Distribution of hereditary blood groups among Indians in South America. VII. In Argentinaкод для вставкиСкачать
Distribution of Hereditary Blood Groups Among Indians in South America VII. I N ARGENTINA' G. ALBIN MATSON,::< H. ELDON SUTTON, JANE SWANSON ABNER ROBINSON Departments of Anthropology and Molecular and Genetic Biology, University of Utah, Salt Lake City; Department of Zoology, University of Texas, Austin; Minneapolis War Memorial Blood Bank, Minneapolis; and Child Research Ceizter of Michigan, DetTozt AND ABSTRACT This seventh and last paper in a series on the distribution of blood groups among Indians in South America reports the findings among Amerinds in Argentina. Blood specimens were procured from putative full-bloods of the following tribes: 38 Diaguita (Calchaqui), 230 Mataco, 90 Chiriguano, 142 Choroti, 51 Toba, 120 ChanQ, 96 Chulupi (Ashluslay), and 178 Araucano (Mapuche). These 945 samples were tested for blood factors in the A-B-0, M-N-S-s, P, Rh-Hr, K-k, Lewis, Duffy, Kidd, and Diego systems. Serum samples were tested for haptoglobins and transferrins. Hemolysates prepared from whole blood were tested for hemoglobin types. The results are presented in tables as phenotype distribution and calculated allele frequencies. Locations of the populations from which blood samples were procured are shown on a map of North and Central Argentina. High frequencies are reported for the 0 allele. Allele frequencies are high also for M , s, Ri (CDe), R2 (cDE), k , Ler* and Fy. They are usually low or absent for alleles B, N , S , Mia, V w , Ro (cDe), r (cde), K , Lei, and fy. The Di allele ranged from 0.013 in the Araucano (Mapuche) to 0.192 in the Toba. Allele frequencies aberrant for Indians were observed more often i n the Araucano (Mapuche) and Diaguita tribes, due probably to greater inflow of non-Indian genes into their gene pool and perhaps also to genetic drift i n small inbred populations. H p i allele frequencies varied from 0.43 i n the Choroti to 0.80 i n the Diaguita. All samples tested for transferrins except six contained the variant Tf C; the six were Bi C present i n samples from one Mataco and six Araucano persons. All the specimens tested electrophoretically for hemoglobin types contained only ( A ) as a major component. Argentina displays a variety of land, topography, and climate in its 1,080,000 square miles distributed through thirtythree and one-half degrees of latitude. In the north lies the near fallow Chaco which has assumed economic importance recently by the finding of oil. On the southern fringe of the Chaco's scrub forests lie plantations of oranges, yerba, tobacco, and sugar. To the west, stand the majestic snow-capped Andes. From the Andes eastward to Uruguay and the Atlantic, and southward to the end of the country lie high mountain lakes, pine forests, wind-swept ranches, the great fertile central plain, the pampas, great oil fields, and finally the bleak nearAntarctic latitudes consisting largely of rocky bays and islets. As varied as the land and its climate were the cultures of the people who lived AM. J. PHYS. ANTHROP., 30: 61-84. there when, in the sixteenth century, the Spaniards conquered and colonized it. Among their descendants are the presently existing Diaguita (Calchaqui), Mataco, Chiriguano, Choroti, Toba, Chanks, Chulupi (Ashluslay) and Araucano (Mapuche) tribes; populations which, by virtue of their donated blood, became the subjects of this study. Relevant information on each of these is briefly presented. The Diaguita (Calchaqui) In pre-Hispanic times the Diaguita (Calchaqui) inhabited arid Argentina and 1 This study is supported by the National Science Foundation grant NSF-G. 22923 and the National Institutes of Health, U. S. Public Health Service Research grants 1 R 0 1 GM. 14837-01, GM-09326 and 5-K3-GM-18.381 and the Universitv of Utah Research Fund. 4 Present address: Zoology Dept., Arizona State University, Tempe, Arizona 85281. 61 62 G. A. MATSON, H. E. SUTTON, J. SWANSON AND A. ROBINSON The Chiriguano The three little farming communities : Carapari, Yacuy and Tatima near Tartagal are inhabited almost exclusively by Indians of the Chiriguano tribe. Their language is of Tupi-Guarani stock. They cultivate maize, pumpkins, sweet potatoes, beans, sweet manioc, peanuts, tobacco and cotton. Roasted grains are pounded into flour from which mush is cooked. They are a proud people whose daring and ferocity have been commemorated in Spanish prose and song (Mktraux, '49). It was from 90 of these putatively fullblood Chiriguano Indians that blood specimens were procured. The Choroti The Choroti (Zolota), like the Mataco tribes subsist primarily through fishing, hunting and the gathering of algarroba pods and other wild products, and produce T h e Mataco Before Europeans arrived, nomadic and fire in the same manner. They speak the warlike tribes inhabited the region of the Matacoan language; are numbered among Argentine Chaco, Formosa and parts of the the foot Indians of the Chaco (Steward, present states of Santa Fe, Santiago del '49); and their dominant form of marriage Estero, and Salta. Among these people were is monogamy (Lowie, '49). According to Mktraux ('46) in 1915 the Mataco-Mataguayo, Choroti, Guayacuabout half of the then total 2,500 Choroti rue and Chiriguano (Levene, '37). lived on the Pilcomayo River near Fortin The Mataco proper (Mataguayo) have Guachalla. Most of the remainder ranged inhabited parts of the Gran Chaco for cen- north to Villa Montes in Bolivia, but a few turies. Among their first contacts with Cau- families roamed inland about 75 to 100 casians were the Jesuit missionaries. The miles from the river. It is probable that latter, however, received little encourage- some descendants of the latter migrants ment from the Mataco, who in 1635, in are the people from whom 91 blood samprevailing aboriginal taste, plotted to kill ples were procured at La Merced, as well them (Mktraux, '46). Today, as then, the as the itinerant laborers at the sugar facMataco dwellings are crudely constructed tory at Tabacal, Salta, from whom 51 samof branches set in the ground, joined over. ples were obtained. head and roughly thatched with rushes, The Toba grass, and palm leaves. Now, as then, the women weave textiles, and gather algarAmong the Guaycurues tribes of the roba pods, and the men make nets, fish Chaco the Toba Indians, because of their and hunt. Today they still light their fires stately stature and general good physical by rotary friction using a plain hand drill condition, have been held in greater esteem in a socketed hearth. Many migrate for than most of the others. They speak a employment on the sugar plantations of Guaicuran language and inhabit the Gran Salta and Jujuy; but on their own part they Chaco both in Argentina and Paraguay engage in agriculture to only a very limited (Mktraux, '46). Subsistence is gained priextent. marily through fishing, hunting, and some The 230 Mataco who contributed blood agriculture, by which they produce maize, for this study live in comparative isolation pumpkins, manioc, beans and tobacco. in the small settlements of San Luis, Santa The 51 blood samples of Toba blood were Maria and Puntana on the Itiyuro River in obtained from inhabitants of Carmedo and Salta's Chaco. Puntana not far from Tartagal. northern Chile. Their independent language, Kakan, differed basically from those of other tribes (Lothrop, '46). They lived in organized towns amid the mesquite, talas, canes and teasels (Bennett, '49). Their diet consisted primarily of maize, peas, gourds, the fruit of the chaiiar tree and prickly pears (Levene, '37). Today they subsist primarily by agriculture and animal husbandry. They make pottery and loom-woven fabrics and work in bronze, gold and silver. Villages are organized under politically autonomous headmen who are succeeded patrilineally from father to son. Polygyny was common and usually sororal (Lothrop, '46; Miranda, '46). The 38 Diaguita from whom blood specimens were procured for this study were employed at Finca Santa Rosa not far from Tartagal in the province of Salta. BLOOD GROUPS OF INDIANS I N ARGENTINA 63 limited; as citizens of the Republic of Argentina they participate in the country’s privileges and responsibilities. The Araucano have acquired more nonIndian genes than have any other Indians of this study with the possible exception of the Diaguita. Araucanian is spoken in the homes unless one of the parents is not Mapuche. All except a few old and isolated women speak Spanish. They subsist primarily by agriculture; cultivating maize, beans, squash, potatoes and small grains on land of private ownership. Such wealth as they possess consists of sheep, cattle, horses, hogs and chickens. The Chulupi (Ashluslay) Ox drawn wooden plows are used; sowing The Chulupi of Argentina and Paraguay and planting is done by hand. Grazing belong to the Matacoan linguistic stock and lands are very often held in common. To inhabit the plains north of the Pilcomayo date these Indians have little or no security River. They subsist by maize and manioc in land tenure and they have been deprived agriculture, fishing in low waters with of the better grazing grounds. The federal conical wicker baskets, hunting, and by government is presently attempting to deal gathering algarroba pods from which they with that problem. The 178 blood samples analyzed in this make flour for cakes. Chafiar fruits are study were procured essentially from farm boiled and the terminal shoots of palms are eaten raw, roasted, or boiled (Mktraux, laborers, their wives and children, residing in the community of Rucachoroy near San ’46). Martin de Los Andes, in the sate of NeuEvery winter, Chulupi bands migrate to work on sugarcane plantations in Argen- quen. MATERIALS AND METHODS ,tina, and it was from 96 of these itinerant Using a sterile Becton-Dickinson vaculaborers at Tabacal, Salta that blood specitainer tube-needle assembly, 5 to 8 mls of mens were obtained. blood were procured from volunteers by The Araucano (Mapuche) arm venipuncture, each tube containing In western Argentina near the Andes in 0.17 ml of citric acid-dextrose solution the State of Neuquen, reside parts of the (13.9 gm citric acid and 50 gm dextrose Araucanian nation, who still speak the dissolved in 1,000 ml distilled water) as Araucanian language. They are a resolute preservative. The clotted specimens, transand sturdy people, who, in pre-Hispanic ported under refrigeration via air express, times, enjoyed great freedom and cultural arrived in good condition at Minneapolis exchange with their kindred in Chile, and where the blood group tests were made. withstood Inca invasion for centuries The serum surrounding each clot was removed and frozen separately. Cells from (Cooper, ’46). Today Araucanian survivors in Argen- each sample were tested or blood group tina reside, for the most part, in Neuquen antigens by techniques recommended by Province and have a population of about antiserum processors. All reagent fluids 5,000, distributed among 26 ‘lofches” or used were specific and potent and had been small tribes. The lofche, consisting of 20 processed by the Minneapolis War Memoto 60 families, is under the control of a rial Blood Bank, gratuitously donated by cacique (chief). In the exercise of his other workers, or procured by purchase or authority the cacique is assisted by two barter. Erythrocytes were tested for blood counselors and receives advice from all the antigens and factors2 in the A-B-0, M-Nmen of the lofche. Clans of gens do not In this paper italics are used as symbols for genes exist in their society. In recent times the and genotypes and regular type as symbols for agglutinogens, phenotypes, blood factors (serological powers of the caciques have been greatly specificities) and the corresponding antibodies. The Chunks The Chanks as well as the Chiriguano have little or nothing in common with the Chaco tribes. Formerly they lived in the sub-Andean range of hills lying north and south of the Pilcomayo River. The C h a d speak Arawakan and differ in this respect from the Chiriguano whom they resemble culturally, except for minor details ( M B traux, ’46, ’48). Blood specimens were procured for 120 Chan6 residing in the two small agricultural communities of Yacuy and Tuijuuti near Tartagal. 64 G. A. MATSON, H. E. SUTTON, J. S W A N S O N AND A. ROBINSON Fig. 1 Map of Central and Northern Argentina. BLOOD GROUPS OF INDIANS I N ARGENTINA S-s, P, Rh-Wr, Kell-Cellano (K-k), Lewis (Le), Duffy (Fy"), Kidd (Jk"), and Diego 65 varez ('39) possessed, in addition, the B and AB blood group antigens. The 282 (Di") systems. Chulupi tested by Mazza ('39) and the 209 The frozen serum samples were flown Andidos examined by Paulotti and Alegria lo Austin, Texas where tests for hapto- ('43) yielded a small distribution of AB globin and transferrin types were made. but no B antigen. The serum protein components were sepaThe 0 allele has its maximum frequency rated by means of the vertical starch gel of 1.000 in the sample of several tribes method of Smithies ('59). The gel was which Mazza and Franke ('27) tested in then sliced into two parts, one of which the state of Salta, in the Chulupi tested by was stained with amidoschwarz 10-B to Mazza et al. ('33); and in the Mataco reveal proteins (Smithies, '55), and the tested by Mazza ('39). Its lowest frequency other with 0-dianisidine (Owen et al., '58) was reported among the Colla (0.797). to detect hemoglobin and haptoglobin- There is no correlation of the variation in hemoglobin complexes. All samples were allele frequencies with the degree of lintested with cyanmethemoglobin added, guistic relationship existent among the Traiisferrins were typed by a micropuri- tribes. The presence in South American fication method in use in the Austin labora- Indians of allele other than 0 in the A-B-0 tory. One drop serum is diluted with three system indicates their possession of nondrops of ferric ammonium citrate solution Indian genes. (0.15% in gel buffer: 0.03 M boric acid, The results of tests for the A-B-0 blood 0.006 M sodium hydroxide, pH ca. 8.4). groups made on samples procured for this To this mixture is added five drops of a study are presented in table 2 as number solution of rivanol (2-ethoxy-6, 9-diamino- and per cent of phenotypes with their coracridine lactate, 0.6% in gel buffer). A responding allele frequencies, which have precipitate forms and a clear supernatant been calculated by the method of Bernstein is obtained by low speed centrifugation for described by Stevens ('38). one minute. The supernatnat is subjected In the 945 samples procured from the to starch gel electrophorcsis (15 v/cm for eight tribes tested, group 0 is consistently 3-4 hours) and stained for protein by high. Indeed, the 51 Toba, 142 Choroti, conventional methods (Smithies, '59). and 230 Mataco presented the maximum Hemolysates, prepared by the method frequency of 1.000 for the 0 allele. This of Drabkin ('46), were frozen and then is not uncommon for isolated, putatively sent to the Child Research Center Labora- full-blood Indian populations in Middle and tories, Detroit, Michigan where they were South America (Matson and Swanson, tested for hemoglobin types by agar electro- '65a,b; Matson et al.,'66, '66a,b, '67). The phoresis at pH 6.5, citric acid-sodium citlowest frequency of allele 0 (0.858) is rate buffer (Robinson et al., '57) and by reported for the Diaguita, suggesting a paper electrophoresjs at pII 9.6, veronal larger heritage of non-Indian genes i n this buffer, ionic strength 0.06 (Zuelzer et al., population. '56). The remaining five populations shown in table 2 contain A' and/or B genes, but RESULTS AND DISCUSSION allele A2 appears only in the Chiriguano The A-B-0 sytsern and Diaguita tribes. The presence of alleles Earlier tests for antigens in the A-B-0 A', A2, or B in Indians of Middle or South system among Indians in Argentina have America suggests some mixture with nonbeen done by four workers. Results of these Indians at some time in the past. The data tests are shown in table 1 and will be shown in table 2 are in good agreement compared with results of this study shown with those shown in table 1 for Indians in Argentina and in general agreement with in table 2. Of the 11 populations listed in table 1, those reported for Amerinds i n Middle three yielded 100% group 0. The remain- and South America (Matson and Swanson, ing eight populations, though largely group '59, '61, '63a,b,c,d, '64, '65a,b; Matson, et 0, presented varying amounts of group al., '66, '67; Santiana, '47, '53; Arteaga, et A. Some of the 227 Mataco tested by Al- al., '52, Salazar-Mallen, '49; Sandoval and 2 1 (’33) Mazza e t al. (’33) 115 46 147 Mazza et al. ( ’331 Mazza et al. 209 194 Paulotti arid Alegria (’43) (’33) 227 135 78 282 55 120 Total numbers tested 73 38 131 198 191 179 130 78 269 55 120 No. 0 63.48 82.61 89.12 94.74 98.45 78.85 96.30 100.00 95.39 100.00 100.00 % Cited bv Salzano ( ’ 5 7 ) . *The locality in which this tribe was studied is not mentioned in the reference cited. Colla * Pillagas & Chaco Choroti Toba Prov. de Jujui Mazza et al. * Andidos Alvarez (‘39) * Matacos (’33) Mazza et al. Mazza (’39) * 2 * Mazza (’39) Prov. de Salta (’33) Mazza et al. Mazza and Franke (‘27) Prov. de Salta 6 2 Investigator Location Matacos Matacos Chunupi (Chulupi) Chunupi (Chulupi) Several Population TABLE 1 42 8 16 8 3 34 5 0 12 0 0 No. A 36.52 17.39 10.88 3.83 1.55 14.98 3.70 0.00 4.26 0.00 0.00 % 0 0 0 0 0 9 0 0 0 0 0 No. Phenotypes B 0.00 0.00 0.00 0.00 0.00 3.97 0.00 0.00 0.00 0.00 0.00 % 0 0 0 3 0 5 0 0 1 0 0 No. AB 0.00 0.00 0.00 1.43 0.00 2.20 0.00 0.00 0.35 0.00 0.00 % ~ r 0.797 0.909 0.944 0.966 0.992 0.879 0.981 1.000 0.975 1.000 0.203 0.091 0.056 0.027 0.008 0.090 0.000 0.000 0.000 0.007 0.000 0.031 0.000 0.000 0.000 0.019 0.002 0.000 0.000 4 0.023 0.000 0.000 P Allele frequencies 1.000 Distribution of the A - B - 0 blood groups in Indians of Arge’ntina-subgroups of “A” not shown (eurlier s t u d i e s ) Monte Carmelo and Puntana (Environs Tartagal) State of Salta Tabacal, State of Salta La Merced and Tabacal, Toba Chulupi Choroti 230 San Luis, Santa Maria and Puntana (Environs Tartagnl) State of Salta Finca Sta. Rosa - Chaco (Environs Tartagal) Province of Salta Mataco Calchaqui (Diaquita) 38 90 Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta Chiriguano 120 Tartagal, Yacuy and ruijuuti, State of Salta 142 96 51 178 Total numbers tested Chan6s State of Salta Rucachoroy (Environs San Martin de Los Andes), State of Neuquen Location Araucano (Mapuche) Population 28 230 81 118 142 95 51 154 No. 73.69 100.00 90.00 98.34 100.00 98.96 0.00 0.00 0.00 0.00 8.89 9 23.68 0 8 1 0.83 0 0 o 7.30 86.52 13 % NO. A1 % ioo.ao 0 0.00 1.11 0.00 0.00 0.00 0.00 0.00 % 1 2.63 0 1 0 0 0 o 0 No. A2 B 0.00 1.04 0.00 6.18 lo 0 0 0 0.00 0.00 0.00 1 0.83 0 1 0 '11 No. Phenotypes 0 0 0 0 0 0 0 0 lo 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 AiB 0 0 0 0 0 0 0 0 No. 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 % AYB "A" s h o w n ( t h i s s t u d y ) No. of Argentina-subgroups of TABLE 2 Distribution of the A-B-0 blood groups in I n d i a n s 0.858 1.000 0.948 0.992 1.000 0.995 0.127 0.000 0.046 0.004 0.015 0.000 0.003 0.000 0.000 O.CO3 0.000 0.030 0.034 0.003 0.005 0.000 0.000 0.000 0.000 0.032 4 0.000 0,OCO PZ 0.c00 0.037 PI Allele frequencies 1.000 0.931 T 68 G . A. MATSON, H. E. SUTTON, Henckel, '54; Salzano, '57; Suarez et al., '64, '65; Nagel and Etcheverry, '63). M-N-S-s s y s t e m The number and per cent of M-N and S-s phenotypes and their corresponding allele frequencies are assembled in table 3. The only other tests reported for M-N types i n Argentina Indians is that of Mazza ('39) in 282 Chunupi (Chulupi) in the Chaco. NIazza's results showed 81.91% M, 11.35% MN and 6.74% N. The allele frequency for M was 0.876 and for N 0.124. Frequencies for the MS, M s , NS, N s chromosomes in this present study were derived from the phenotypic data, in part by gene counting and in part by a n extension of the maximum likelihood method described by Boyd ('54). In all tribes except the Diaguita the allele frequency is high for M and low for N . There is a n equitable distribution of the M and N frequencies for the Diaguita (M = 0.500). The frequency for M in the Araucano agrees well with that reported for the same tribe in Chile (Matson et al., '67), and those in the Chulupi and Toba agree rather well with frequencies reported for the same tribes in Paraguay (Matson et al., '67b). These high frequencies for the M allele resemble those reported for putatively pure Amerinds elsewhere (Arteaga et al., '52; Wiener et al., '45; Boyd, '39; Brown et al., '58; Chown and Lewis, '53; Levine et al., '35; Matson and Piper, '47; Matson and Roberts, '49; Matson and Schrader, '33; Matson and Swanson, '59, '61, '63a,b,c,d, '64, '65a,b; Matson et al., '36. '54, '63, '66, '66a,b, '67). The lower frequency for allele M in the Diaguita may be accounted for, perhaps, on the basis of a greater flow of non-Indian genes into this population than among the other tribes tested and/or by genetic drift. High M and low N is usually associated with racial purity in Amerinds. The highest frequency for the s allele and lowest for S is recorded for the Diaguita ( s =- 0.789). The lowest s allele frequency is reported for the Choroti (0.549). It is usual to find high frequencies for s and low for S alleles in American Indians. Frequencies for the four chromosomes M S , M s , N S and Ns vary somewhat with the tribe examined. The M S chromosome J. SWANSON AND A. ROBINSON frequency is higher and the Ms lower in the Choroti than ordinarily expected in South American Indians and in the Diaguita the N s chromosome is higher than usual. Since the M and s allele frequencies are usually high and the N and S allele frequencies low in Amerinds, the expected distribution would be high M s and very low N S , with M S and N s frequencies appearing somewhere between. Departures from this general distribution usually observed in full-blood Amerinds may suggest inflow of non-Indian genes or possibly genetic drift in small inbred populations. Either, or both, mechanism may be suspected by the higher than usual frequency for N in the Diaguita and the higher than usual S frequency in the Choroti. Miltenberger (Mi")and V e r w e y s t (Vw)antigens Of the 945 blood samples tested for the Miltenberger (Mi") and Verweyst (Vw) antigens only one of each was observed. The 230 Mataco specimens yielded both antigens in the same person. Negative tests are quite usual in American Indians, Miltenberger (MP) having been reported previously i n only one tribe, i.e. in four of 239 Quechua in the northern plateau in Ecuador (Matson et al., '66); and Verweyst (Vw) in only one person among 162 Kekchi at CobAn, Guatemala (Matson and Swanson, '61). The alleles corresponding to these antigens are extremely rare not only in American Indians but in Europeans as well, Mi" having been reported in them in a proportion of about 1 in 500, one half of which are coupled with Vw and one half with Vw - (Hart et al., '54). + P system Specimens from the Argentine Indians were tested only with anti-P (anti?,) serum for antigens in the P system. The distribution of phenotypes of the P system in these populations and the calculated frequencies of their corresponding alleles are presented in table 4. The maximum allele frequency of 1.000 or the PI gene in the Diaguita, and its low frequency of 0.289 in the Araucano (Map3,The notations PI and P? (Sanger, '55; Matson et al., 59) are employed, PI being the antigen previously called P and Pz the one formerly called p. 4.78 7 11 90 (Diaguita) C alchaqui Mataco 38 230 7.78 8 120 Chan6s Chiriguano 21 142 Choroti 0 10 96 Chulupi 0.00 6.67 14.79 10.42 5.88 3 51 0.56 % Toba s 1 No. 178 Araucano (Mapuche) Population Total numbers tested TABLE 3 4 48 20 30 35 22 6 35 No. c/b 25.28 96 6 15.79 16.09 37 20.87 10.53 18.89 17 22.22 21.67 9.16 13 26 23.96 27.45 S 23 14 45 No. 25.00 24.65 22.92 11.77 19.66 ss M 6.25 1.96 5.06 % 10.53 1.74 4 4 7.78 6.67 4.93 S 7 8 7 6 1 9 No. % 2 49 19 20 5.26 21.30 21.11 16.66 23.94 15.62 15 34 17.65 16.85 SS 9 30 NO. MN Phenotypes 12 45 13 21 23 16 12 37 No. $6 31.58 19.57 14.44 17.50 16.20 16.67 23.53 20.79 S 0.00 0 0.00 1 0.43 0 2 4 5 5.26 1.74 5.56 8 31 2 6 1 0.83 0.00 0 7 1 0.70 5 1 0.70 0 0.00 8 No. 4 0.00 % 6.74 ss 0.00 0 0 1 1.96 % IVO. 12 s 1 0.56 No. N 2.22 5.00 4.93 4.16 9.80 4.50 96 21.05 13.48 S 0.121 0.252 0.279 0.278 0.402 0.302 0.182 0.379 0.378 0.427 0.460 0.309 0.464 0.485 0.509 Ms 0.090 0.037 0.121 0.068 0.049 0.057 0.063 0.118 NS 0.410 0.333 0.173 0.194 0.240 0.177 0.270 0.213 Ns Chromosome frequencies 0.160 MS Distribution of phenotypes and chromosome frequencies in the M-N-S-s system in Indians o f Argentina 70 G. A. MATSON, H. E. SUTTON, J. SWANSON AND A. ROBINSON TABLE 4 Distribution of antigens and allele frequencies in the P system i n Indians of Argentina samples tested with anti-PI serum only Phenotypes Population Location Total numbers tested P1-C - . % Araucano (Mapuche ) Rucachoroy (Environs San Martin de Los Andes), State of Neuquen 178 88 49.44 Toba Monte Carmelo and Puntana (Environs Tartagal) State of Salta 51 47 92.16 Chulupi Tabacal, State of Salta 96 76 79.17 Choroti Tabacal and La Merced, State of Salta 142 114 80.28 Chants Tartagal, Yacuy and Tuijuuti, State of Salta 120 106 Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta 90 Mataco San Luis, Santa Maria and Puntana (Environs Tartagal) State of Salta 230 Calchaaui (Diaguita) Finca Sta. Rosa - Chaco (Environs Tartagal) Province of Salta 38 1 The - Allele frequencies _. PI Nn NO. Chiriguano P, PZ % 90 50.56 0.289 0.711 4 7.84 0.720 0.280 20 20.83 0.544 0.456 28 19.72 0.556 0.444 88.33 14 11.67 0.658 0.342 80 88.89 10 11.11 0.667 0.333 195 84.78 35 15.22 0.610 0.390 38 100.00 0 0.00 1.000 0.000 notations PI and Pa (Sanger, '55; Matson et al., '59) are used where PI = former P and PZ= former p. uche) are noteworthy variants in opposite directions. The frequencies for PI presented in table 4 for the remaining tribes range between 0.544 and 0.720. These fall within the range usually reported for this allele in American Indians (Chown and Lewis, '53; Matson et al., '54; Matson and Swanson, '59, '61, '63a,b,c,d). No p or Pk was noted in the Argentine Indians tested. It is noteworthy that the low PI allele frequency (0.289) in the Araucano at Rucachoroy differs so widely from that determined for the Araucano on the western slope of the Andes at Longuimay, Chile, where a population of 141 present a frequency of 0.831. The reasons for these deviations in opposite directions for P' allele frequency in two isolates of the same tribe are not immediately apparent. Until more information is at hand concerning the history and migrations of this tribe and their relationships with other populations, east and west of the Andes, perhaps it can only be assumed that any or all of such factors as isolation, inbreeding, genetic drift and, more recently, miscegenation with Caucasians may have played a role. Prevalence of the maximum frequency of 1.000 for allele P' in the 38 Diaguita is also remarkable, but no less so than the same observation in 51 Guayaki of Paraguay (Matson et al., '67b). Similar high frequencies have been reported also for this allele in other small, isolated, inbred populations of putatively pure Indians in South America e.g., in 14 ChAcoba in the Bolivian Beni (Matson et al., '66b), 14 Isconahua and 9 Yagua in Peru (Matson et al., '66a). In such small populations the chances of introducing bias is great; however, very high frequencies for PI have been observed also in 109 Itonama in the Bolivian Beni; 93 Ticuna, 93 AymarB, 151 Aguaruna, and 181 Quechua in Peru (Matson et al., '66a). BLOOD GROUPS O F INDIANS IN ARGENTINA Rh-Hr s y s t e m Five antisera were employed in testing the blood samples for antigens in the RhHr system, i.e., anti-rh' (C), -Rho ( D ) , -rh" (E), -hr' (c) and -hr" (e) *. Anti-hr" (antie ) was used only to determine the zygosity status of the rh" ( E ) factor. Results of the tests with the above antisera are presented as number and per cent of phenotypes in table 5 and as calculated allele frequencies in table 6. In only two populations, the Araucano and Chanks, did the rh (ccddee) phenotype appear and only in the Chanks and Diaguita was the Rho (ccDee) phenotype present, These two phenotypes are rare indeed or completely absent among fullblood Indians and are thought to be determined by genes of non-Indian origin. The Rhlrh phenotype ranged from 0.00% in the Mataco to the high values of 18.54% in the Araucano and 10.53% in the Diaguita. It is noteworthy that the phenotype kh,Rhz, usually high in full-blood Amerinds, had lower values in the Araucano and Diaguita than in the other tribes shown in the table, suggesting again the greater presence of non-Indian genes or perhaps gentic drift in these two tribes. Phenotype data can be observed better when reduced to allele frequencies. These have been calculated from the data in table 5 and presented in table 6. Absent in all the tribes were alleles T' ( C d e ) , r" ( c d E ) and rY ( C d E ) . The allele r ( c d e ) appeared in the Araucano, ChanBs, and Diaguita, but it was absent in the other populations. The R" ( c D e ) allele was present in all except the Araucano, but it presented its highest frequency in the Diaguita. Alleles R1 ( C D e ) and R2 ( c D E ) were present in all tribes, but the frequency for Rz ( c D E ) , usually high in full-blood Indians, was highest in the Mataco (0.715) and lowest in the Araucano (0.289). Allele R" ( C D E ) , ordinarily present in Amerinds, was absent in the Araucano but present in all the other tribes. In Amerinds the R' ( C D e ) and R2 ( c D E ) allele frequencies are higher and the T ( c d e ) and R" ( c D e ) lower than in other ethnic groups with the possible exception of Polynesians (Matson et al., '54; Matson and Swanson, '59, '61, '63b,c, '64, '65a; Chown and Lewis, '53; 71 Matson and Piper, '47; Pantin and Kallsen, '53; and Brown et al., '58). Kell-Cellano s y s t e m Anti-K and anti-k sera were both employed in testing the 945 Argentine Indian blood specimens for Kell and Cellano antigens. All samples tested negative for Kell (K) and positive for Cellano ( k ) as is usually the case in putatively pure Indian populations; for homozygous K has not been noted in Amerinds and heterozygous (Kk) only in a few instances (Chown and Lewis, '57; Matson and Swanson, '59, '63b; Fernandes et al., '57; Salzano, '61; Best et al., '62; Allen, '59; Ellis et al., '63; Layrisse and Wilbert, '60; Nunez Montiel et al., '57; Etcheverry et al., '66, '66a,b). Only Pantin and Junqueira ('52) have reported any appreciable amount of Kell in Amerinds (20% or more in Indians of Mato Grosso, Brazil). Presence of Kell in American Indians strongly suggests a nonIndian source for the responsible gene. Lewis s y s t e m Both anti-Lel and anti-Lea sera were used in testing for antigens in the Lewis system. The results of these tests are presented in table 7. Only two populations in this series presented phenotype Lel i.e., the Araucano, in which there were two among 178 persons and the Choroti, having four in 142. The complete absence of Let in the remaining tribes conforms to the pattern odserved in full-blood Amerind populations generally (Matson and Swanson, '59, '61, '63b,d, '64, '65a,b; Matson et al., '66b, '67; Chown and Lewis, '53; Gray and Laughlin, '60; Layrisse et al., '61, '62, '63a). The presence of the Lel phenotype in the Araucanian and Choroti populations may be a further indication, therefore, of non-Indian genes in their gene pools. Tests were not made on saliva to determine the prevalence of the A-B-H secretor character in these Indians. The few studies that have been accomplished on the distribution of the A-B-H secretor and its cor4The existence of more than one nomenclature for this system makes it desirable to follow the Rh-Hr notations with CDE equivalents in arentheses. 5 Lewis notations recommended f y Wiener ('65) are used i n this paper where: Lel =Le(a+b-), Lez = (a-b+), le = (a-b-); anti-Lei = anti-Lea and anti-Lea = anti-Leb. 140 119 90 230 Tabacal, State of Salta Tabacal and La Merced, State of Salta Tartagal, Yacuy and Tuijuuti, State of Salta Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta San Luis, Santa Maria and Putana (Environs Tartagal) State of Salta Chulupi Choroti ChanCs Chiriguano Mataco 38 % No. % CcDee Rhlrh No. % CCDee RhlRhl % CcDEe No. % % 0 0.00 1 2.63 0 0.00 0 0.00 0 0.00 0 0.00 1 0.84 2 1.68 0 0.00 0 0.00 0 0.00 0 0.00 0 0.00 0 0.00 5 9.80 30 58.83 1.43 33 23.57 58 41.43 1.04 19 19.79 38 39.58 3.96 1.11 18 20.00 42 46.67 4 10.53 6 15.79 12 31.58 0 0.00 12 5.22 93 40.43 1 4 3.36 27 22.69 48 40.34 2 1 1 5.88 9 17.65 % 2 3.92 1 1.96 7 18.42 1.31 113 49.13 4 10.53 3 4 4.44 13 14.44 8 6.72 22 18.49 3 2.14 35 25.00 2 5.26 2 5.26 2 0.87 7 3.04 6 6.67 6 6.67 5 4.20 2 1.68 7 5.00 1 0.71 2 2.08 22 22.92 10 10.42 3 3.13 3 No. &DEE Rh,Rha 0 0.00 0 0.00 % CCDEe No. ccDEE No. ccDEe No. Rh.Rhl RhzRhz Rhzrh Phenotypes RhlRhz 10 5.62 0 0.00 33 18.54 45 25.84 45 25.28 30 16.85 14 7.87 % ccDee No. ccddee No. Rh, rh ‘ % 0 0.00 0 0.00 0 0.00 0 0.00 1 0.71 1 1.04 0 0.00 0 0.00 No. CCDEE RhzRhz Sera 1 None of the following phenotypes were observed: rh’rh (Ccddee); rh’rh’ (CCddee); rh’rh” (CcddEe); rh”rh (ccddEe); rh”rh” (ccddEE); rh,rh‘ (CCddEe); rh,rh” (CcddEE); rhyrhy (CCddEE). Finca Sta. Rosa - Chaco (EnvironsTartagal) Province of Salta 96 Monte Carmelo and Puntana (Environs Tartagal) State of Salta Toba Calchaqui (Diaguita) 51 Rucachoroy (Environs San Martin de 10s Andcs), State of Neuquen Araucano 178 Location Population Total numbers tested Distribution of Rh-Hr blood groups in Indians of Argentina samples tested with anti-Rho (D), anti-rh’ (C), anti-rh“ ( E ) , anti-hr‘ ( c ) and anti-hr” ( e ) TABLE 5 73 BLOOD GROUPS OF INDIANS I N ARGENTINA TABLE 6 Allele frequencies f w Rh-Hr blood groups calculated f r o m data o n table 5 Population Araucano (Mapuche) Toba Chulupi Total numbers tested Allele frequencies 7 Re R1 Rz Re cde cDe CDe cDE CDE Rucachory (Environs San Martin de Los Andes), State of Neuquen 178 0.233 0.000 0.478 0.289 0.000 Monte Carmelo and Puntana (Environs Tartagal) State of Salta 51 0.000 0.041 0.420 0.508 0.031 Tabacal and La Merced, State of Salta 96 0.000 0.017 0.452 0.452 0.079 Location Choroti Tabacal, State of Salta 140 0.000 0.012 0.471 0.486 0.031 Chanks Tartagal, Yacuy and Tuijuuti, State of Salta 119 0.045 0.033 0.464 0.426 0.032 90 0.000 0.030 0.470 0.431 0.069 San Luis, Santa Maria and Puntana (Environs Tartagal) State of Salta 230 0.000 0.007 0.258 0.715 0.020 Finca Sta. Rosa - Chaco (Tartagal Environs) Province of Salta 0.001 0.139 0.387 0.413 0.060 Chiriguano Mataco Calchaqui (Diaguita) Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta 38 responding Se gene in Indians of South America is discussed in paper number VI of this series (Matson et al., '67b). Generally, in studies of A-B-H secretors among American Indian populations in which frequencies for the Se gene are less than 1.000, the deviation has been attributed to the presence of non-Indian genes in the Indian gene pool (Matson and Piper, '47; Chown and Lewis, '53; Pollitzer et al., '62; Ellis et al., '63; Salzano, '64). In Europeans, the distribution of A-B-H secretors is low and Lel secretors high (Race and Sanger, '62), whereas in Amerinds the reverse has been found. Therefore, as expected, in American Indians having European admixture the Lei phenotype becomes higher and A-B-H secretors lower than in €ull-blood Amerinds. D u f f y System Duffy blood group factors were determined in the Argentine Indian samples by the use of anti-Fy" serum only. The phenotype distribution and allele frequencies determined from the results of the tests are presented in table 8. Phenotype Fy( a ) is consistently high for the populations represented, ranging from 91.11% in the 90 Chiriguano to 100% in the 51 Toba and 142 Choroti. In contrast, tests performed on samples from 100 Toba at Laguna Por6 in the Paraguayan Chaco yielded only 83.00% positive for the Fy" antigen, which was the lowest value found for Fy" in Paraguayan Indians. The reason for this inconsistency in the frequency of the Fy gene in the Toba in separate locations in the Gran Chaco is not known, one is constrained to think that basic principles such as migration, isolation, inbreeding, miscegenation and/or genetic drift have played their parts. The certainty of what happened must remain, for the present at least, a close-locked secret of their distant past. More similar, however, were the values for the prevalence of the Fy(a +) phenotype in the Chulupi of Argentina and Paraguay i.e., 96.88% (allele frequency 0.823) + TABLE 7 2.82 4 96 142 120 90 230 38 Tabacal and La Merced, State of Salta Tabacal, State of Salta Tartagal, Yacuy and 'Tuijuuti, State of Salta Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta San Luis, Santa Maria and Puntana (Environs Tartagal) State of Salta Finca Sta. Rosa - Chaco (Environs Tartagal) Province of Salta Chulupi Choroti Chanbs Chiriguano Mataco 0 0 0 0 0 32 203 81 109 122 80 42 162 No. % 6 27 9 11 le = Le(a-b-), 15.79 11.74 10.00 9.17 0.397 0.343 0.316 0.303 0.336 0.408 0.420 0.281 Ze anti-Lei = anti-Lea 0.603 0.657 0.684 0.697 0.664 11.27 16 0.580 0.719 0.592 17.65 7.87 I Le Allele frequencies 16.67 le 16 9 14 NO. = Le(a-b+), 84.21 88.26 90.00 90.83 85.91 83.33 82.35 91.01 Ley Leg Phenotypes = Le(a+b-1, 0.00 0.00 0.00 0.00 0.00 1 Lewis notations recommended by Wiener ('65) are used in this paper where: Lei and anti-LeH = anti-Lea. Calchaqui (Diaguita) 0.00 0 51 Monte Carmelo and Puntana (Environs Tartagal) State of Salta 1.12 Toba 2 178 % Rucachoroy (Environs San Martin de Los Andes), State of Neuquen Araucano (Mapuche) No. Lei Location Population Total numbers tested Distribution of phenotypes and allele frequencies for the Lewis system in Indians of Argentina samples tested with anti-Lel and anti-Len sera TABLE 8 51 96 142 120 90 230 38 Monte Carmelo and Puntana (Environs Tartagal) State of Salta Tabacal, State of Salta Tabacal and La Merced, State of Salta Tartagal, Yacuy and Tuijuuti, State of Salta Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta San Luis, Santa Maria and Puntana (Environs Tartagal) State of Salta Finca Sta. Rosa - Chaco (Environs Tartagal) Province of Salta Toba Chulupi Choroti Chan6s Chiriguano Mataco Calchaqui (Diaguita) 178 Rucachoroy (Environs San Martin de Los Andes), State of Neuquen Araucano (~Mapuche) --".-- Location Population Total 36 224 82 115 94.74 97.39 91.11 95.83 100.00 96.88 93 142 100.00 93.26 % 51 166 No. - sampbs 2 6 8 5 0 3 0 12 No. Phenotypes Distribution of phenotypes and allele frequencies f o r the D u f f y system in Indians o f Argentina 5.26 2.61 8.89 4.17 0.771 0.838 0.702 0.796 1.000 0.823 3.12 0.00 1 .ooo 0.740 0.00 6.74 % Allele freai 0.229 0.162 0.298 0.204 0.000 0.177 0.000 0.260 tested w i t h anti-Fp serum only 76 G. A. MATSON, H. E. SUTTON, and 98.65% (allele frequency 0.847) respectively. Although the Fy( a+ ) phenotype presents an erratic distribution in American Indians, ranging from 100% positive in the Kutchin (Lewis et al., '61), the Yahgan (Matson et al., '67), the Guayaki (Matson et al., '67b) and in the Toba and Choroti of this study, to 100% negative in one group of Brazilian Indians (Pantin and Junqueira, '52), the values are more frequently high than low (Matson et al., '54; Pantin and Kallsen, '53; Matson and Swanson, '59, '61, '63a,b,c,d, '64, '65a,b; Matson et al., '66, '66a,b, '67; Chown and Lewis, '53; Hulse, '57; Brown et al., '58). In Caucasians moderately low frequencies for the Fy allele have been reported (Cutbush and Mollison, '50; Matson et al., '54). Kidd s y s t e m Only anti-Jk" serum was available for testing the Argentine Indian specimens for the homologous antigen in the Kidd system. Phenotype and allele frequency data presented in table 9 were determined €rom the results of these tests. A wide variation exists in the distribution of the Jk(a +) phenotype and the frequencies of its responsible allele among Indian tribes in Argentina. The range as shown in table 9 is from a low of 46.07% (allele frequency 0.266) in the Araucano to a high of 88.24% (allele frequency 0.657) in the Toba. The range is practically the same as among Indians in Paraguay (Matson et al., '67b) but the Toba of Paraguay presented a frequency of only 0.490 for the J h allele. The frequencies for J h in the Chulupi in the Chaco of both countries varied less than in the Toba i.e., 0.490 in Argentina and 0.368 in Paraguay (Matson et al., '67b). Elsewhere in Middle and South America frequencies for the Jh allele resemble those appearing in table 9. Low frequencies for Jk have been noted among the Araucano (Mapuche) of Chile i.e., (0.113) (Matson et al., '67), the AymarA and Chama in Bolivia (frequencies 0.389 and 0.270 respectively, Matson et al., '66b) and in others e.g., the Aguaruna in Peru, Quechua in Ecuador, Guaymi in Panama, T6rraba in Costa Rica, Mam in Guatemala (Matson J. SWANSON AND A. ROBINSON et al., '66, '66a; Matson and Swanson, '65b, '61). Frequencies of 0.500 or more for J h are numerous however in Indians of the Americas (Chown and Lewis, '53; Allen, '59; Matson and Swanson, '59, '61, '63a,b, c,d, '64, '65a,b; Matson et al., '66, '66a,b; Tejada et al., '61; Pollitzer et al., '62; Layrisse et al., '61, '62, '63, '63a; Best et al., '62 and Gershowitz et al., '67). Although frequencies vary greatly for the Jk allele in Amerinds, they present no easily discernable ethnological pattern at the present time, and they are not statistically very different from those reported for the English (Race et al., 51) or for American Whites (Allen et al., '51; Rosenfield et al., '53). Chinese in New York have been reported, however, to have a low frequency for the Jk allele (Rosenfield et al., '53). Diego s y s t e m A potent and specific anti-Di" serum obtained from Dr. Layrisse of Caracas, Venezuela was used to test the Argentine Indian bloods for the Diego (Di") antigen. The results of these tests are presented in table 10 as number and per cent of phenotypes and calculated allele frequencies. The allele responsible for the Di" antigen was present in all the tribes tested, but the distribution of the phenotype varied greatly between tribes ranging from 2.60% Di( a +) (allele frequency 0.013) in the Araucano to 34.74% Di(a +) (allele frequency 0.192) in the Chulupi. The D i allele frequencies in the Toba and Chulupi of this study were in fair agreement with those reported for the same tribes in the Chaco in Paraguay (Matson et al., '6%). So also was there agreement between the low values determined for Di in the Araucan0 (Mapuche) in Argentina and in Chile, Distribution of phenotype Di( a +) and frequencies of its corresponding allele varies widely among American Indians (Layrisse and Wilbert, '60), but no orderly ethnographic gradient or cline appears. Layrisse and Wilbert ('60) have attempted to explain this situation on the basis of migration waves; an early one is postulated to have had a low frequency of the Di allele, followed by a later wave of high D i frequency. They endeavor to correlate given physical and serological types with specific cultural and sociological traits. TABLE 9 88.24 73.96 45 71 96 142 120 90 230 38 Tabacal, State of Salta Tabacal and La Merced, State of Salta Tartagal, Yacuy and Tuijuuti, State of Salta Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta San Luis, Santa Maria and Puntana (Environs Tartagal) State of Salta Finca Sta. Rosa - Chaco (Environs Tartagal) Province of Salta Chulupi Choroti Chanbs Chiriguano Mataco Calchaqui (Diaguita) 29 118 53 90 111 76.32 51.30 58.89 75.00 78.17 46.07 51 82 Monte Carmelo and Puntana (Environs Tartagal) State of Salta 178 Toba % Rucachoroy (Environs San Martin de Los Andes), State of Neuquen Araucano (Mapuche) No. Location JW a+ 1 9 112 37 30 31 25 6 96 No. JWa-1 23.68 48.70 41.11 25.00 21.83 26.04 11.76 53.93 % - samples tested Phenotypes Population Total numbers tested Distribution of phenotypes and allele frequencies for the Kidd system i n Indians of Argentina 0.513 0.302 0.358 0.500 0.533 0.490 0.266 Jk 0.487 0.698 0.642 0.500 0.467 0.510 0.734 ik Allele frequencies with anti-Jka serum only r $ 8 0 + 2 3 * 0 80 td 230 28 Monte Carmelo and Puntana (Environs Tartagal) State of Salta Tabacal, State of Salta Tabacal and La Merced, State of Salta Tartagal, Yacuy and Tuijuuti, State of Salta Carapari, Tatima, Yacuy (Environs Tartagal) State of Salta S a n Luis, Santa Maria and Puntana (Environs Tartagal) State of Salta Finca Sta. Rosa - Chaco (Environs Tartagal) Province of Salta Toba Chulupi Choroti Chanes Chiriguano Mataco 81 118 142 95 51 154 Rucachoroy (Environs San Martin de Los Andes), State of Neuquen Araucano (Mapuche) Total numbers tested Location Population Calchaqui (Diaguita) TABLE 10 4 69 7 19 37 33 12 4 No. 2.60 % 14.29 30.00 8.64 16.10 26.06 34.74 24 161 74 99 105 62 39 150 No. Phenotypes 23.53 Wa+) =(a-) 85.71 70.00 91.36 83.90 73.94 65.26 76.47 97.40 % Distribution of phenotypes and allele frequencies f o r the Diego (Dia) antigen i n Indians of Argentina 0.074 0.163 0.045 0.084 0.140 0.192 0.125 0.013 Di 0.926 0.837 0.955 0.916 0.860 0.808 0.875 0.987 di Allele frequencies 2 H v) ? tl 5. m F % 0 .1 cn 0 ? 79 BLOOD GROUPS OF INDIANS I N ARGENTINA TABLE 11 Distribution of haptoglobin types in Indians of Argentina Population Location Araucano (Mapuche) Rucachoroy (Environs §an Martin de Los Andes), State of Neuquen Toba Monte Carmelo and Puntana (Environs Tartagal) State of Salta Total Phenotypes 0 Allele frequency Hpl 11 0 0.79 23 14 0 0.49 0.44 numbers tested 1-1 207 132 64 50 13 2-1 2-2 Chulupi Tabacal, State of Salta 100 23 42 35 0 Choroti Tabacal, and La Merced, State of Salta 142 24 75 43 0 0.43 Tartagal, Yacuy and Tuijuuti, State of Salta 120 26 76 18 0 0.53 Chiriguano Carapari, Tatirna, Yacuy (Environs Tartagal) State of Salta 109 29 52 25 3 0.52 Mataco San Luis, Santa Maria and Puntana (Environs Tartagal) State of Salta 205 47 103 54 1 0.48 Finca Sta. Rosa - Chaco (Environs Tartagal) Province of Salta 38 24 11 2 1 0.80 Chanes Calchaqui (Diaguita) Nee1 and Salzano ('66) have criticized this typological approach proposed by Layrisse and Wilbert ('60). (0.78), Mapuche (0.72) in Chile, (Matson et al., '67; Nagel and Etcheverry, '63); the Aymara (0.69) in Peru (Matson et al., '66a; Giblett and Best, '61); the Quechua Haptoglobins (0.78) and Colorado (0.89) in Ecuador Haptoglobins differentiated by starch-gel Matson et al., '66); the Caingang (0.79) electrophoresis (Smithies, '55) are pre- and (0.73) in Brazil (Salzano and Sutton, sented in table 11 as the four different '63, '65); and the Guahibo (0.67), Piraroa phenotypes and calculated frequencies of (0.77) and Waica (0.80) in Venezuela the autosomal Hp' allele. (Arends and Gallango, '62, '64). The usual three phenotypes Hp 1-1, Hp It appears that the Hp' allele frequencies 2-1, and Hp 2-2 were the only ones demon- in the Indians of this study range from strable, except in the Chiriguano, Mataco, 0.44 or the Chulupi to 0.80 in the Diaguita. and Diaguita, in which populations were The higher frequencies for Hp' in the revealed very low prevalences of Hp 0 Araucano and Diaguita are in line with (apparent ahaptoglobinemia), the inheri- their deviations from the other Argentine tance. of which is not well understood. The Indians with respect to blood group allele distribution of the three phenotypes Hp frequencies. However, these Hp' allele fre1-1, Hp 2-1, and Hp 2-2 forms a roughly quencies cannot be explained on the basis uniform 1 , 2, 1 pattern throughout the of inflow of non-Indian genes. At least, no tribes except in the Araucano and Diaguita one has reported a population with Hp' tribes in which the prevalence of Hp 1-1 frequencies near 1.0 except in very small phenotype far exceeds the other two. This isolates. Genetic drift may explain some relationship is clearly depicted in the Hpf aberrant allele frequencies in the Diaguita allele frequency column. Other high fre- and Araucano better than significant forquencies reported for the Hp' allele in eign admixture. Apparently the Araucano South America are: The Atacameiios and Diaguita differ from the other Indians (0.67), Araucano (0.75), Pehuenches tested in the distribution of many genes. 80 G . A. MATSON, H. E. SUTTON, J. SWANSON AND A. ROBINSON The Hp' allele frequencies shown in the All these contained hemoglobin (A) as a table for the remaining tribes are in es- major component. No unusual major hemosential agreement with values reported globin component was encountered. generally for Indians in Middle and South Among Indians in Middle and South American Indians e.g., the Alacaluf (0.48) America abnormal hemoglobin types have in Chile (Matson et al., '67); the San Blas been rarely found, though the following Cuna (0.38) and Choc6 (0.46) in Panama exception to hemoglobin (A) have been (Matson et al., '65b); the Terraba (0.52), noted: in southern Mexico, two hemoglobin Cabkcar (0.44), and Boruca (0.44) in (N), one (AS) and one (F) were detected Costa Rica (Matson et al., '65a); the Pan- in 87 Chiapaneca (Sutton et al., '60; Matare (0.35), Pamon (0.38), Paraujuano son et al., '63); in Costa Rica, two (AS) (0.53) in Venezuela (Arends and Gallango, '62); the Xavante (0.46) in Brazil were found in 67 Thrraba; in Panama five (Neel et al., '64); and the Aweikoma and (AS) and one homozygous ( S ) in 153 Guarani (0.52) in Brazil (Salzano and Guaymi, and two (AS) in 350 San Blas (Matson and Swanson, '65b). These few Sutton, '63). aberrancies may well have been determined by genes introduced into the gene pool Transferrins Serum samples tested for haptoglobins from non-Indians. were subjected to electrophoretic mobility ACKNOWLEDGMENTS tests to determine the presence of variants This study could not have been accomof the iron-binding pl globulin, transferrin. Of the more than a dozen variants that plished without the interest and cooperahave been defined (Giblett, '62; Parker tion of numerous persons and agencies. and Bearn, '61), only two were detected Genuine gratitude moves us to mention in the serum specimens from the Argen- the following: tine Indians. All the samples except six In the United States. The Department contained the variant Tf C; the six were of State and the U. S. Customs Service asBIC, present in samples from one Mataco sisted by official sanction and in the expeand five Araucano individuals. ditious movement of blood samples through With few exceptions South American portals of entry into the United States. The Indian tribes tested for transferrin variants, technical assistance of Mrs. Annette Arosahave presented 100% values for Tf C (Mat- mena, Minneapolis War Memorial Blood son et al., '66, '66a,b; Salzano and Sutton, Bank, in performing blood grouping tests is '65; Giblett and Best, '61; Neel et al., '64; much appreciated. Parker and Bearn, '61). The exceptions, I n Argentina. The following persons asbesides the BIC variant, reported in this sisted in official and non-official capacities paper haven been very low frequencies of in the procurement and handling of blood slow Tf DI (or Dchi) in the Piro in Peru samples from the Indians: (Matson et al., '66a), the Quechua, Cayapa In Buenos Aires. Dr. Arturo Oiiatavia, and Secoya in Ecuador (Matson et al., ' 6 6 ) , El Ministro de Asistencia Social y Salud and in the Paraujuano and Yupa in Vene- Publica; Dr. Frederick W. Brown, Scientific zuela (Arends and Gallango, '62, '64) and Attach&,American Embassy; Mr. Robert A. the B2C variant in the Quechua in Peru Hancock, Counselor of Embassy, American (Giblett and Best, '61) and the Caingang Embassy; Dr. Bernard0 A. Houssay, Presiin Brazil (Salzano and Sutton, '63). dente, Podor Ejecutivo Nacional, Consejo Nacional de Investigaciones Cientificas y Hemoglobin types Technicas; Professor Enrique Palavecino, Hemolysates were prepared from a total Director, Museo Etnografico de la Faculof 353 samples of whole blood. The tribes tad de Filasofia y Letras, Universidad de represented and the number of samples Buenos Aires; Dr. Armando S. Parodi, Catested from each were the following : Arau- tearc de Microbiologia, Faculdad de Medcan0 (Mapuche) 39, Toba 24, Chulupi 37, kina, Universidad de Buenos Aires; Mr. Choroti 92, Chan6s 54, Chiriguano 55, Richard Baxendale, American Embassy; Mataco 26, and Diaguita (Calchaqui) 26. Sr. Jose B. Gaon, Gerente de Cargas, Pan BLOOD GROUPS OF INDIANS IN ARGENTINA American-Panagra Air Lines; Sra. Maricela Gil Pietranera de Pombo, Florida, 947. In Salta, Province of Salta. Dr. Oscar Rivero; Dr. Danton Cermesoni, Sr. Ministro de Asuntos Sociales y Salud Publica de Salta; Dr. Nestor Rodriguez Campoamor; Padre Lozano; Senorita Nelly Perkz; Senorita Sonia D’Andrea; Victor Amable Guerra, Hito 1”” Sargento, Jefe Hito 1, Salta. In Tartagal, Province of Salta. Padre Jose Buttinelli, Parroco de Tartagal; Rev. Francis Tomkins; Sr. Jefe de Seccidn, Gendarmeria Nacional; Sr. Humberto Aguero, Jefe Servicio Lucha Antipa-Barrio Mitre; Sr. Julio Andre Ferreyra, Recopilador del Folklore del Chaco Salteno; Sr. Juan Albert, Driver; Sr. Arnerico Gonzalez, Cook; Juan Teodoro Gueros, Sargento l“”,Santa Maria; Jefe Seccidn Santa Victorio “Este” de Gendarmeria Nacional; Rev. and Mrs. Olof Jonsson, Missionaries to the Mataco. In San Carlos de Bariloche, Province of Rio Negro. Jose M. Iglesias, M.D. “Ruca Lelen”; Federico Caeiro, M.D.; Emilio Feliu, M.D. LITERATURE CITED Allen, F. H.,Jr. 1959 Summary of blood group phenotypes in some aboriginal Americans. Am. J. Phys. Anthrop., 17: 86. Allen, F. H., Jr., L. K. Diamond and B. Niedziela 1951 A new blood group antigen. Nature, 167: 482. Alvarez, A. G. 1939 Comprobaciones bioldgicas en aborigenes argentinos - Consideraciones sobre los grupos sanguineos de los Matacos. Comision Honoritria de Ruduciones de Indios. Buenos Aires. Publ. no. 6: 25-34. Arends, T., and M. L. Gallango 1962 Haptoglobin and transferrin groups in Venezuela. Proc. 8th Cong. Internat. SOC.Blood Transf., Pp. 379-382. 1964 Frequencies of haptoglobin types in various South American populations. Proc. 9th Cong., Internat. SOC.Blood Transf., 463471. Arteaga, C., M. Salazar-Mallh, E. L. Ugalde and A. Velez-Aronzco 1952 Blood agglutinogms of the Mexicans. Ann. Eugen., 16: 351-358. Bennett, W. C. 1949 A cross-cultural survey of South American Indian Tribes, Architecture and Engineering, Habitations. B. A. E. Bull., 143, H.S. A.I., 5: 1-20. Best, W. R., M. Layrisse and R. Bermejo 1962 Blood group antigens i n Aymark and Quechua speaking tribes from near Puno, Peru. Am. J. Phys. Anthrop., 20: 321-329. Boyd, W. C. 1939 Blood groups of American Indians. Am. J. Phys. Anthrop., 25: 214-235. 1954 Maximum likelihood method for estimation of gene frequencies from MNS data. Am. J. Hum. Genet., 6: 1-10. 81 Brown, K. S., B. L. Hanna, A. A. Dahlberg and H. H. Strandskov 1958 The distribution of blood group alleles among Indians of Southwest North America. Am. J. Hum. Genet., 10: 175195. Chown, B., and M. Lewis 1953 The ABO, MNSs, P, Rh, Lutheran, Kell, Lewis, Duffy and Kidd blood groups and the secretor status of the Blackfeet Indians of Alberta, Canada. Am. J. Phys. Anthrop., 11: 369-383. 1957 The Kell antigen in American Indians. Am. J. Phys. Anthrop., 15: 149-156. Cooper, M. M. 1946 The Araucanians. B.A.E. B ~ l l . ,143, H.S.A.I., 2: 687-760. Cutbush, M., and P. L. Mollison 1950 The Duffy blood group system. Heredity, 4: 383389. Drabkin, D. L. 1946 Spectrophotometric studies; crystalographic and optical properties of hemoglobin of man in comparison with those of other species. J. Biol. Chem., 164: 703-723. Ellis, F. R., I. P. Cawley and G. W. Lasker 1963 Blood groups, hemoglobin types, and secretion of group-specific substance at Hacienda Cayalti, North Peru. Human Biology, 35: 26-52. Etcheverry, R., C. Guzman, A. Hille, R. Nagel, E. Covarrubias, A . Montenegro and Norma Duran 1966 Blood groups and other genetic blood characters in the Mapuche Indians. Rev. Med. Chile, in press. Etcheverry, R., E. Boris, C. Rojas, J. Villagran, C. Guzman, M. Muranda, C. Regonesi and Norma Duran 1966a Blood groups and other genetic characters in Fuegians Indians: Alacalufes and Yaganes. Rev. Med. Chile, in press. Etcheverry, R., E. Boris, C. Guzman, R. Nagel, R. Cruz-Coke, H. Blanc, C. Regonesi, M. Muranda, N. Duran and L. Suarez 196613 Blood groups and other genetic blood characters in natives of Easter Island. Rev. Med. Chile, in press. Fernandes, J. L., P. C. Junqueira, H. Kalmus, F. Ottensooser, R. Pasqualin and P. Wishart 1957 P. T. C. thresholds, colour vision and blood factors of Brazilian Indians. I. - Kaingangs. Ann. Hum. Genet., 22: 16-21. Gershowitz, Henry, P. C. Junqueira, F. M. Salzano and J. V. N e d 1967 Further studies on the Xavante Indians. 111. Blood groups and ABHLea secretor types in the Simoes Lopes and Sao Marcos Xavantes. Am. J. Hum. Genet., 19: 502-5 13. Giblett, Eloise R. 1962 The plasma transferrins. Progress i n Medical Genetics, 2nd ed., A. G. Steinberg, Grune and Stratton, New York. Giblett, E. R., and W. R. Best 1961 Haptoglobin and transferrin types in Peruvian Indians. Nature, 192: 1300-1301. Gray, M. P., and W. S. Laughlin 1960 Blood groups of Caddoan Indians of Oklahoma. Am. J. Hum. Genet., 12: 86-94. Hart, Mia V. D., H. Bosman and J. J. van Loghem 1954 Two rare human blood group antigens. Vox Sanguinis, 4: 108-116. Hulse, F. S. 1957 Linguistic barriers to geneflow. The blood groups of the Yakima, Okanagon and Swinomish Indians. Am. J. Phys. Anthrop., 15: 235-246. 82 G. A. MATSON, H. E. SUTTON, Layrisse, M., and J. Wilbert 1960 E l antigen0 del sistema sanguine0 Diego. Editorial Sucre, Caracas, Venezuela, 160 pages. Layrisse, M., Z. Layrisse and J. Wilbert 1963 Blood group antigen studies of four Chibchan tribes. Am. Anthropol., 65: 35-55. 1963a The blood groups of Northern Continental Caribs. Human Biology, 35: 140166. Layrisse, M., Z. Layrisse, E. Garcia and J. Wilbert 1961 Blood group antigens tests of the Yaruro Indians. S. W. J. Anthrop., 27: 198-204. - 1962 Blood group antigens of the Pemon Indians of Venezuela. Am. J. Phys. Anthrop., 20: 411-420. Levene, R. 1937 A history o f Argentina. The University of North Carolina Press, i-xiii plus 565 pages. Levine, P., G. A. Matson and H. F. Schrader 1935 Distribution of blood groups and agglutins gen M among Indians “Blackfeet” and “Blood” tribes. Proc. Suc. Exper. Biol. Med., 33: 297299. Lewis, M., J. A. Hildes, H. Kaita and B. Chown 1961 The blood groups of the Kutchin Indians a t Old Crow, Yukon Territory. Phys. Anthrop., 19: 383-389. Lothrop, S. K. 1946 The Diaguita of Chile. B.A.E. Bull., 143, H.S.A.I., 2: 633-636. Lowie, R. H. 1949 Social and political organization of the tropical forest and marginal tribes. B.A.E. Bull., 143, H.S.A.I., 5: 313-410. Matson, G. A., and C. L. Piper 1947 Distribution of the blood groups M-N, Rh types and secretors among the Ute Indians of Utah. Am. J. Phys. Anthrop., 5: 357-368. Matson, G. A., and H. J. Roberts 1949 Distribution of the blood groups, M-N and Rh types among Eskimos of the Kuskokwim Basin in Western Alaska. Am. J. Phys. Anthrop., 7: 109122. Matson, G. A., and H. F. Schrader 1933 Blood grouping among the Blackfeet and Blood tribes of American Indians. J. Immunol., 25: 155163. Matson, G. A., and J. Swanson 1959 Distribution of hereditary blood antigens among the Maya and non-Maya Indians in Mexico and Guatemala. Am. J. Phys. Anthrop., 27: 49-74. 1961 Distribution of hereditary blood antigens among Indians in Middle America: I. Lacandon and 0th- Maya. Am. Anthrop., 63: 1292-1322. 1963a Distribution of hereditary blood antigens among Indians in Middle America: 11. Tzotzil and other Maya. Am. J. Phys. Anthrop., 22: 1-14. - 1963b Distribution of hereditary blood antigens among Indians i n Middle America: 111. In Guatemala. Am. J. Phys. Anthrop., 22: 301-318. 1963c Distribution of hereditary blood antigens among Indians in Middle America: IV. In Honduras. Am. J. Phys. Anthrop., 21: 319-334. 1963d Distribution of hereditary blood antigens among Indians in Middle America: V. In Nicaragua. Am. J. Phys. Anthrop., 21: 545-559. J. SWANSON AND A. ROBINSON 1964 Distribution of hereditary blood antieens among Indians in Middle America: VI. fn British Honduras. Am. J. Phys. Anthrop., 22: 2 7 i - x ~ . 1965a Distribution of hereditary blood antigens among Indians in Middle America: VII. In Costa Rica. Am. J. Phys. Anthrop., 23: 107-121. 1965b Distribution of hereditary blood antigens among Indians in Middle America: VIII. In Panama. Am. J. Phys. Anthrop., 23: 413-426. Matson, G. A., E. A. Koch and P. Levine 1954 A study of the hereditary blood factors among the Chippewa Indians of Minnesota. Am. J. Phys. Anthrop., 12: 413426. Matson, G. A., P. Levine and H. F. Schrader 1936 Distribution of the subgroups of A and the M and N agglutinogens among the “Blackfeet” Indians. Proc. SOC.Exper. Biol. Med., 35: 4647. Matson, G. A., J. Swanson and Abner Robinson 1966b Distribution of hereditary blood groups among Indians in South America: 111: In Bolivia. Am. J. Phys. Anthrop., 25: 13-34. Matson, G. A,, H. Eldon Sutton, Jane Swanson and A. R. Robinson 1963 Distribution of haptoglobin, transferrin, and hemoglobin types among Indians of Middle America: Southern Mexico, Guatemala, Honduras and Nicaragua. Human Biology, 35: 474-483. 1966a Distribution of hereditary blood groups among Indians in South America. 11: In Peru. Am. J. Phys. Anthrop., 24: 325-349. 1967b Distribution of blood groups among Indians in South America. VI: In Paraguay. Am. J. Phys. Anthrop., 29: 81-98. Matson, G. A., H. Eldon Sutton, Raul Etcheverry, B., Jane Swanson and Abner Robinson 1967 Distribution of hereditary blood groups among Indians in South America. IV: In Chile. With inferences concerning genetic connections betweem Polynesia and America. Am. J. Phys. Anthrop., in press. Matson, G. A., H. E. Sutton, J. Swanson, A. E. Robinson and A. Santiana 1966 Distribution of hereditary blood groups among Indians in South America. I: In Ecuador. Am. J. Phys. Anthrop., 24: 51-69. Matson, G. A., J. Swanson, J. Noades, R. Sanger and R. R. Race 1959 A ‘hew” antigen and antibody belonging to the P blood group system. Am. J. Mum. Genet., 11: 26-34. Mazza, S. 1939 Los factores M y N en sangre de indigenas del Chaco argentino comparados con 10s de nativas de Buenos Aires. SOC. Arg. de Patol. Reg., 9a. Reunidn, Mendoza, 3: 1916- 1918. Mazza, S., and I. Franke 1927 Grupos sanguineos de indios y de autcktonos del norte argentino. Prensa Med. Arg., 14: 408-409. Mazza. S., K. Schurmann and H. Gutdeutsch 1933 apud Steffan, P., and S. Wellisch, 1936. Die geographische Verteilung der Blutgruppen. Z. f u r Rassenphysiologie, 8: 3 8 4 7 . Mbtraux, A. 1946 Ethnography of the Chaco. B.A.E. Bull., 143, H.S.A.I., 1: 197-370. 1948 Tribes of the Eastern Slopes of the Bolivian Andes. B.A.E. Bull., 143, H.S.A.I., 3: 465-485. BLOOD GROUPS O F INDIANS IN ARGENTINA - 1949 Warfare, cannibalism, and human trophies. B.A.E. Bull., 143, H.S.A.I., 5: 383-409. Miranda, F. M. 1946 The Diaguita of Argentina. B.A.E. Bull., 143,H.S.A.I., 2: 637-654. Nagel, R., and R. Etcheverry 1963 Types of haatonlobins i n Araucanian Indians of Chile. Nature, 197: 187-188. Neel, J. V., and F. M. Salzano 1966 A prospectus for genetic studies on the American Indians. The Biology of Human Adaptability. P. T. Baker and J. S. Weiner, eds. Oxford Clarendon Press, 1, 245-274. Neel, J. V., F. M. Salzano, P. C. Junqueira, F. Keiter and D, Maybury-Lewis 1964 Studies on the Xavante Indians of the Brazilian Matto Grosso. Am. J. Hum. Genet., 16: 52-140. Nunez Montiel, J. T., R. A. Perez and A. Nunez Montiel 1957 Estudio hematologica en grupos indigenas del Estado Zulia, Sistemas ABO, MN, Rh, Duffy, Kell y Diego. Acta Cient. Venezol., 8: 10-13. Cited by Salzano ('61). Owen, J. H., J. Silberman and C. Got 1958 Detection of haemoglobin, haemoglobin-haptoglobin complexes and other substances with peroxidase activity after zone electrophoresis. Nature, 182: 1373. Pantin, A. M., and J. C. Junqueira 1952 Blood groups in Brazilian Indians. Am. J. Phys. Anthrop., 30: 395-405. Pantin, A. M., and R. Kallsen 1953 The blood groups of the Diegueno Indians. Am. J. Phys. Anthrop., 11: 91-96. Parker, W. C., and A. G. Beam 1961 Haptoglobin and transferrin gene frequencies in a Navajo population: a new transferrin variant. Science, 134: 106-108. Paulotti, 0. L., and L. G. Alegria 1943 Grupos sanguineos de 10s nativos de la Puna Jujeila. A n del Mus. Arg. de Cienc. Nat., 41: 21-28. Pollitzer, W. S., R. C. Hartman, H. Moore, R. E. Rosefield, H. Smith, S. Hakin, P. J. Schmidt and W. C. Leyshon 1962 Blood types of the Cherokee Indians. Am. J. Phys. Anthrop., 20: 33-43. Race, R. R., and R. Sanger 1962 Blood Groups i n Man, 4th Ed., Springfield: Charles C Thomas. Race, R. R., R. Sanger, F. H. Allen, L. K. Diamond and B. Niedziela 1951 Inheritance of the human blood-group antigen Jka. Nature, 168: 207. Robinson, A. R., M. Robson, A. P. Harrison and W. W. Zuelzer 1957 A new technique for differentiation of hemoglobin. J. Lab. Clin. Med., 50: 745-752. Rosenfield, R. E., P. Vogel, N. Gibbel, H. Ohno and G. Haber 1953 Anti-Jka: three new examples of the isoantibody. Frequency of the factor in Caucasians, Negroes and Chinese of New York City. Am. J. Clin. Path., 23: 12221225. Salazar-Mall&, M. 1949 El agglutinogeno Lewis en la sangre de 10s Mexicanos. Bol. Inst. Estud. Med. Biol. Mex., 7: 25-28. Salzano, F. M. 1957 The Hood groups of South American Indians. Am. J. Phys. Anthrop., 15: 555-580. 1961 Studies on the Caingang Indians. 111. Blood groups. Am. J. Phys. Anthrop., 19: 391404. 83 1964 Salivary secretions of Indians from Santa Catarina Brazil. Am. J. Hum. Genet., 16: 301-310. Salzano, F. M., and H. E. Sutton 1963 Haptoglobin and transferrin types in Southern Brazilian Indians. Acta Genet., Basel, 13: 1-8. 1965 Haptoglobin and transferrin types of Indians from Santa Catarina, Brazil. Am. J. Hum. Genet., 17: 280-289. Sandoval, L., and C. Henckel 1954 The ABO, MNS, and Rh-Hr blood groups of the Mapucbe Indians of Cautin Province, Chile. Human Biology, 26: 324-329. Sanger, R. 1955 An association between the P and Jay systems of blood groups. Nature, Lond., 176: 1163-1164. Santiana, Antonio 1947 Los grupos sanguineos de 10s Indios del Ecuador. Comunicacion Definitive Imp. de la Univcrsidad, Quito, Ecuador, 1-47. 1953 Los Indios del Ecuador y sus caracteristicas serologicas. Boletin de Informaciones CientScas Nacionales, 6: no. 55, 52-64. Smithies, 0. 1955 Zone electrophoresis in starch gels; group variation in serum proteins of normal human adults. Biochem. J., 61: 629641. 1959 A n improved procedure for starchgel electrophoresis: further variations in the serum proteins of normal individuals. Biochem. J., 71: 585-587. Stevens, W. L. 1938 Estimation of blood group gene frequencies. Ann. Eugen., 8: 362-375. Steward, J. H. 1949 South American Cultures: a n interpretative summary. B.A.E. Bull., 143, H.S.A.I., 5: 669-772. Suarez, M. O., B. Broder and A. Loayza 1964 Antigenos de Grupos Sanguineos e n Indios Chiyapas de Bolivia. Anales de la Academia Nal. de C. Cuaderno 2. Serie Ciencias de l e Naturaleza Publ. no. 4, 8 pages, 3 plates. Suarez, M. O., E. Linares I. and J. Gutierrez B. 1965 Grupos Sanguineos en Poblaciones del Altiplano Andino con Bocio Endemico. Anales de la Academia Nal. de C. Cuaderno 2. Serie Ciencias de la Naturaleza Publ. no. 5, 6 pages, 1 map, 3 plates. Sutton, H.E., G. A. Matson, A. R. Robinson and R. W. Kousky 1960 Distribution of haptoglobin, transfenin, and hemoglobin types among Indians of Southern Mexico and Guatemala. Am. J. Hum. Genet., 12: 338-347. Tejada, C. M. Sanchez, M. A. Guzman, E. Bregni and N. S. Scrimshaw 1961 D,istribution of blood antigens among Guatemalan Indians. Human Biology, 33: 319-334. Wiener, A. S. 1965 Lewis blood types: theoretical implications and practical applications. Techn. Bull. Reg. Med. Techs., 35: 388-392. Wiener, A. S.,J. P. Zepeda, E. B. Sonn and R. H. Poliyka 1945 Individual blood differences in Mexican Indians, with special reference to the Rh blood types and Hr factor. J. Exp. Med., 81: 559-571. Zuelzer, W. W., J. V. Neil and A. R. Robinson 1956 Abnormal hemoglobins: Progress in Hematology. Grune Stratton, Inc., New York, 1: 103-104.