Genetics 17, 280-289. Spielman, R. S., Migliazza, E. C. & Neel, J. V. (1974). Regional linguistic and genetic differences among Yanomama Indians. Science 184, 637-644. Szathmary, E . J . E . & Reed, T. E. (1978). Calculation of the maximum amount of gene admixture in a hybrid population. AmericanJournal of Physical Anthropology48, 29-33. Tchen, P., Bois, E., Lanset, S. & Feingold, N. (1981). Blood group antigens in the Emerillon, Wayampi, and Wayana Amerindians of French Guiana. Human Heredity 31, 47-53. Tchen, P., Bois, E., S~ger, J., Grenand, P., Feingold, N. & Feingold, J. (1978). A genetic study of two French Guiana Amerindian populations. I Serum proteins and red cell enzymes. Human Genetics45, 305-315.
M. A. Mestriner A. L. Sim6es Departamento de Gendtica, Faculdade de Medicina, Universidade de Sdo Paulo, 14100 Ribeirdo Preto, SP, Brazil
Demography and Genetics of the Sater6-Maw6 and their Bearing on the Differentiation of the Tupi Tribes of South America Information concerning age and sex distribution, inter-racial marriages, migratory movements, fertility and mortality was obtained in one community of the Sater6-Maw6 Indians of the Brazilian Amazon. In addition, 29 genetic systems were studied from the blood and two from the saliva of 170 persons. A high level of exogamy was found, but low marital distances (average 56 km). The Sater&Maw6 are characterized by high frequencies ofL M~, Le, PGM I and GLO l, but low prevalences ofL Ns and ESD I. The data obtained in this group were compared with those of six other Tupi tribes, and the pattern of geographical variation of 18 alleles was analyzed. The population structure of the community studied suggested ample opportunity for genetic differentiation, and the genetic data confirmed this prediction. Some regularities in the distribution of several alleles were observed; in general the southern Tupi groups are more different from the Sater&Maw~ than the Amazonian ones.
Maria J. de Melo e Freitas Instituto de Ci&cias Bioldgicas, Universidade Federal do Amazonas, 69000 Manaus, AM, Brazil
Received 18January 1985 and accepted 21 February 1985 Keywords: Brazilian Indians,
migration patterns, polymorphisms, language and genetic diversification. Journal of Human Evolution (1985) 14, 647-655
1. I n t r o d u c t i o n W h e n the Portuguese discovered Brazil in 1500 they found, scattered t h r o u g h o u t the littoral areas, I n d i a n s s p e a k i n g l a n g u a g e s that were classified afterwards as belonging to one of the more w i d e s p r e a d South A m e r i c a n stocks: the T u p i - G u a r a n i . Pericot y G a r c i a (1962) asserts t h a t the nucleus of dispersion of people who spoke l a n g u a g e s of this stock should u n q u e s t i o n a b l y be p l a c e d between the P a r a g u a y a n d Paranfi rivers, from which, by the X i n g u river a n d other ways, they r e a c h e d the A m a z o n , the G u i a n a s a n d the A t l a n t i c coast (Figure l). Such a wide dispersion gives the o p p o r t u n i t y for a new a p p r o a c h to the p r o b l e m of the r e l a t i o n s h i p between l a n g u a g e a n d genetic diversity ( S p i e l m a n et al., 1974; Salzano el al., 1977; Black et al., 1983; Salzano et al., 1985). Therefore, when we recently s t u d i e d m e m b e r s of a T u p i tribe, the S a t e r & M a w ~ , we decided to evaluate the results o b t a i n e d trying to a n s w e r the following questions: (a) I n what w a y can the p r e s e n t p o p u l a t i o n s t r u c t u r e a n d d e m o g r a p h i c p a t t e r n of this tribe give clues a b o u t the factors that influenced the m i g r a t i o n of T u p i groups?; a n d (b) w h a t is the degree of genetic diversification p r e s e n t a m o n g T u p i - s p e a k i n g populations? C a n g r a d i e n t s or clines be distinguished? T h e results of this i n q u i r y are r e p o r t e d herein. 0047-2484/85/070647 + 09 $03.00/0
9 1985 Academic Press Inc. (London) Limited
648
F . M . SALZANO
ETAL.
E~~ri lon ai ~ pi \
~o.~.~Guarani Nucleus of
e~s,o~j~.~ ~ e r
] /
=G~oyoki
Figure 1. Map of South America showing the geographical location of the tribes considered, the probable nucleus of dispersion of the Tupi groups, and some of the rivers that provided the ways for their dispersion.
2. Subjects and Methods The Sater~-Maw~ (synonyms: Mau6, Mahu6, Mauh6, Maguaz, Magu6, Mau-au; Sater~ is the name of their main subdivision) have traditionally lived between the Tapajoz and Madeira rivers, close to the Amazon (Figure 1). They were found there by the Jesuits in the 17th century, and in that same region were partially grouped in missions, submitted to slavery or killed, depending on their attitude towards the church and the colonizers. But they never gave up, and despite three and half centuries of contact they still maintain today their cultural and biological identity. Their total population is presently estimated at 2500,
649
STUDIES ON SOUTH AMERICAN INDIANS
Table I
Demographic characteristics of the population studied
Age and sex distribution Age interval Group or characteristic
0-14
Males Females Total Percentage Sex ratio
100 100 200 54 100
15-30
31 and more
Age not determined*
Total
Estimated average age X + S.D.
44 53 97 26 83
38 38 76 20 100
166 99 265 -168
348 290 638 -120
19 + 16 19 _+ 18 19 + 17 ---
Interracial marriages Type of marriage Saterfi-Maw6 x Saterfi-Maw6 Sater6-Maw6 x Mestizo Sater6-Maw~ x Neo Brazilian Mestizo X Mestizo Mestizo x Neo Brazilian Total
Number
Percentage
49 8 2 2 1 62
79 13 3 3 2 --
Number
Percentage
4 5 47 56 44 56
7 9 84
Marriage and migratory movements Type of marriage Both partners born in Marau One partner from another population Both partners from outside Total number of marriages Number of populations involved Average marital distance (km)
Fertility and mortality Sample All families Completed families
Opportunity for selection+
No. femMes
No. live births
Mean + S.D.
Premature deaths (%) t
60 12
274 65
4.6 -+ 2"5 5.4 _+ 2.5
20
Im
If
If/ps
I
0"25
0"20
0"25
0'50
* Persons referred in the interviews as living in the village, but who were not seen. ~"Before age 15. ++Im = pd/ps where pd = premature deaths and ps = proportion surviving or 1 - p d . I f = Vf/Yc2 where V f = variance in offspring number in completed sibships and Yc= mean number of live births per woman who was past reproductive age. I = Im + If/ps = Index of Opportunity for Selection (Crow, 1958).
650
v.M.
SALZANO E T A L .
scattered over 33 communities (Pericot y Garcia, 1962; Malcher, 1964; Ribeiro, 1970; Ricardo el al., 1984). Ethnographic information about them can be found in Pereira (1954) and Leacock (1964). Details about their language are given by G r a h a m & G r a h a m (1978). Bellizzi (1957, 1958) has studied their dental health and anthropometric variation. Demographic information, blood and saliva samples for the present study were obtained in July 1982 at the Indian Post Marau, located at the margins of the Marau river. Access to the Post can only be made by water, and it took about seven hours to reach it from the town of Maufis. After collection the samples were kept under refrigeration and transported to M a u l s and Manaus. Blood group determinations and the preparation of aliquots were performed at the latter city, and subsequently the material was carried to Porto Alegre and RibeirS~o Preto, for further testing. The typing procedures have been described or referenced in Harris & Hopkinson (1976) and Salzano et al. (1980).
3. Results The demographic data arc summarized in Table 1. The estimated avcragc agc of thc population is low (19 years, independent of sex), with 54% of the individuals from whom age could be obtained falling in the range 0-14 years. The sex ratio shows some deviations that could be due to male emigration in the range 15-30 years and to preferential recalling of males outside the studied group. As expected from the history of the group, part of the marriages (21%) involved persons with acknowledged non-Indian ancestry. To avoid complications in the analysis of the genetic data, these individuals were not considered in the tabulation of the results. The migration data present a pattern that seems to be characteristic of Amazonian Indians: high levels of exogamy (only 7% of marriages in which both partners had been born in Marau), large number of places (44) involved in the exogamic unions, but relatively low marital distances (average: 56 km). Fertility can be regarded as moderate (five live-born children per woman who was past reproductive age), as well as mortality (20% of deaths before age 15). As a result, the Index of Opportunity for Selection also shows a moderate value (0'50). Table 2 presents phenotype and gene frequencies for 31 genetic systems studied among the Sater~-Maw~, and in Table 3 the results on 18 of the variable alleles are compared with those from six other Tupi tribes, as well as with South American Indians in general. The occurrence of two Kell positives, one Lu (a+), and six carriers of Gm3;5 or Gmt; 5 among the Sater6-Maw~ (Table 2) indicates that despite our efforts in trying to eliminate admixed individuals, some non-Indian genes did occur in the sample. This problem will be considered again in the Discussion section. The Saterfi-Mawfi present high frequencies of L Ms (0'76, range in South American Indians, RSAI: 0'19-0"84), Le (0-76, the highest found thus far in Indians of this continent), PGM~ (0-96; RSAI: 0"55-1"00), and G L O 1 (0'36; RSAI: 0"17-0.30); but low prevalences of L Ns (0'03; RSAI: 0"00-0'41) and E S D 1 (0"62; RSAI: 0"43-1"00). On the other hand, the Tupi tribes as a whole can be characterized as having low values o f R z and A C P A, and moderate frequencies of L Ms, Rl,,]k ~ and Gml,2; 21. Since the Sater~-Maw~ occupy a somewhat intermediate geographical position considering the other Tupi tribes tested (Figure 1), it could be asked whether significant clines or gradients of frequencies occur in the distribution of these 18 loci. First, it should be pointed out that if we examined the Sater~-Maw~ frequencies and those of the other Tupi tribes, there is no constant increase of the differences with geographical distance. But some
651
STUDIES ON SOUTH AMERICAN INDIANS Table 2
Phenotype and gene frequencies for 31 genetic systems studied among the Sater~-Maw~
System
No. tested
Phenotypes found
Numbers found
Gene or haplotype
Frequency
ABO MNSs
170 105
1.00 0"16 0"76 0.05 0"03
170
pt
0-55
Rh
170
R1 R2 Rz R0
0.61 0.24 0-12 0.03
Kell
170
k
0.99
Duffy
160
b~
0"67
Kidd
170
Jk ~
0"36
Lutheran
125
Lug
<0"01
Diego
170
Di n
O"14
ABH Secretion
165
Se
0'83
Le~ Secretion
164
Le
0"73
Hemoglobin
170
170 29 59 1 !2 3 1 136 34 27 60 t1 58 5 7 2 2 168 I43 17 100 70 1 124 45 125 160 5 152 12 170 170 79 91 170
I0 L Ms L Ms L Ns L N~
P
O MSs Ms MNS MNSs MNs NSs Pt P2 CDEe CDe CcDE CcDEe CcDe oDE cDEe K+ Ka+ aa+ aa+ aa+ aSec. Non-sec. Sec. Non-sec. A A~ B B A
Hb~ A Hb~a2
1"00 1"00
Gd~
1.00
PGDA
1"00
1-1 2-1 1-1 1-1 1-1 A AB B A 1-1 2 1 2-2 1-1 2-1 2-2 1-1 1- 1 1 1 1-1
156 14 170 170 170 4 40 126 168 60 89 20 26 69 74 169 168 t68 168
PGMI ~
0"96
PGM21 AK I ADAL
1"00 1.00 t'00
A CP A
O"14
ESAA~_~ ESD~
1"00 0"62
GLO l
0"36
CA~ l PEPA ~ PEPB l PEPC 1
1"00 1.00 t.00
Glucose-6-Phosphate Dehydrogenase Phosphogluconate dehydrogenase Phosphoglucomutasel
Males: 79 Females: 91 170
Phosphoglucomutas% Adenylate kinase Adenosine deaminase Acid phosphatase
170 170 170 170
170
Esterase A~ .~ Esterase D
168 169
Glyoxalase I
169
Carbonic Anhydrase 2 Peptidase A Peptidase B Peptidase C
169 168 168 168
1"00
652
F. M. SALZANO E T A L .
Table 2 continued No. tested
Phenotypes found
Numbers found
Gene or haplotype
Frequency
Haptoglobin
170
0"71
170 170 170 80
79 72 12 7 170 170 170 33 41 4 1 1
Hp I
Transferrin Ceruloplasmin Albumin Gm*
1-1 2-1 2-2 0 C B A 1;21 1,2;21 1;5,21 1,2;5,21 1,3;5,21
System
Tff
1"00
Cp B Al A
1.00 1.00
1;21 1,2;21 3;5 1;5
0"65 0-31 0.01 0-03
* D a t a reported by H a m e l et al. (1984).
Variability on 18 alleles among seven Brazilian Tupi tribes compared with that observed among South American Indians in general
Table 3
South A m e r i c a n Indians'~
T u p i tribes*
Allele L Ms L M~ L Ns L Ns pl R1 R2 Rz R~ Fy~ arka Di a Le PGM~ 1 ACP a ESD t Hp ~ Gm t,'2;21
No. of rare alleles++ No. of indiv. studied
SaterfiMaw~ 0"16 0'76 0"05 0"03 0"55 0"61 0"24 0"12 0"03 0"67 0-36 0.14 0"73 0.96 0.14 0-62 0"71 0.31
WaiS, pi 0"38 0'31 0"02 0"29 0"24 0"50 0"49 0"01 0"00 0"79 -0'08 -0.91 <0.01 1.00 0-51 0"33
ParakanS. E m e r i l l o n
Sirion6
Guayaki
Guarani
Average
Range
--
0"00-0'72 0"19-0'84 0'00-0'42 0.00-0.41 0'09-1 '00 0"09-0'96 0'03-0'73 0"00-0'33 0"00-0'23 0"24-1'00 0.11-1.00 0.00-0.44 0.34-0.63 0.55-1.00 0.01 0'37 0-43-1"00 0"20.0'83 0-00.0"53
0"19 0'81 0"00 0"00 0"68 0"71 0"08 0"00 0'16 1"00 0.55 0"39 -0'98 0.11 0'36 0'78 --
0"12 0"35 0-41 0"12 0"29 0'60 0'30 0'09 0"01 0'72 0"25 0.41 0'28 1.00 0.11 0.70 0'56 0-21
0'22 0"09 0'00 0"69 0"54 0'31 0"69 0"00 0"00 0'81 0"57 0.02 0.62 0-55 0.01 -0.34 0-16
0"13 0"59 0"00 0"28 1"00 0"61 0'29 0"00 0'10 1"00 0.63 0"00 1.00 --0.22 0.31
-0'50 0.22
0'22 0"50 0'06 0"22 0'45 0'57 0'34 0'05 0'02 0'69 0.40 0.12 0'51 0.85 0.11 0.73 0"62 0.25
--0"46 0"72 0"12 0"07 0"09 0-41 -0"23 ---
0
1
4
1
0
0
0
--
--
170
135
100
64
108
62
34
--
--
* D a t a reported by Fernet et al. (1964), Salzano (1964), Salzano & Steinberg (1965), Salzano & Sutton (1965), M a t s o n et al. (1966, 1968), Brown et al. (1974), Quiliei (1975), T c h e n et al. (1978, 1981), Black et al. (1980, 1983), H a m e l et al. (1984) and in the present c o m m u n i c a t i o n . "~ Samples w i t h at least 50 individuals. ++T h e rare alleles are the following (frequencies in parentheses): Waifipi: TJ Dchi (0'07). Parakangt: ry (0.05), PGM.2 I1 (0"01), CP ACAY 1 (0'07), TJ TM (<0"01). Emerillon: PGM26 (0.04).
S T U D I E S ON
SOUTH
AMERICAN
INDIANS
653
regularities can be discerned; always considering the Sater~-Maw6 as point of reference, they can be summarized as follows: (a) The frequencies ofR z and A C P a decline and those from f y a increase (with one exception) in all directions; (b) The Parakan~ sometimes present the opposite picture, when compared with the differences found at the north and south of the Saterfi-Mawfi. Thus, the Parakan~ prevalences of L Ms and H p 1 are higher and those o f L Ns and E S D 1 lower than those of the Sater~-Maw~, while the contrary is true for the other tribes; (c) Considering the differences as a whole, and disregarding the Guarani because there are no data for them in relation to many loci, it could be seen that there are less differences between the Sater6-Maw~ and the three other Amazonian tribes (Parakanfi, Emerillon, Wai~tpi) than between them and the two southern tribes (Guayaki and Sirion6).
4. D i s c u s s i o n
Before we discuss the questions posed in the introduction, it is important to consider the problem of the race admixture present among the Sater~-Maw6. As was mentioned in the previous section, alleles at the Kell, Lutheran and Gm systems indicate the occurrence of non-Indian genes among them. How significant are they? Fortunately, as the number of polymorphic loci amenable to study increases,' the possibility of correctly answering this question also increases. Fifteen of the systems investigated (ABO, Rh, Kell, Lutheran, Hemoglobin, G6PD, PGD, PGM2, AK, ADA, CA2, Hp, Tf, Cp and Gm) have specific markers that can detect non-Indian alleles if they were present. Considering them together, and using the method of Szathmary & Reed (1978) we estimate that the minimum and maximum amount of admixture present among the Sater~-Mawd are 1% and 5%, respectively. This value is in good agreement with the average calculated using the Gm markers only, which is of 4% (Hamel et al., 1984). Given the other factors that can influence gene frequency estimates (non-random samples, deterioration of reagents and/or blood samples, typing and transcription mistakes, for instance) we consider that this amount of admixture is not high enough to vitiate our comparisons. If the patterns of dispersion observed among the Sater~-Maw~ are at least an indication of how the Tupi groups moved in the past, they suggest that these migrations might have occurred at a slow pace. But a movement of 50 km per generation is compatible with the colonization of the French Guiana region, located some 2000 km from the postulated center of dispersion, in 800 years (taking 20 years as the period of one generation: 2000/50 = 40 • 20 = 800). Another question is the amount of tribal intermixture that might occur after these groups have settled at different places. Among the Sater~-Maw~ not a single instance of intertribal marriage has been observed, though the situation might be different among other Tupi communities. We conclude that the population structure of these groups provides ample opportunity for genetic differentiation. The genetic data reviewed here confirm this prediction. Although constant regular trends were not found (and would also be difficult to expect, given the conditions prevalent in our species), the southern Tupi groups differ more from the Sater~-Maw6 than the Amazonian ones; and despite the noted "average" pattern o f T u p i gene frequencies, wide departures can be observed among them. Thus, considering the 18 loci listed in Table 3, the mean of the differences between the extreme values of the Tupi tribes was as high as 45%.
654
F . M . SALZANO E T A L .
T h a n k s are d u e to t h e F u n d a ~ o N a c i o n a l do I n d i o ( F u n a i ) w h i c h g a v e p e r m i s s i o n to s t u d y the I n d i a n s a n d p r o v i d e d o t h e r assistance; a n d to G i r l e y V. S i m 6 e s a n d A l t a i r F. Santos, for h e l p in t h e field. F i n a n c i a l s u p p o r t was p r o v i d e d b y the C o n s e l h o N a c i o n a l de D e s e n v o l v i m e n t o Cient~fico e T e c h n o l d g i c o ( P r o g r a m a I n t e g r a d o de G e n ~ t i c a e T r d p i c o U m i d o ) , F u n d a ~ f i o d e A m p a r o ~ P e s q u i s a do E s t a d o do R i o G r a n d e do Sul, a n d special f u n d s f r o m the F e d e r a l U n i v e r s i t i e s o f R i o G r a n d e d o Sul a n d A m a z o n a s .
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STUDIES ON SOUTH AMERICAN INDIANS
655
Salzano, F. M. & Steinberg, A. G. (1965). The Gm and Inv groups of Indians from Santa Catarina, Brazil. American Journal of Human Genetics 17, 273-279. Salzano, F. M. & Sutton, H. E. (1965). Haptoglobin and transferrin types of Indians from Santa Catarina, Brazil. AmericanJournal of Human Genetics 17, 280-289. Spielman, R. S., Migliazza, E. C. & Neel, J. V. (1974). Regional linguistic and genetic differences among Yanomama Indians. Science 184, 637-644. Szathmary, E . J . E . & Reed, T. E. (1978). Calculation of the maximum amount of gene admixture in a hybrid population. AmericanJournal of Physical Anthropology48, 29-33. Tchen, P., Bois, E., Lanset, S. & Feingold, N. (1981). Blood group antigens in the Emerillon, Wayampi, and Wayana Amerindians of French Guiana. Human Heredity 31, 47-53. Tchen, P., Bois, E., S~ger, J., Grenand, P., Feingold, N. & Feingold, J. (1978). A genetic study of two French Guiana Amerindian populations. I Serum proteins and red cell enzymes. Human Genetics45, 305-315.