DOCUMENT RESUME

ED 063 980 RC 006 143

TITLE Biomedical Challenges Presented by the American Indiah, Proceedings of the Special Session of the PAHO Advisory Committee on Medical Research (7th, Washington, D.C., June 25, 1968). INSTITUTION Pan American Health Organization, Washington, D. C. REPORT NO Sci-Pub-165 PUB DATE Sep 68 NOTE 188p.

EDRS PRICE MF-$0.65 HC-$6.58 DESCRIPTORS *American Indians; Anthropology; Biochemistry; *Biological Influences; Conference Reports; *Genetics; *Health; *Human Geography; Tables (Data)

ABSTRACT At each meeting of the Pan American Health Organization Advisory Committee on Medical Research, a special 1-day session is held on a topic chosen by the committee as being of particular interest. At the 7th meeting, which convened in June of 1969 in Washington, D.C., the session surveyed the origin, present iistribvtion, and principal bioJogical and medical issues of the American Indian--including the Lroblems of newly contacted Indian groups and those of groups well along in transition. Recorded in this volume are the following papers and ensuing discussions:(1) The Origin and Dispersion of American Indians in North America; (2) Theses for Meditation on the Orijin and Dispersion of Man in South America;(3) Biological Subdivisions of the Indian on the Basis of Physical Anthropology;(4) Biological Subdivisions of the Indian on the Basis of Genetic Traits;(5) The American Indian in the International Biological Program; (6) Survey of the Unacculturated Indians of Central and South America; (7) Medical Problems of Newly Contacted Indian Groups;(8) The Problem of Gallbladder Disease among Pima Inlians;(9) Hyperglycemia in Pima Indians;(10) Malaria in the American Indian;(11) Food and Nutrition of the Maya Before the Conquest and at the Present Time;(12) Iodine Deficiency Without Goiter in Isolated Yanomama Indians; and (13) Study of Endemic Goiter in the American Indian.(Author/LS) BIOMEDICAL CHALLENGES PRESENTED

BY THE AMERICAN INDIAN

F roceedings of the Special Session held during the Seventh Meeting of the PAHO Advisory Committee on Medical Research 25 June 1968

U.S. DEPARTMENT OF HEALTH, EDUCATION WELFARE OFFICE OF EDUCATION THIS DOCUMENT HAS BEEN REPRO- DUCED EXACTLY AS RECEIVED FROM (HE PERSON OR ORGANIZATION ORIG- INATING IT. POINTS OF VIEW OR OPIN- IONS STATED DO NOT NECESSARILY ,RORESENT OFFICIAL OFFICE OF EDU- CAT)ON POSITION OR POLICY.

Scientific Publication No. 165 September 1968

PAN AMERICAN HEALTH ORGANIZATION Pan American Sanitary Bureau, Regional Office of the WORLD HEALTH ORGANIZATION 525 Twenty-third Street, N.W. Washington, D.C., 20037 NOTE At each meeting of the Pan American Health Organization Advisory Committee on Medical Research,a special onezy session is held on a topic chosen by the Committee as being of particular interest. At the Seventh Meeting, which convened in lune 1968 in Washington, D.C., the' session surveyed the origin, present distribution, and principal biological subdivisions of the American Indian and considered the specific scientific and medical issues calling for clarification, including the problems ofnewly contacted Indian groups and those of groups well along in transition. This volume records the papers presented and the ensuing discussions. PAHO ADVISORY COMMITTEE ON MEDICAL RESEARCH

Dr. Herr& Alessandri Dr. Alberto Hurtado Ex-Decano, Facultad de Medicina Rector, Universidad Peruana Cayetano Heredia Universidad de Chile Limq, Perti Santiago, Chile

Dr. Otto G. Bier Dr. Walsh McDermott Diretor, Departamento de Microbiologia e Imu- Chairman, Department of Public Health nologia Cornell University Medical College Escola Pau lista de Medicina New York, New York, J S.A. São Paulo, Brasil Dr. James V. Neel Dr. Roberto Caldeyro-Barcia Chairman, Department of Human Genetics Jefe, Departamento de Fisiopatologia University of Michigan Medical School Facultad de Medicina Ann Arbor, Michigan, U.S.A. Universidad de la RepUblica Montevideo, Uruguay Dr. Marcel Roche ( Vice-Chairman) Dr. Crtr los Chagas Director, Instituto Venezolano de Investigacio- Chief, Brazilian Delegat ion to UNESCO nes Cientificas Paris, France Caracas, Venezuela Dr. Philip P. Cohen Chairman, Department of Physiologicri Chemis- Dr. James A. Shannon try Director, National Institutes of Health The University of Wisconsin Bethesda, Maryland, U.S.A. K-?dison, Wisconsin, U.S.A.

Dr. Rene Dubos (Chairman) Dr. John C. Waterlow Member and Professor Director, Tropical Metabolism Research Unit The Rockefeller University University of the West Indies New York, New York, U.S.A. King:-.ton, Jamaica

Dr Hermon E. Hilleboe Director, Division of Public He5ith Practice Profe' sor Abel Wolman School of Public Health and Administrative Emeritus Professor of Sanitary Engineering and Mcdicine Water Resources Columbia University The Johns Hopkins University New York, New York, U.S.A. Baltimore, Maryland, U.S.A.

Dr. Bernard, Houssay Director, Instituto de Biologfa y Medicina ;Ex- Dr. Salvador Zubirain perimental Director, Instituto Nacional de la Nutrición Buenos Aires, Argentina Mexico, D.F., Mexico

Dr. Mauricio Martins da Silva, Secretary Chief, Office of Research Coordination Pan American Health Organization Washington, D.C., U.S.A.

PAN AMERICAN HEALTH ORGANIZATION Pan American Sanitary Bureau Dr. Abraham Horwit7,, Director Special Session on INDIAN BIOMEDICAL CHALLENGESPRESENTED BY THE AMERICAN Moderator: Dr. James V.Neel PARTICIPANTS Dr. James B. Griffin Dr. Tulio Arends Museum of Anthropology Hematologia Experimental Instituto Venezolano deInvestigacionesCien- The University of Michigan Ann Arbor, Michigan, U.S.A. tificas Caracas, Venezuela Dr. William S. Laughlin Dr. Jose Barzdatto Department of Anthropology Hospital del Salvador The University of Wisconsin Universidad de Chile Madison, Wisconsin, U.S.A. Santiago, Chile Dr. Moises Behar Dr. Miguel A. Layrisse Instituto de Nutrición deCentro America y SecciOn de Fisiopatologia Investigaciones Cien- Panamá nstituto Venezolano de Guatemala, Guatemala tificas Caracas, Venezuela Dr. Thomas A. Burch Clinical Field Studies Unit Dr. Ruben Lisker National Institute of Arthritis andMetabolic Instituto Nacional de la Nutrición Diseases Mexico, D.F., Mexico Phoenix, Arizona, U.S.A. Dr. William Centerwall Dr. Max Miller D Tartment of Human Genetics Depanment of Medicine The University of Michigan Western Reserve University Ann Arbor, Michigan, U.S.A. Cleveland, Ohio, U.S.A. Dr. Napoleon A. Chagnon Department of Anthropology Dr. Jerry D. Niswander Human Genetics Branch The University of Michigan National Institute of Dental Health Ann Arbor, Michigan, U.S.A. Bethesda, Maryland, U.S.A. Dr. Juan Comas * Sección de Antropologia Dr. Noel Nutels Instituto de Investigaciones Históricas Serviço Nacional de Tuberculose Universidad Nacional Autónoma de WY iCo Ministerio de Saiide Mexico, D.F., Mexico Rio de Janeiro, Brasil Dr. Edmundo Covamibias Dr. Marcel Roche Departamento de Genetica Investigaciones Cien- Facultad de Medicina kstituto Venezolano de Universidad de Chile tificas Santiago, Chile Caracas, Venezuela Dr. Jose M. Cruxent Dr. Francisco M. Salzano Sección de Antropologia Departamento de Genitica Instituto Venezolano de InvestigacionesCien- Universidade Federal do Rio Grande do Sul tificas Pôrto Alegre, Brasil Caracas, Venezuela Dr. George Giglio li Dr. William J. Schull Malaria Laboratory Department of Human Genetics Ministry of Health The University of Michigan Georgetown, Guyana Ann Arbor, Michigan, U.S.A. Unable to attend.

iv CONTENTS

Opening Statementlames V. Ned 1

The a igin and Dispersion of American Indians in North America James B. Griffin 2

Theses for Meditation on the Origin and Dispersion of Man in South AmericaJose M. Cruxent 11 riscussion William S. Laughlin 17 General Discussion 19

Biological Subdivisions of the Indian on the Basis of Physical AnthropologyJuan Comas 22

Biological Subdivisions of the Indian on the Basis of Genetic TraitsMiguel Layrisse 35 Discussion Tulio Arends 40 Ruben LiAer 43 General Discussion 46

The American Indian in the International Biological Program fames 17 Ned 47 Discussion 55

Survey of the Unaccultufated Indians of Central and South AmericaFrancisco M. Salzono 59 Discussion 67 Medical Problems of Newly Contacted Indian Gro,..psNoel ZVutds 68 Discussion Willard R. Centerwall: A Recent Experience with Measles in a "Virgin-Soil" Population 77 General Discussion 80

The Problem of Gallbladder Disease among Pima IndiansThomas A. Burch, Leonard 1. Comets, and Peter H. Bennett 82

Hyperglycemia in Pima Indians: A Preliminary Appraisal of Its Significana Max Miller, Peter W. Bennett, and Thomas A. Burch 89

Malaria in the American IndianGeorge G. Giglioli 104 Food and Nutrition of the Maya Before the Conquest and at the Present Time Moises Behar 114

Iodine Deficiency Without Goiter in Isolated - Yanomama Indians: Preliminary Note René Ritnere, Dominique Comar, Mentech Colonomas, Jean Desenne, and Marcel Roche 120

Study of 7I:ndemic Goiter in the American Indian José Barzelatto and Edumundo Couarrubias 124 Discussion 1. D. Nistvander 133 Edtnundo Covarrublas 136 General Discussion 137

Appendix: Tabulations of Phenotype and Gene Frequencies for 11 Different Genetiz Systems Studied in the American IndianR. H. Post, J. V. Ned, and W. I. Schull 141

vi OPENING STATEMENT

James V. Neel, Moderator

The American Indian continues today, as he Further, it begins to appear as if once that has ever since the discovery of the Americas, totransition is accomplished the Indian may pre- present a wide variety of questions and chal- sentsomeratherspecialdiseaseproblems, lenges ranging from the purely scientific to the whose satisfactory solution will challenge all intensely humanitarian. In today's Special Ses- our medical ingenuity.' sion, we shall be abie to consider only a few of There is one important biomedical problem these many challenges. of the Indian that this session will leave com- Specifically, we shall deal first with some of pletely untouchedthe problem of high-altitude the unusual opportunities the Indian still offers populations. This subject was covered both at for the investigation of basic problems in ge-the Special Session of 19661 and at a WHO/ netics and anthropology. These oppottunitiesPAHO/IBP conference held here in November include studies of the nature and rate of human 1967. Under these circumstances, a further brief dispersion, of the emergence of human diversity, treatment at this time was deemed redundant. and of the biology of primitive man, wi-h all The number of participants in the Session is its ramifications. somewhat greater than usual, thanks to the In the latter half of the program, we shall turn generous financial assistance of boththe U.S. our attention to some of the moremedical and Nationql Committee for the International I3io- humanitarian problems presented by the Indian. logical Program and the Human Adaptability In many respects, these problems are similar to Section of the Commi.tee for the International those of the cam penno or caboclo throughout Biological Program of the International Union South America, to whose gene pool the Indian of Biological Sciences. has made a significant contribution. Today's formal presentations will be followed However, the Indian also presents some spe- tomorrow by an informal session atwhich prob- cial problems. One of these is the transition, in lems of common interest to the participants here the relatively unacculturated Indian, from "small today will be discussed, with particular reference band" epidemiology to "large herd" epidemi- to the International BiologicalProgram. ology. As we are all aware, this transition has It is customary at such a session to designate repeatedly had a fatal impact on Indian groups. two rapporteurs. This year Dr.Napoleon Chag- The medical challenge to blunt this impact is non and Dr. Edmund Covarrubiashave kindly clear ane, persistent. In addition, by taking it up agreed to discharge that function. we preserve at least temporarily oneof the last great resources for the study of primitive man, one of the last opportunities to attempt tofathom the nature of the forces to which man was re- The proceedings appear in Lik at High Altitudes, sponding during the course of human evolution. PAHO Scicntifi, Publication 140.

1 THE ORIGIN ANDDISPERSION OF AMERICANINDIANS IN NORTH AMERICA

James B. Griffin

tations and can be read bythose who care to The historically effectivediscovery of the New World by Europeans in 1492 openedthe Ameri- do so. cas tolarge-scale colonization and to along The temporal framew ork for earlyIndian groups ultimate origin of period of speculation on the human groups almost four hun- The time of arrival of the first the New World peoples. For likely to remain a framework was is far from settled and is dred years the chronological research problem for some time. For purposes based on biblical sources and was notadeo,uate of this paper I shall refer tothe period of time cultural diversity of New World to explain the from the first arrival of man toabout 8000 B.C. culture. Prominent among thespeculations on the Paleo-Indian period,and allcultural origins were the multiple-migrationhypotheses, as complexes and skeletal materialof this age, if which provided civilized Mediterranean,Euro- any, will beincluded within this period. Arapid pean, and even SoutheastAs;an groups to bring review of evidence from theUnited States and higher culture into the Americas and a level of Canada includes only onefairly homogeneous produce the more complex civilizationsof Meso- and widely dispersedcomplex, a few examples america and the Andes. Suchexplanations were of other early Paleo-Indianperiod peoples, and also employed to explain a presumedmultiracial a numberof purported sites that are not re- origin of the American Indian. garded in this paper as soundlyestablished. As early as the sixteenth century somewriters The Fluted Point Hunters arethe sole recog- recognized the predominantly Asiaticrelation- nizable group with a continent-wide occupancy ships of the native Americanpopulations and in the latter part of thePaleo-Indian period from postulated that the movement ofpeople tookabout 10,000 to 8000 B.C. Theearliest radio- place from northeast Asia to northwesternNorthcarbon dates in association with anestablished America. Thisis,of course, the only routecultural context are about 9300 to9000 B.c., of entry seriously considered by contemporaryfrom sites in Arizona, NewMexico, Colorado, scholars, but thereisconsiderable room for and Oklahoma. This is referred to asthe Llano disagreement in many phases of the originand complex, with the Clovis fluted point as amajor dispersion of American Indianpopulations. For diagnostic tool. The Sandia finds areregarded this paper I have chosen to refrainfrom present- as a potentialpredecessor, with an uncertain age ing many reference citationsto support theprobably within a thousand yearsof Clovis. In points of view included in the paperand fromthe western plains the laterFolsom assemblages directly citing other views. InsteadI have in- at several siteshave been dated between 9000 United States and cluded in the references a list ofbooks and other and 8000 B.C. In the eastern from thc South articles that afford a wide spectrumof interpre-Canada, fluted points are known

2 Atlantic and Gulf coasts to the Great Lakes and North America for which considerable antiquity Ontario and into New England, and asfar has been claimed as places inhabited by early northeast as Nova Scotia. At the Debert site in Indians. Even whole books have been published Nova Scotia, a series of radiocarbon dates sug-on non-sites. The reasons why it is now difficult gest that man occupied the area between 9000 or impossible to include such "finds" here varies and 8500 B.C. Provisional correlations of the dis- from location to location; a detailed dissent is tribution of fluted points in the Great Lakes not within the scope of this paper. In this cate- area have proposed an age of from 10,000 to gory I would include the claims for occupation 8000 B.C. for that area, and the evidence from at Tule Springs, Nevada, of the order of 20,000 the Southeast also suggests an antiquity for to 30,000 years ago; on Santa Rosa Island, much fluted points of abuut the same order. before 8000 B.c.; at the Scripps Institute bluff at In the area west of the Rocky Mountains there La Jolla, California, slightly over 20,000B.C.; are a number of complexes that may begin at Lewisville, Texas, more than 37,000 B.C.; at before 8000 B.c., but as yet this has not beenSheguiandah, Manitoulin Island, Ontario, for adequately demonstrated. Among these are thea cultural complex much before 7000 B.c.; for Desert Culture, Lake Mohave and the Sana completely pebble-tool cultureinnorthern Dieguito culture of southern California, and aAlabama of extreme antiquity; for a chopper/ number of sites and complexes from Oregon tochopping-tool complex of aninterglacial or British Columbia and Washington compressed interstadial period, or of a simple bone-tool tra- by some archaeologists into an Old Cordilleran dition of any age; or for Pleistocene man in the culture. In extreme northwest Canada, the Brit- Trenton, New Jersey, gravels. ish Mountain complex is assigned considerable In Mexico particularly and alsoin Central antiquity, butitsage, except on typologicalAmerica, there are indications of the presence grounds, is not known, and no archaeologicalof man before 8000 B.c., but many of these assemblage from Alaska is as old as 8000B.C. identifications were made quite a number of The oldest radiocarbon dates in association years ago and suffer from a lack of sound dates with human habitation in the United States are or are isolated artifacts inadequate for the re- from Wilson Butte Cave in south-central Idaho, construction of a cultural assemblage. Some of where from the lower zone of Stratum C therethese finds from the Late Pleistocene Upper Be- is a date of 12,550 -± 500B.C.(M-1409) and cerra formation may well record human occupa- from Stratum E a date of 13,050 ± 800B.C. tion, and continuing excavations in Mexico will (M-1410). The context of the material in the eventually place the temporal position and in- cave indicates the presence of man with extinct dustrial activities during the Pa leo-Indian period fauna such as camel and horse and some Boreal-. on a firmer basis. At present investigations are zone microfauna, but the few artifacts found being conducted in the Valsequillo gravels of are not particularly diagnostic (7). Puebla andatTlapacoya intheValley of Fluted points are known from Mexico and as Mexico. As usual, thereissome uncertainty far south as Guatemala and perhaps Costa Rica. about the temporal correlation of gravel deposits In none of these instances, however, is there an between one area and another, about radio- associated industry. Most of the few fluted points carbon dates of 35,000 to 24,000 years ago of are found in northern Mexico, where they arehigh antiquity but not directly associated with on the southern fringe of the Llano and Folsom adequate artifacts, or about the precise age of a concentrations in the Southwest and western cultural complex on a buried living surface. Texas. There are no soundly established cultu- While the South American evidence for early ral assemblages in Mexico and Central America Indian occupations is to be evaluated by Profes- directly dated by radiocarbon before 8000 B.C.sor J. M. Cruxent at this session, I shall include There are a large number of locations in some observations on this area because of its

3 importance in assessing the probable age of thc islands. A radiocarbon date of 12,800-± 120B.C. firstinhabitantsof North America. On the(GIN-97) has recently been obtained on fossil north side of the Strait of Mar flan, in Fellsbone from the lower cultural level of Marta Cave, there is a date of 8760B.C.(W-915) ob- near Irkutsk, often attributed to the older Upper tained by Junius Bird. From eastern Brazil, atPaleolithic of the Irkutsk area. A date of 18,950 Lagoa Santa, there is evidence of occupation -± 300B.C.was obtained on charcoal from the around 8000 to 7000 B.c., and a sambaquis on lower cultural level of Afontova Gora II in the the southeastern Brazilian coast has provided a Upper Yenesei Valley near Krasnoyarsk (TO. date close to 6000B.C.In northeastern Vene-From Kamchatka there is a date of close to zuela, at the Muaco site, there is a possible asso- 18,000 B.C. on charcoal irom the Uski Isite. ciation of man and extinct fauna in the period Other sites in southwest Siberia are presumed to aom 14,500 to 12,000 .J.c. In the northern An-date substantially earlier, and sites in Japan of dean area, artifact complexes have been given an Upper Paleolithic cast date well back toward ages in the neighborhood of 8000B.C. the 20,000-year range. The South American data, with their wide The southern Siberian complexes of about geographic spread of early man around or be- 30,000 to 12,000B.C.have a strong early rela- fore 9000 to 8000B.C.,imply the arrival of mantionship to the late Mousterian stone industries on that continent substantially before the known of eastern Europe. As the Upper Paleolithic dated complexes. Similarly, in Mexico and Cen- developed, there was corresponding modification tral America the wide distribution of human in Siberia, but the patterns of change in Siberia occupations just before or after 8000B.C.implies are sufficiently different from those of the better- that the arrival of the first human groupsVIM known areas to the west that an easy alignment substantially before this date. If the proposedhas not been possible. The industrial develop- dates for man in the Valley of Puebla and the ment of these Siberian populations was the re- Valley of Mexico are confirmed to be between sult of the long Eurasian cultural development, 40,000 and 24,000 years ago, there will be much which became adapted to the late Pleistocene. work for archaeologists to do in the future toThe level of occupationisattributedtothe find substantiating evidence in the rest of thelower section of deposits on Terrace II of the New World. The North American geographical Yenesei. The faunal composition represet* cold spread of dated evidence and the considerableperiglacial conditions, and the date corrwonds diversity of assemblages shortly after 8000 B.C. to about the maximum of the last major Siberian imply an antiquity of man in North Americaglaciation, called the Sartan. Also in the Middle considerably greater than thc known agc of the Yenesei,at the Kokorevo sites,radiocarbon Fluted Point Hunters or of the occupants ofdates range from about 14,000 to 11.000B.C. Wilson Butte Cave. In summary, an agc of about In the same arca at the Marta site, a datc of 15,000 years for man in thc New World is about 7000B.C.was obtained on a Mesolithic- viewed as reasonable, with thc possibility thatlike complex located on the higher areas of the it may be considerably greater. Terrace I floodplain deposits. In these Siberian sites there are crude heavy The temporal framework for northeast Asia chopping tools, a variety of flake implements Sound dating of the Late Pleistocene occupa- including scrapers and knives, discoidal corcs, tions of Siberia is just beginning; in fact, ade-and some bifacially flaked poims or knives. quate investigation of ancient man in north-There is a trcnd toward greatc 'se true eastern Siberia has only recently been initiated. blades made from prepared cores t? r., J the Most of the sites with an agc of morc than four fashioning of end and sidc sccar rs, or five thousand years are along the southcrn perforators, gravers and burins, and an incre borders of Siberia from Russia to the Japancscin bone tools and ornaments. The animals on

4 which the people fed are those from arctic toTheir way of life was developed from southern subarctic and cold arid steppe environments. Russia to south-central Siberia during the latter They made skin clothing and had substantialpart of the Pleistocene; it was based on a late houses in the construction of which they used Mousterian industry, modified by the initial ele- the bones of large mammals such as woollyments of Upper Paleolithic emphasis on blade rhinocerus and i.mmoth. Probably the most tools and the beginnings of a bone industry that important animal was the reindeer in the tundra was an aid in the production of skin clothing area. It might be said that the spread of manand shelter. This gradaal expansion northward into North America awaited the presence ofinto new territories from northeastern Siberia arctic-alpinetundraspecies,during thelate to Alaska would have taken place without resis- Pleistocene, on which man could liveas he tance from resident hunters. If it took place in hunted his way across northeastern Siberia into the time period suggested, then a substantial North America. number of hunting camps must now be under While the extent of the mountain glaciers in ocean water, but some will eventually be found eastern Siberia is not satisfactorily known, itis in favorable areas such as elevations overlooking certain that only a small part of the land masspasses followed by game animals in moving was glaciated and that most of the area wasfrom one feeding ground to another. occupied by xerophytic arctic tundra or alpine This early population spread isbelieved to tundra. A long tongue of steppe or periglacialhave been diverted south along the west side of steppe extended from southwest Siberia east-the McKenzie Valley. A number ofrecent ward between the Central Siberian Plateau and papers (9, 11 , 28) have emphasized the difficulty Lake Baikal as far as Yakutsk on the Middleof passing from the Lower McKenzie Valley Lena. From this area the best access route to the into the eastern Rocky Mountain slopes of the north was down the Lena Valley to the ArcticUnited States because of the presence of the Ocean. coalesced continental and cordilleran ice from The fall of the sea level during the last major Montana to the Yukon Territory along the glacial advance of the Wisconsin-Wurm is esti- eastern margin of the Canadian Rockies. The mated to have produced a land bridge at the evidence for the closing and opening of the Bering Strait from about 24,000 to 8000 ex., with corridor between these sheets is not so firmly two periods of submersion of the highest partestablished that sound datings for these events of the shelf correspcnding to major ice-melting are available. The position adopted in this paper phases of the retreat of the Wisconsin ice. Theis that the corridor would have been closed only size of the exposed land was considerable. Most at the maximum of the Wisconsin glaciation for of it was not forested but was occupied bya few thousand years, about 19,000 to 15,000 tundra vegetation similar to that of tne Siberian years ago. arid-steppe tundra. During the last glacial domi- If the early hunters came into the United nance, betwcen 23,000 and 10,000 Lc., the arctic States before 17,000 B.c., then archaeologists in trees and shrubs were more restricted in their the United States either have been unlucky or distribution than they are today and the climate have not been able to correctly evaluate the evi- was colder than it is now. dence for his occupancy before the 13,000 Lc. The movement of early man into date mentioned at Wilson Butte Cave. Entry North America shortly before 17,000 B.c. would allow ample Keeping in mind the lack of direct evidence timeforpenetrationinto extreme southern for the presence of early man in northwestern South America for the known occupancy there, North America and northeastern Siberia, we but would not accommodate the proposed Valse.- can still present an acceptable hypothesis for a quillo and Tlapacoya occupations in Central spread of hunting bands from west to east. Mexico.Ifthe corridor was closed between

5 21,000 and 10,000 p.c., archaeologists are faced an early post-Valders invasion of the east by with at least as impressive dilemmas in the form animalforms now associatedwith western of an absence of sound data representing man in prairie environments, and this would correlate North Amerka before 21,000 B.C. and the long with recent similar hypotheses of prairie vegeta- period from then to 13,000 B.c., or in accepting tion movement eastward at an early period (8). the speed with which man moved from Alberta The park-tundra and coolprairie would be to Tierra del Fuego. suitable for barren-ground caribou, which have The environmental changes in North America been identified in Michigan and New York. Did as a result of the retizat of the Wisconsin icethey penetrate this far south before thelast would have had an effect upon the way of lifeWisconsin advance, or did they arrive with the of ancient man thrlugh the shift of climatic "reopening" of the corridor? zones, vegetation, and animal life. The expan- sions of the Canadian continentalicesheets The Fluted Point Hunters of North America effectively obliterated the vegetation and animal By 10,000 to 8000 p.c. the people of the Late life from much of Canada. The expansion of the Pa leo-Indian period had occupied sparsely most ice into the northern sections of the United of the area south of the present Canadian boreal States markedly altered the biota, and com- forest to South America and from the Pacific to pressed and interdigitated elements of previousthe Atlantic. Most of these populations were periods into assemblages distinctive to late Wis- strongly dependent on hunting, as we know consin times. The climatic conditions during the from the spear and dart points, knives, and life of the western mountain glaciers lowered the scrapers of various kinds to work skins and tree line, changed the faunal associations and from the fact that these tools have been found in distributions, and produced thousands of lakes association with a small number of large game in the now-dry basins of the western Plains, in animals. This latter fact gave rise for a time to the Southwest, in the intermontane plateau re-the idea of almost a limitation of diet to big gion, and in the Pacific Coast states and Mexico. game animals. Data from sites such as Linden- The lowered forest zones and more extensivemeier in Colorado, Graham Cave in central and effectivegrasslands supportedthelarge Missouri, and others prove that the meat diet grazing and browsing animals ofthelate was quite varied, and at least in the east there Pleistocene fauna. There were more streams, is very little evidence of early man killing the with corridors of pine and spruce crossing the mammott and mastodon. In addition, the early grassland. Fluted Point Hunters would have recognized a The changes in climatic regime accompanying large variety of the plant foods available, from the withdrawal of the Wisconsin ice had already nuts to berries and tubers. The diet of early man produced notable shifts from thefullglacial was not likely to stay restricted to a few classes environments by the time of the early Flutedof foods when he entered environments with a Point Hunters of 10,000 to 9000 B.C. The shift wide variety of them. It should be possible to in vegetation and accompanying animal life took discover sites that will reflect varieties of food place on a large scale over North America, gathering and processing activitiesthat were causing some shifts in hunting and collectingpart of the life of the Fluted Point Hunters. areas, and assisted in the displacement or dis- They should have had some seasonal activity appearance of a small number of game animals. patterns. A number of students of the Pa leo- The northward movement of musk ox and Indian cultures are beginning to recognize re- mammoth is thought to have been in a park- gional tool and behavior complexes that will aid tundra vegetation zone which initially occupied our understanding of this earliest known com- the soilsleft free of glacial ice in the Great plex. Lakes area. It has been suggested that there was The wide distribution of Fluted Point bands

6 12. and the relative homogeneity of the implements prairie provinces, continues to reflect the exis- recovered implies a rather rapid spread of thesetence of a hunting economy with bison as an early hunting people, and apparently intoareas important supplier of food, tools, and clothes. not hitherto occupied. There is also the implica-At some locations there is clear evidence ofmass tion that there would have been continuingcon- killings, evidently the result of communal drives. tact between neighboring bands, perhaps forIndications of variability of animal foodcomes group hunting or other food-procuring activities from sites where giant beaver, pronghornante- at favorable seasonal locations or at locations lope, elk, deer, raccoon, coyote, and smaller favorable for shelter during the winterseasons. mammals as well as bisonwere part of the food Such collective activity would have permittedsupply. At other sites thereare indications of exchanges of new cultural developmentsingrinding and milling stones, burins, sandstone terms of sources of food, raw materials, manu- abraders, and whetstones. facturing techniques, hunting technologyand West of the Rocky Mountains from around of people. It is doubtful that individual bands 8000 B.C., and continuing for many millenia, wouid have been isolated from othergroups for archaeologistsrecognizetheDesertCulture, extensive periods or that peoples moving into which has a number of named regional variants new regional environments would have been cut from Mexico into Oregon and Washington. off by those environments from culture-sharingThese variants emphasize the gathering and with peoples in their former territory. preparation of small seeds by hand and milling stones; the hunting of a wide variety of animals; Archaic-period adaptations in North Americaand the extensive utilization of wood, bone, In the long Archaic period in the United hide, and vegetational sources for tools,orna- States between 8000 B.C. and the effective intro- ments, and containers. It was a gradually de- duction of agriculture around 1B.c., many re- veloping adjustment tothe essentiallydesert gional cultural developments occurredas the environment, which supported only small bands Indian groups became mcre familiar with local and in which population density remained low resources and develov.c.,the knowledgefor up to the historic period. It was not, howeva, a successfully exploiting _hem. As they didso, static complex, for many significant changes successful adaptations lo particular environments took place in the technology, some of themrep- tended to restrict band and group activity toresenting almost continent-wide shiftsintool these environments and to producea higherforms, and new weapons, and shiftsinthe level of exchange c; culture and people within techniques of manufacturing baskets. Important these areas than between them. This is reflected variants are recognized in areas along streams archaeologically by the growth of distinguishableand lakes, in upland forested or alpine areas, regional cultural traditions. and where minor shifting climaticpatterns One of the best-documented cultural continui- allowed the expansion of desert bands intosome- ties from the Pa!eo-Indian period Fluted Point times better watered areas or the penetration of Hunters to later complexes is in the western foreign groups into the Desert Culture region. Plains states. The production of fluted points Between 8000 and 6000 B.C. along the North- was gradually abandoned and nonfluted points west Coast and into the interior along the major and knives of essentially thesame basic formrivers, at least some part of the year was spent continued in use along with the rest of the stone in obtaining food from the spawning runs and tools. New tools appe r.such as the specialized in otherwise exploiting the food supply asso- Cody knife, and new techniques suchas theciated with the streams and coastalareas. The parallel flaking of the Scottsbluff and Eden latter is a reasonable inference, for some parts forms. All the evidence from sites in this region,of what was then the coast are now under water. from the Rio Grande north into the CanadianIt was to be a long time, however, before the

7

I striking Northwest Coast sea-adapted complex and a main hearth area was the Bering Strait on would develop between Washington and the both sides of the International Date Line, where Alaskan peninsula. people have been moving across in both direc- In the interior, from the Yukon territory to tions for many millenia. Idaho, there is an Old Cordilleran complex that In the woodland area of the eastern United may be viewed as an Asiatic-derived parent to Statesa gradual transitionisrecognized in the Fluted Point, as a collateral contemporane- several areas from the Fluted Point Hunters ous variant, or as the result of the northern andcomplex to assemblages maintaining the same western expansion of the Llano to Plano tradi-basic manufacturing stone-working techniques tion. At present there would seem to be a basic and tools, but with the dt:velopment or adoption relationship, and current radiocarbon dates indi-of nonfluted projectile forms very similar to cate the time period of Old Cordilleran as not some of the early Plano forms of the Plains. By over 8000 B.C. 7000 to 6000 B.C. stemmed and notched pro- InsouthernCaliforniathe SanDieguito jectile forms become common, and an increasing hunting culture with percussion-flaked lanceo-diversity of regional areas through time reflects late points, knives, scrapers, and choppers isthe increasing specialization of groups learning known from about 8000 B.C. to 6000 B.C. Shortly to exploit the resources of these local regions. afterward there is a development of a number Many oi the changes are related in form and of areal specializations in coastal, desert, and function to those of areas to the west, but the forest environments, which during the last few east acquires a distinctive flavor ofits own thousand years resulted in an unusually densethrough the development of a series of wood- pOpulation for a hunting-gathering populationworking tools and ground and polished stone in the oak-forest area of central California. forms. While regional specializations are present Southwestern Alaska was sparsely populatal in the eastern Archaic complexes, there is also by around 8000 to 6000 B.C. by people with a evidence of increasing exchange of raw materials unifacial core and blade industry whose move-or manufactured objects as travel or trade routes ment into Alaska is likely to have been from the become established. Pacific side of the Chuckchi peninsula into Many archaeologists view most of the cultural coastal Alaska and south to the eastern Aleutian complexes of Mexico from about 8000 to 4000 area. If these early coastal-adapted groups wereB.C. as a southern extension of the Desert Cul- the first Aleuts, as is implied, it would suggestture. Certainly the general hunting-gathering that the Bering Strait area was occupied bypattern is similar, and there arc some tool forms Eskimoan-speaking peoples longer ago than has that are held in common with early Archaic and been thought. In interior and northern Alaska Desert Culture groups from California to Texas. thereis a considerable variety of assemblages The most important feature of the Mesoamen- reflecting inland and coastal developments withcan scene is the early domestication of plants, continuing ties with Siberia, and also influences from Tamaulipas to Chiapas, which has been fromnorthward-spreadinggroupsprimarily demonstrated wheresystematiceffortshave moving with the expansion of bison and other been expended to search for such evidence. This game animals. area shows a very gradual increase in the num- Between 3000 and 2000 B.C.the Denbigh ber of plants domesticated in the several regions Flint Complex, with a marked coastal adapta-and in the proportion of domesticates consumed. tion, spread with surprising speed eastwird toMarked population increase in some areas such provide the first successful occupation of theas the Valley of Mexico, the Valley of Oaxaca, eastern Arctic. The first Eskinio bands reached the coastal lowland of southern Vera Cruz and northern Greenland by 2000 B.C. Eskimo cul-Tabasco, and thePacific Coast of adjacent tural traditions have a considerable time depth, Chiapas and Guatemala is observed by 1000 B.C.,

8

14 when agricultural practices were well developed man to move into North America and spread and the Early Formative cultures were becom- throughout the New World. ing established. The main access route into interior North Ameri.ca was east of the Rocky Mountains, and Summary dispersion into most of the Americas was by All the supportable evidence available indi- this route. Population increase and any physical cates that the first human occupants of Northdillerentiation of human groups south of the America came from northeastern Asia. Some Arctic area is derived primarily from the popula- archaeologists support the view that this first tions of the Pa leo-Indian period. There arc no occurred from 30,000 to 40,000 or more yearsindications in the prehistoric record of any later ago, others believe it was from about 25,000 to substantialmigrating groups influencing the 20,000 years ago, and some have contended that cultural life of the residents of North America it could not have been until about 12,000 years south of the Alaskan and Canadian Arctic ago. The time of arrival has not been settled. Some archaeologists emphasize the Mouste- region. rian origins of the first emigrants, believing that The archaeological evidence, except in rare the earliest American cuhural complexes indi-instances, supports the view that, in spite of cate a spread into North America before ele- regional adaptations to food supplies and raw ments of Upper Paleolithic origin had reached materials, there was a continuous exchange of eastern Asia. Many archaeologists, however, be- new developments between regions, with the lieve that Upper Paleolithic developments wereadditional implication of population interaction a part of the cultural mechanisms that allowed as well.

REFERENCES 1. AVELEYRA ARROYO DE ANDA, L. El Segundo 8. GUILDAY, J. Grizzly bears from eastern North Mamzet FoK1 de Santa Isabel Iztapan, Mexico, y America. Amer. Midland Nat. 79:247-250, 1968. Artektos Asociados. Mexico, D. F., Instituto Na- 9. HAYNES, C. V., JR. Fluted projectile points: cional de Antropoligia e Historia,1955.61pp. Their age and dispersion. Science 145:1408-1413, (DirecciOn de Prehistoria, PublicaciOn 1) 1964. 2. AVELEYRA ARROYO DE ANDA, L. Antigfiedad del 10. HAYNES, C. V., JR. Muestras de C-14, de Hombre en Mexico y Centroamerica:Cata logo Tlapacoya, Estado de Mexico. Mexico, D. F., Insti- Razonado de Localidades y Bibliografia Selectatuto Nacional de Antropologia y Historia, Boletin (1867-1961). Mexico, D. F., Universidad National 29, 1967, pp. 49-52. AutOnoma de Mexico, 1962. 72 pp. (Cuadernos del 11. HOPKINS, D. M., ed. The Bering Land Bridge. Instituto de 'Astoria, Serie Antropologica, 14) Stanford,California,StanfordUniversityPress, 3. CUSHING, E. J., and H. E. WRIGHT, JR., eds. 1967. 495 pp. Quaternary Paleoecology. Nein Haven and London, 12. IRWIN, C. lnfornie preliminar sobre las exca- vaciones en Valsequillo. Mexico, D. F., Archivos Yale University Press, 1967. 443 pp. del Instituto Nacional de Antropologia c Historia, 4. GOODLIFFE, E. M. Un sitiopleistocenica en 1965. Tlapacoya, Estado de Mexico. Mexico, D. F., Insti- 13. JENNINGS, J. D., and E. NORBECK, eds. Primi- two Nacional de Antropologia c Historia,Eolethi tive Man in the New World. Chicago, University 23, 1966, pp. 30-32. of Chicago Press, 1964. 633 pp. 5. GRIFFIN,J.B. Some prehistoric connections 14. KLEIN, R. G. Radiocarbon dates on occupation between Siberia and America. Science 131:801-812, sites of Pleistocene age in the U.S.S.R. Arctic An- 1960. thropology 1V:224-226, 1967. 6. GRIFFIN, J. B. Eastern North American archae- 15. LORENZO, J. L. A fluted point from Durango, ology: A summary. Science 156:175?191, 1967. Mexico. Amer. Antiquity 18:394-395, 1953. 7. GRUHN, R. Two early radiocarbon dates from 16. LORENZO, J. L. Dos puntas acanaladas en la thelower levels of Wilson Butte Cave, South region de Chapala, Mexico. Mexico, D. F., Jnstituto Central Idaho. Tebiwa (Idaho State Museum) 8:57, Nacional de Antropologla e Historia, Boletin 18, 1965. 1964, pp. 1-6.

9 15 17. LORENZO, J. L. La etapa Utica en Mexico. tocen-.:o de Tlapacoya, Mexico. Mexico, D. F., Insti- Mexico D. F., Instituto Nacioval de Antropologia tituto Nacional de Antropologia c Historia, Boletin e Historia, 1967. 49 pp.(Depat tamento de Prehis- 29, 1967, pp. 37-41. toria, Publicaci6n 20) 26. MOOSER, F. Tefracronologia de la Cuenca de 18. MACDONALD, G. F. Debert: APaleo-lndian Mexico para los tiltimos treinta mil aiios. Mexico, Site in Central Nova Scotia. Ottawa, NationalMu- D. F.Instituto Nacional dc Antropologia e His- seum of Canada, 1968. 207 pp.(Anthropological toria, Boletin 30, 1967, pp. 12-15. Papers, 16) 27. WAUCHOPE, R. Lost Tribes and Sunken Conti- 19. MACNEISH, R. S. Preliminary archaeological nents: Myth and Method in the Sttulyof the investigations in the Sierra de Tamaulipas, Mexico. American Indian. Chicago, University of Chicago Amer. Philo. Soc. Trans. 48, 1958. Press, 1962. 155 pp. 20. MACNEISH, R. S. Ancient MesoamericanCivi- 28. WENDORP, F. Early man in the New World: lization. Science 143: 531-537, 1964. Problems of migration. Amer. Nat. 100253-270, 21. MARTIN, P. S., and H. E. WRIGHT, JR.,eds. 1966. Pleistocene Extinctions: The Search for a Cause. 29. WEST, F. H., ed. Early Man in the Western New Haven and London, Yale University Press, American Arctic: A Symposium. College, Anthropo- 1967. logical Papers of the University of Alaska 10(2), 22. MARTINEZ DEL Rio, P. Los origenesameri- 1963. canos. 3a. ed. Mexico, D. F.,PAginas del Siglo XX, 30. WEST, R. C. Handbook of MiddleAmerican 1953. Indians, Vol. I. Natural Environment and Early 23. MASON, R. J. The Paleo-Thdian Tradition in Cultures. Austin, University of Texas Press,1964. Eastern North America. Current Anthropology 3: 570 pp. 227-278, 1962. 31. WILLEY, G. R. An introduction toAmerican 24. MAYER-OAKES, W. J. Life, Land andWater. Archaeology. Vol. 1. North and MiddleAmerica. Proceedings of the 1966 Conference on Environ-Englewood Cliffs, New Jersey, Prentice-Hall, 1966. mental Studies of the Glacial Lake' Agassiz Region. 526 pp. Winnipeg, University of Manitoba, 1967, 414 pp. 32. WRIGHT, H. E., JR, and D. G. FREY, eds.The (Occasional Papers, Department of Anthropology, 1) Quaternary of the United States. Princeton, Prince- 25. MIRAMBEI L, L. Excavaciones en un sitio pleis- ton University Press, 1964. 922 pp.

10 16 4, MESES FOR MEDITATION ON THE ORIGIN AND DISTRIBUTION OF MAN IN SOUTH AMERICA

José M. Cruxent

This paper does not constitute an extensiveeven knowing anything about them is a respon- study of the origin and early settlement of sible thing to do. In many cases I think it may South America. So many important works onbe precipitate. I am convinced that a review of the subject have been published that I am re- the old literature in the light of modern research lieved of that duty. I need only cite Hester(10), might be very useful and might lead to surpris- Krieger(12),Comas(3),Lanning and Patter-ing conclusions. son(13),Lynch(14),and Bosch Gimpera(8). Out of the great variety of working hy- Unfortunately, despite all the valuable recent potheses on the origin and dispersion of man in documentation, we still cannot really prove justSouth America, which I consider of little help how South America was peopled. to those concerned with human biology, I find As we all know, concern over the problem ofone positive fact: even today we cannot be sure the origin of American man goes back to thethat man in South America came uniquely by time of Columbus. The conquistadors even land and by way of Panama. wondered whether Indians were human, and so After years of trying to find an answer in it is understandable that Pope Paul III shouldarchaeology and other sciences, I believe that have issued a papal bull explaining that theSouth America was peopled not only overland Indians were in fact human and could receivebut also, and to a large extent, by sea and river the Faith(18). routes, primitive though the means of naviga- Since the sixteenth century numerous hy-tion must have been. I do not see how men potheses about the origin of American mancould otherwise have crossed so many rivers, have been suggested, and most of them haveswamps, and lakes with their entire families been discarded as fantastic. Today we feel that women, children, and the agedto occupy the we have long outlived that era, that -ur work whole of the South American continent. is imbued with a scientific spirit and our hy- If the route was terrestrial, the only entrance potheses are based on valid archaeological docu-is through the Darien jungle in Panama and ments. Nevertheless, I find it difficult to accept the Atrato region in Colombia. I know that area that all this vast ancient documentation is worth-well; I have crossed it on foot from the Pacific less. I say this in full awareness of how difficultto the Atlantic and back again. In this labyrinth itis for a present-day archaeologist to consultthere is no sun, only a weak green light. At and study this copious literature, which not only every stepisa river or a marsh. Whether is hard to find but is sometimes available onlythrough the low ground or along the mountain in archaic languages. But I wonder whethercrests, walking is extremely difficult; hunting prejudging allthose ancient studies without would have been practically impossible. In the 4

"winter" or rainy season no one could haveNormans. Apart from the reasons I mentioned moved. And there are many other Dariens, and above, there is the general discreditto which many rivers to cross. From my travels in South these theories are subject because they have been America I suspect that Pa leo-Indianman mustput forward by untn ined persons guided by have had to go at least part of the way by water ignorance or by a desire to sell books. Thescorn (and I make bold to suggest eaat hemay haveof responsible archaeologists, and their unwill- known how even before he entered the New ingness to risk their prestige by examining them World); if so, this would explainmany of thecritically, is understandable. But thereare some gaps we find when we try to trace his migration.recent finds that if verified would be of great Unfortunately, a great deal of the possible evi-interest. I do not believe that the seriousness of dence has disappeared under risingwater levels such studies as those of Hui-lin Li (11)or since Pa leo-Indian times (10). Sauer (22) can be doubted. The immigrants were unquestionably required One of the old chronicles tells of finding Old to adapt themselves to a great variety of en-World coins in America, and I myself know of vironments, but if they traveled by water thea North American who bought a pot full of cultural changes would be less strong and vio- Roman coins, which seems to have been found lent than if they came only by land, because in a cave, from a farm worker in Falcon State, there would be fewer stages to their journey.Venezuela. The coins are now in the Smith- Genetically, they might have had fewer contacts sonian Institution. I have also seen in the Ja- or mixtures with other migratory groups because maica Museum a pedestal with a Roman inscrip- the continent was so sparsdy populatedat thetion that was found off the Jamaicancoast. I rime. But if we accept the possibility ofa water have heard that a Mexican publication, whk route, we must also accept the possibility that I have not seen, mentionsa small Roman statue some groups reached South America some other unearthed in that country. Thenewspapers are way than through Darien. forever reporting discoveries of ancient objects The "autochthonous"theoryof Americanof European and African origindiscoveries, man, which originated in a misreading of Qua-. however, that are seldom controlledor studied ternary fossil remains and in the belief that by professional archaeologists. I bring all thisup Pa leo-Indian immigration would have been im- merely to inquire whether it isproper for us to possibly difficult, seems now to have beenre- consider such informationa priori false and of jectedamong other reasons because thereis no interest. no convincing evidence to support it. I should Many colleagues have asked me: "If these like to make the point that a good way of laying arrivals did occur, where is the evidence? Where it to rest once and for all would he to prove that does archaeology find any appreciable traces in man came from elsewhere and to prove how he the indigenous cultures?" This is the problem came.I may say in thisrespectthat, even to which we must find an answer, not only from though I cannot prove the contrary, Iam notthe cultural standpoint but also from the bio- at all satisfied with the Bering Strait as the sole logical. entrance route. How could man have spread As to the former, let me say that the cultural over such a vast territory and created such a impact of a small immigrant group on an mosaic of cultures and ethnic groups inso short American aboriginal population is in most cases a time, 20,000 years at most, if he had entered practically nil. We have an example in Balboa's by a single route? siege of Careta: "...the chroniclers agree... I have the impression that not enoughatten- that Balboa found helpful compatriots already tion has been paid to the ancient accounts of the installed there.. . They had adapted them- travels of such navigatorsas the Phoenicians, selves to their new environment without reserve, the Romans, the Arabs, the Vikings, and the and. .all were 'as naked as the Indians and

12 as plump as the caponsthat housewives fatten Once again, therefore, I postulate navigation in their cellars' " (20). This probably happened as a possibility that could havespared the Pa leo- on many other occasions, and notonly at theIndian innumerable obstacles on his way and time of the Conquest. When I was exploringthat furthermore would have permitted migra- the Orinoco Delta region I came upon a French- tory movements without requiring the existence man (he eventually confessed thathe had es- of an ice-free corridor east of the Rocky Moun- caped from Devil'sIsland),living with an tains. Indian wife and mestizo children; it was only by his slightly lighter skin and his heard that IOrigin and antiquity of man in South America was able to identify him. Ihave heard of other All recent studies on the origin of man in examples, and there must be many more. South America agree that migration was chan- As can be seen, the cultural influence of theneled from north to south across the Isthmus of intruder is slight or nonexistent. His biologicalPanama. Evidence of this migration is limited, influence I cannot saythat is up to the biolo- perhaps because there has been very little ex- gists. ploration in this area. So far, no other theory The possibility of immigration by sea is sup- seems to have been confirmed, riot eventhat of ported by the fact that navigation is much easier Mendez Correa (15), who suggests a route than is supposed and much older than is gen-through the Antarctic from Australia to Tierra erally accepted. In the Old World, from pre-del Fuego. As far as 1 know, n) transatlantic historic times until the fifteentl, century, boats routes have been proposed. However, extraordi- hugged the coasts, for the fear of the open sea nary similarities have beenshown between the was great and navigation wasrudimentary. Butstone implements of western Africa andnorth- any storm might pull the boat out to sea,and eastern South America (5). These parallels are then the crew had to resign itself to the currents much closer than between Pa leo-Indian material and the wind. When it is recalled that some offrom Siberia and from North America. the remote Pacific islands were peopled by the As to the antiquity of Pa leo-Indian man time the Europeans came, the logical conclusion using this term for a hunting and gathering cul- that man willingly or unwillingly made longture that coexisted with animals nowextinct journeys across the oceans cannot be avoided. and is known by its cro,tie c.,koppio9toois The most fortunate encountered islands or conti- no accepted chronology yetexists. We nents; some learned in time how to getback may estimate thatthe Pa leo-Indian disappeared again, and others did not. about 7,000 years ago, and very tentatively we Columbus'diaries,citing encounters with may propose an arrival datein South America This .., canoes that traveled theCaribbean from island or Venezuela of 15,000 to 20,000 years ago. to island, support the ideathat sea journeys are latter date is calculated on the basis of finds for possible even in small, primitive boats. We alsowhich the following ages have been obtained: have the many incredible transatlantic voyages El Jobo complex, 10,000 years; Las Lagunas and of sportsmen (16) and contemporary politicalEl Camare, more than 16,000 years; and Muaco, refugees who have brought small, crammed 14,740 to 16,580 years (21). I do not know boats across the ocean in search of freedom andof any earlier dates anywhere else in South a new life in America.I do not believe thatAmerica. Lanning and Patterson (13) estimate Bombard (1) was the first to obtain potable the Chuqui complex in Chile and the Tortuga water from fish; this vital technique maywell and Red Zone complexes in Peru as the most have been known to the ancients. In preceramic ancient in the Pacific Andean region, both being times they could have stored it in light-weight13,000 to 14,000 years old. For Argentina we vessels such as gourds or skins, which whenhave a date of 7,970:I:10t)at Ayampitin- empty could have been used asfloaters. Int; huasi, and for Brazil a date of 7,501 at

13 19 Sambaqui de Maratua(18).Current excavationswhich 1 accept for both Meso-Indian and Neo- in Chile showing man in association with masto- Indian times. I may cite herepurely for refer- dons and a variety of other animals promise to ence purposes, since I am not sure of their produce extremelyimportantdocumentation validity for the CaribbeanHester's observations (2). (10)on the rising of the sea level since Paleo- From these and a few other dates that might Indian times. Many islands would be intercon- be mentioned, we can make the following ten- nected, thus making navigation and migration tative timetable for Pa leo-Indian man in Amer- easier. The C-14 ages of some Meso-Indian sites ica: North America, 15,000 to 30,000 years (9); support the idea of inter-Antillean navigation: northern South America, 10,000 to 20,000 years; Rooi-Rincon, Curacao, 4490-160 years; Mor- western and southern South America, 8,000 todan, Dominican Republic, 4120 ± 160 years; 14,000 years. The figure for North America Punta Gorda, Venezuela, 4150 ± 80 years. deserves a question mark; the experience of As to Paleo-Inclian man in the Caribbean, at South America leads one to believe that further the Mordan site we have found a more ancient studies will uncover earlier dates(13).Indeed, level of chipped-flint implements, which we although it is quite possible that some specialists have called Alejandrina (Cruxent and Chanlatte, may consider these dates too ancient, I find them work in progress). The ypology of the Alejan- not only logical but overly conservative and I drina complex can be r Aated to that of the expect future studies to at least confirm them. El Camare complex,.-.-;.11large plano-convex knives, scrapers, chisels, and other tools. This Principal lithic complexes material has not yet been fully studied. The Lanning and Patterson(13)have proposed a artifacts are made of excellent-quality flint and three-group typology for the South American are in strict accord with the typology of flint lithic complexes:(1)bifacialindustries, (2) industries, which is based in part on the nature burin or graver industries, and (3) chopper in- of the substance. The same is true of the Man- dustries. An examination of the principal Vene- zanillo and El Camare implements, the first of zuelan complexes shows that the typology ofwhich are of silicified wood and the second of that country presents certain characteristics that quartzite. I am one of those who believe thata are not present in those of Argentina, Chile,single group of artisans trying to manufacture Peru, and Ecuador. Earlier this year, when he a certain type of instrument with different ma- consulted our collectionsin the Venezuelan terials would have to produce different typolo- Institute for Scientific Research, Dr. Lanninggies. This must be taken into account in com- was able to relate the Camare complex with the paring complexes, for it is easy to make mistakes. "Bifacial Andean Horizon" (large chipped arti-There is also the possibility of the fortuitous facts, large bifacial stones, absence of projectileuse of objects as implements in certain circum- points). He could not relate the Chuoui, Tor- stances. This may be the answer in the case of tuga, Red Zone, or Exacto (Ecuador) complexes Taima-Taima, whichiscompletelyatypical; with those of northeastern South America, and on the other hand, it is just as likely that we call he found some of te Lagunas and El Job° it atypical because this is the first time we have material (plano-convex scrapers, spear points, seen such implements in association with Pleisto- and other articles) to be quite unlike anything cene fauna. in the regions he had studied. The presence of Paleo-Indian culture on the To me this circumstance is extremely impor-strategic island of Hispaniola raises a series of tant. It permits a working hypothesis as to the problems. The principal one is how the Ale- existence of another possible entrance route tojandrina people lived, for no fossil remains South America: the chain of Caribbean islandsof mastodons, glyptodonts, or megatheres have from Florida to northeastern South America, been found. Considering that they were island-

14 dwellers, we must conclude, if they were greatposition of having no erudite citations tomake hunters, that if large animals were not to bein support of my arguments.) From thisexperi- found on land they may well have beenfound ence, I do not think we candiscard the possi- at seaperhaps manateeand seals, both ofbility that there were Palto-Indian groups with- which were abundant in the Caribbean in those out stone implements any morethan we can times (7 , 23). ignore the likelihood that they knew how to Yothing is known of the origin of the Paleo-navigate. Indians of the Alejandrina complex, but they I also think it a valid working hypothesis that probably came from the north. Nor is it knownPaleo-Indians without a lithic culture, which I whether they reached Venezuela, though some call "primary culture groups," could wellhave slight contact between the Alejandrina complex,been the ancestors of certain groups offorest or an older one ofthe same sort, and the Elgatherers and primitive agriculturists in South Camare and Las Lagunas complexes would not America. Lithic culture evolved through aseries have been impossible. Later groups may alsoof phases before arriving at triangular peduncu- have had some influence on the early Meso- fated points; it is significant that this tradition Indian or late Paleo-Indian complex of Canaimasurvives to the present day in our llanos and in Venezuelan Guiana (21), the Rio Claro com-savannas but that Ihave not found it in jungle plex in Brazil (4), and the Gran Sabanaand or thickly forested areas. Paragua material. This could indicate that a diversity ofdiets I hope in the near future to be able to clear existed from the start or else that it cameabout up this problem of another route toeastern at the end of Paleo-Indianand the beginning South America in Paleo-Indiantimes. This of Meso-Indian times, when the greatvertebrates would extend the validity of the H theory pro-disappeared. Unfortunately, no conclusion can posed by Osgood and Howard (17) as a graphic be reached at present, for we do notknow explanation of migratory routes in general. whether the jungle-dwellers descended from an- Before concluding, 1 wish to point out thatofaer group of great hunters or from a group many times we fail totake into account a hy- that entered the New World alreadycarrying a pothesis that should not be ignored indrawing primary culture. Itis also possible that some conclusions. I am referring here to the possibility groups abandonedthe flake industry in adapta- that there may have been Paleo-Indian groups tion to an ecological area that did notallow its without a lithic industry. 1 do not understanduse. Geography is notdetermining, but it un- why we always imagine Paleo-Indian manwith doubtedly affects many aspects ofthe life of their habitat a stoneimplement in his hand; we are even primitive peoples, who obtain from presumptuous enough todemand a specific type the elements essential to theirneeds and their of implement, an officially accepted one.We security. must not forget that until veryrecently projec- In conclusion, I believe that many ofthe gaps tile points were the only implements recognized; we observe inPaleo-Indian migration can be all others were considered "blanks." For many logically explained by taking navigationinto years 1 have beenaccumulating experience with account. I do not rejectthe idea of terrestrial In an archaeologist's eyein many Indian commu- migrations; I wish only to complement it. nities. This in vivo archaeology has given me any case, I amoptimistic about the possibility many answers that wouldhave been hard toof solving the problem of theorigin and dis- come by from purelyacademic studies. (Of persion of man in South America,because most course, this often puts me inthe unfortunategood archaeologists do not regard it assettled.

15

, 21 REFERENCES 1. BOMBARD, A. Naufrage Volontaire. Paris, 1958. Columbian transatlantic travel by Arab ships. Har- 2. CASAMIQUELA, J. MONTANE M.- and vard 1. Asiatic Studies 23, 1961. R. SANTANA A. Convivencia del hombrecon el 12. KRIEGER, A. D. Early man in the New World. mastodonte en Chile central. Mus. Nac, Hist. Nat. In J. D. Jennings and E. Norbeck (eds.), Prehistoric Noticiario Mensual (Santiago) XI:132, 1967. Man in the New World. Houston, Texas, Rice Uni- 3. COMAS, J. Los problemas de la antropologia versity Press, 1964, pp. 23-81. fisica ante el origen del hombre americana. SID 13. LANNING, E. P., and T. C. PATTERSON. Early Paulo, Encontros Intelectuais, 1961. man in South America. Scientif. Amer. 217(5):44- 4. CRUXENT, J. M. Noticia sobre litos de silex del 50, 1967. Brasil. Bol. Mus. Cienc Nat. (Caracas) IV, 1958; 14. LYNCH, T. F. The nature of the Central V, 1959. Andean precerarnic. Occasional Pipers Idaho State 5. CRUXENT, J. M. Les Cultures lithiques Pa leo- Univ. Mus. 21:78-82, 1967. Indiennes du Venezuela: un curieux phenombse de 15. MENDEZ CORREA, A. A. Nouvelle hypothese convergence typologique es fonctionnelle avec lessur le peuplement primitif de PAmerique du Sud. cultures foresteres centrafricaines.Tervuren Bel- Anais Fac. Sociologia do Porto XV, 1928. gique. Koninklijk Museum voor Midden-Africa, 16. MERRIEN, J. Les Navigateurs Solitaires. Paris, 1960. 1957. 6. CRUXENT, J.M., and I. ROUSE. Arqueologia 17. OSGOOD, C., and G. D. HOWARD. An Archeo- logical Survey of Vene.zuda. New Haven, Yale Uni- cronológica de Venezuela.2vols.Washington, Unión Panamericana, 1961. versity Press. (Publications in Anthropology, 27) 18. PERICOT y GARCIA, L. 7. EnciclopediaUniversal America indigena. IlustradaEuropeo- Tomo I. El hombre americano. Los pueblos de Americana. Barcelona, Espasa, 1924, Vol. 24, p. 218. America. 2a ed. Barcelona, 1962. 8. CIMPERA, B. P. L'Arn;:rique: Paliolithiqueet 19. REMAN, E. The Norse Discoveries and Ex- misnrshique. In Andre Varagnac, Paris, L'Homme plorations in America. Berkeley, University of Cali- avant Picriture, 1959, pp. 165-187. fornia Press, 1949. 9. GRIFFIN, J.B. The origin and dispersion of 20. Romou, K. Balbo.of Darien: Discoverer of American Indians iTI North America. In Biomedicalthe Pacific. Garden City, New York, Doubleday, Challenges Presented by Me AmericanIndian. 1953, p. 94. Washington, Pan American Health Organization, 21. ROUSE, I., and J. M. CRUXENT. Arqueologia 1968, pp. 2-10. Venezolana. Caracas, Ed, 1,/e..a., 2962. 10. HESTER, J.J. Late Pleistocene environments 22. SAUER, C. 0. Neblina norteiia. Berkeley, 1968. and early man in South America. Amer. Naturalist 23. SIMPSON, G. G. The principles of classification 100:914, 1966. and a classification of mammals. Bull. Amer. Mac. I I. Hui-LINLi.Mu-Ian-Pi: A caseforpre- Nat. Hise. 85, 1945.

16 DISCUSSION

William S. Laughlin

I feel a little like an Eskimo shaman coming think, the advantage of a richer ecological zone. before the College of Cardinals to discuss theThey had the marine mammals. They had the doctrine of the Immaculate Conception. But an fish. And, of course, they had the birds, as well Eskimo shaman would not hesitate to do this,as those things that grow in theintertidal zone, and so neither will I. and, very importantly, invertebrates, mollusks,.1 octopus., and so forth.hts cAptateiiNiciS C35111 A great deal of ground has been covered verybccessibe 44warm's.ech t,NetWte expertly by both these scholars, and they have I suspect that the ancestors of the Aleuts (of raised a number of very good problems. I canthe Aleutian Islands) and the Eskimos (of the make only a few comments and perhaps add onemainland extending over to Greenland) may or two olrervations by way ofinterpretation. have originated or taken their primary form on I would suggest that most or all of the peoplethe southern part of the Bering land bridge, who entered the New World came by way of the and that when the land became submerged they Bering land bridge. This, of course, does notsimply backed up and remained on the coast. exclude other points of entry. But at least theOf course, as the waters rose there was more

other groups that"Vave come L have left nocoastline rather than less, and therefore they distinguishable genetic heritagethat can be prospered rather than disappeared. sorted out. The ancestors of the Paleo-Indians, who more As Professor Griffin pointed out, the Bering likely came through the interior, may have never land bridge was a large extent, over 1,000 kilom-seen the people on the coast, beingseparated by eters from north to south. This leaves, then, anseveial hundreds of miles, or have had only opportunity for at least two routes across the intermittent contacts with them. They were big- land bridge and may help to explain some ofgame hunters hunting mammoth,caribou, and the differentiation between the less Mongoloidpossibly musk-ox. These people may have lived American Indians and the more Mongoloid Es-on the Bering landbridge for some time, but kimos and Aleuts of the North. when the waters rose they had only two options. It is very likely that some of the early peopleOne was to learn to live under water; the other lived permanently on the land bridge. Theywas to go back to Siberia or on tothe New were there for a long periodof time. They didWorld. Obviously, a number of them went on not simply run across in order to get tothe to the New World. other side in order to be discovered by people Now, when we look at the early sites, we find who came later. And for all practical purposes very good documentation for theperiod between they were unamwe that they were going to be9,000 and 12,000 years ago. There are a great submerged. So for a few thousand years theymany dates that I think can beaccepted as being enjoyed the real estate they held. well rep!icated. But getting beyond the 12,000- Now, those who lived on the coast had, I year barrier is difficult.

17 The most recent piece of evidence isone thatChina, as was originally observed by Franz Dr. T. Dale Stewart called to my attention Weidenreich. This similarity probably indicates yesterday. Dr. Leakey has rediscoveredan old some genetic continuity. If that is), this again discovered skull in Southern California,at La- reflects a rather minor amount of external visible guna Beach, which has been dated on the bone morphological change and suggestsa slow rate itselfnot on associated materialsat 17,000 of morphological evolution. years. The skull has not arrived here yet, and One interesting point we must consider then we do not know what it looks like. is the possibility that the morphological diver- In any event, if that date should be substan-gence between peoples within the New World tiated, we would be ina position to say thatand between the inhabitants of the New World man had to come prior to the 14,000- or 15,000- and the Old may reflecta slow rate of human year period and must therefore have come before evolution, morphologically speaking. Therefore, 24,000 years ago. This, however, has not yet the fact that some of the earliest physical remains been established. do not took like Neanderthal manor like Now, turning to South America, one thingAustralians does not mean that they have not I should like to mention is the trait complexbeen here for some time. associated with the Lag& Santa crania of Brazil. And there is something else to be emphasized You will recall that these were discovered in thethat the amount of morphological change does last century; they are now in the Museum of not accurately reflect that of other tissue systems. Zoology in Copenhagen. Serological evolution, dental evolution, changes Lag& Santa crania are dated around 8760B.C. in different tissue systems, are proceedingat The interesting thing isthat although they different rates. When we look to the past,we are appear to be this old, they look like those of limited to the skeleton; we cannotassume that contemporary Indians, and not only in general change in many genetic traits, including behav- features of morphology but also in discontinuous ioral traits, has not gone on at perhapsa much traits that may be more similar to single-gene faster rate. traits, though this is not firmly demonstrated. The other point I should like to make is the They have shovel-shaped incisors, whichare fact that the majority of the people coming found in all New World occupants, both Eskimo across Bering Strait were exposed to a cold and Indian. They have dehiscences of thetym- filter. All botanists, glaciologists, geologists, and panic plate. And there are severalcases of inter- anthropologists are agreed that, whatever condi- parietal bones, the separate Inca boneon thetions were, they were cold. The inhabitants posterior of the skull, which reachesa highadapted to the cold for the fewor several frequency in South American Indians. thousand years they livedon the Bering land Taking this example to raisea point, it would bridge. This may possibly be related to some appear that there has been very little morpho-of the epidemiological and immunological as- logical change in American Indiansover a period pects of contemporary populations. Once they of 9,000 years. got south, of course, they could adapt to several We can say the sar..e thing for North Ameri- other varieties of zones. But they all had topass can, though we do not have as good a series as through a cold filter. for Lag 6a Santa. Nevertheless, theamount of The last point I should like to mention is the morphological evolution does notappear to have rate of dispersion and how long it would have been great. taken to get to the Strait of Magellan. I simply We can go one step further and pointout that call your attention to the fact that the Aleuts the Eskimos and Aleuts, who forma single and Eskimos are spread over a difficult expanse coherent population system, demonstratea simi-of 8,000 miles, and it took them considerably larityto Middle Pleistocene man of northern less than 8,000 yearspossibly as littleas 2,000

18 yearsto ga that way. Therefore, the trip toWhat I asked is this: You seem to give the South America may Rot have been as long as impression that in North America we find older we might otherwise think. ages than in Mesoamerica,and in turn the ages in Mesoamerica are greater than those in South America. Could this be taken as additional evi- General Discussion dence for a north-to-south migration? Salzano: Since Dr. Laughlin mentioned the Cruxent: Yes, it seems to confirm the theory Lag Oa Santa material,I want to say a few that the migratory current was from north to words about the prehistoric remains of Brazilian sou th. Indians. Moderator: Professor Griffin, you have a name A few years ago a researcher from the Na-for being a bit of a skeptic in these matters. tional Museum, Professor Marilia de M. Alvin,Would you care to respond at all to Dr. Crux- did a review of the anthropometric data at hand ent's presentation? and found a considerable heterogeneity between Griffin: The evidence of man in South the different Lagoa Santa sites. We cannot say, America, both what can be established and what therefore, that these people are just one homo- has been claimed, indicates an antiquity of man examined the shell- geneous group. She also in South America greater than that in North mound material of different locations and they America, and evidence in which I have some were even more heterogeneous. confidence suggests that we have not found man Moderator: Do you mean to imply that the in North America of the antiquity necessary for heterogeneity in the Lagoa Santa material is his appearance in South America at the age greater than might be found in contemporary claimed. I think itis possible that we simply Indian populations in Brazil? have not found the evidence in North America. Salzano: That is difficult to say, because the I should like to comment on one point of material available for study is not very extensive only about 80-odd individuals. But even inCruxent's paper: If through man's penchant for self-destruction the evidence for our astronauts this small sample the heterogeneity is very great. there would be very I am not saying, however, that it is greater than were completely destroyed, little in the earth's atmosphere to tell the people among present-day Indians. of the future that man had actually gone around Roche: I have a question for Professor Crux- the world. Cruxent's "Paleonauts," if I may ent. He gives a possible ai;e for man in North coin a phrase, are extremely difficult to trace in America of from 15,000 to 30,000 years, in the sea waters. So that while he hinted broadly western and southern South America of 8,000 at the possibility of man's making voyagesfrom to 14,000 years, and in northern South America western Europe or western Africa and perhaps of 10,000 to 20,000 years. These ages seem to across thc Pacific, myposition is that he may less. Does he consider this as go from more to have done so but I have seen no traces of the north-to-south migration? one more proof of boats, or whatever they used, and thus have Cruxent: I do not think the last word has yet no evidence for such aninterpretation. been said as to ages. We have not been able to date the oldest complexes we have in South I also wish to comment on the presumed con- European, African, or America, in the alluvial terraces at great alti- tact of Mediterranean, tudes, because all the material is on the surface relatively civilized Asiatics with the New World. and has been exposed to the air. It happens inSuch ideas have been current for a long time. some places in the States.But I feel that little They were originally based on the philosophical by little, here in the States, earlier ages will be position that it would have been impossible for found. We keep finding older and older ma- Indians to achieve high cultures without benefit terial in America, and I think that will continue. from groups that had achieved reasonably high Roche: That was not exactly my question.cultural status in the Old World. But that posi-

19 tion has been abandoned. We no longer need being the rather small Eskimo type and the to bring high culture into the New World fromother the North American type. Is thereany the Old because now we can clearly documentevidence that they came fromtwo different the gradual development of the higher civiliza-families or groups in central Asia? tions in the New World, both in South America Laughlin: I think the answer isno, there is and in Mesoamerica. no evidence of two separate groups that can be It is true that a great many contact areas aredisting.,ished today. However, the recordin known. For example, there are stories of thenorth China and Japan doessuggest that evolu- Irish being present in North America, primarily tion has been proceeding there and that people around the hinterland of the Boston area. Phoe-would differ according to when they sampled nicians are recognized in eastern Pennsylvania that particular area. around Lebanon. A major center for pre-Colum- So the fact that the more recent (if indeed bian Scandinavian influence in North America they are more recent) Eskim-)s and Aleuts look is in the area of Minnesota and Wisconsin. Butmore like Japanese and Chinese may reflect some of us are extremely skeptical about theeither their recency or the fact that theywere appearance of Roman coins or statues or various drawn up the no-th Pacific riinJapan, the other objects, because they can be reasonablyKuriles, Kamchatkaand followedtizecoast explained on the basis of man's pack-rat and rather than coming from northern Sibefia. souvenir complex. There is no particular group that could be And so I admit to somewhat of a skepticalidentified today, and I think it would be haz- position on many of these matters that Dr. ardous to look for one, since those populations Cruxent has so delightfully called to our atten-have been evolving for the same period of time tion. as those in the New World. McDermott: Dr. Griffin, is there any linguistic But there is, as Dr. Comas and others have evidence acceptable to reasonable man,as you pointed out, a major dichotomy between the put it, on this question of migration from North Aleuts and the Eskimos and the rest of the to South America? Indians. This includes several serological factors Griffin: I am told there was tremendous lin- as well as growth rates, disease patterns, and guistic diversity among the American Indian other morphological differences. groups and that none of the New World lan- I would add just one other thing, and that guage stocks can be traced back to those of Asiais that in our views we should take intoaccount with any degree of certainty. I am also told that that Japanese anthropologists have again pointed the linguisticdiversityin,for example, theout that they think the Ainu of Hokkaido, in eastern United States could reasonably havenorthern Japan, are a variant of Mongoloids taken place within a period of some 10,000 toand not of archaic whites who were surrounded. 12,000 years from a single ancient linguistic The old thesis, originally proposed by Professor stock. Birdsell, that there was a group of Australoids, The archaeological evidence we have indicates archaic whites, or Ainu-like people living in to some of us that the original populations of Siberia is no longer tenable if as much hetero- the eastern United States were relatively homo- geneity as that between Ainu and Japanese can geneous in cultural material and, further, that be recognized among Mongoloids. cultural extremes can best be explained primarily Griffin: If our archaeological evidence is any as regional developments based on the original good, then the first migrations seem to have been cultural groups. from the central Siberian area, with cultural Water low: Dr. Laughlin suggested thattwo complexes that date back somewhere around groups of people came over by different routes20,000 years or so. These populations seem to northern and southernthrough Alaska,one have moved primarily along the northern coastal

20

26 areas of Beringea into theNorth Americanprimitive music shows much less variation and interior. The evidence is that the populationsevolution than primitive languages, that origins Dr. Laughlin has been talking about with hiscan be recognized muchbetter through music proto-Eskimo or prehistoric Eskimo of 8,000than through language. Practically nothing has years ago or more aredistinctly different from been done in this area. For what it is worth, he the earliest movers into the New World andfeels that the Latin American music he has had a cultural complex formulated primarilyheard, particularly the very primitive, such as on the Pacific side ofnorthern Asia, at a later that of the Caribbean, has very strong relation- time. It was chiefly the first group, moving intoships with northern Asia, particularly Japan. the New World along the northern rim, that Moderator; At this point we have succeeded populated North America and I think movedin getting the Indian to the Americas,although into South America, either on migratory-birdthere is something less than unanimity as to wings, across water by navigation, or ploddinghow this was accomplished. In the next section through and slowly adapting to various environ-of the program we propose to look at the ments. physical and genetic diversity present inthe Roche: I want to make just a brief comment Indian.Thedistinctionbetween in relation to Dr. McDermott's question. There American have been studies on linguistics, but there have "physical" and "genetic" is of course artificial, been very few comparative studies on the music since the basis of much physical diversityis of the Indians. I have been told by the headof genetic. It is a convenient distinction,however, the Institute for Music Study in Berlin that and we shall adhere to it today.

2127 BIOLOGICAL SUBDIVISIONS OF THE INDIAN ON THE BASIS OF PHYSICAL ANTHROPOLOGY

Juan Comas

In accepting the invitation to participate intheir color and make are so nearly alike," and this Special Session, I was guided primarily by the consequent acceptance of the somatic unity the great interest this subject has always hadof the pre-Columbian population of the New for me. I well realize, however, that my knowl-World as a definitive fact, received the support edge of the problem and my information, both of such renowned anthropologists as Samuel G. direct and indirect, leave much to be desired.Morton (1842), Timothy Flint (1826), Ales I present my excuses in advance and ask forHrdlicka (1912), and Arthur Keith (1948). leniency on the part of my readers. The opposite campcomposed of those who For a better understanding of the vah1/4: and recognized the existence of obvious biological scope of the principal biological subdivisions ofdifferences among the Indian groups, describing the Indian on the basis of physical anthropology, them as "races," "varieties," or "sub-species"--in- I believe it necessary to first pose three questions eluded, among many others, Humboldt (1811), and go on from there to some provisional con- Desmoulins (1826), D'Orbigny (1839), Retzius clusions. The questions are these: (1842), Aitken Meigs (1866), Topinard (1878), 1. Do the American Indians constitute a bio-Deniker (1889), Virchow (1890), Ten Kate logically homogeneous population or have they, (1892), Haddon (1909), Biasutti (1912), Wisier on the contrary, a certain variability which per, (1922) Dixon (1923), Rivet (1924), Eickstedt mits thinking in terms of subdivisions? (1934), Hooton (1937), Imbelloni (1937-58), 2. To what cause or causes may the biologicalCount (1939), Neumann (1952), and Schwi- variability of the American Indian be attributed? detzky (1952).1 3. What attempts have been made at a racial It is interesting to point out that, while the taxonomy of the American Indian? believers in the somatic unity of the Indian re- Let us examine these questions consecutively. mained a minority, in -heir forefront was Mor- ton, of whom Stewa t and Newman(56) say Biological variability of American aborigines as shrewdly: against classical conception of "American so was hisinfluencethathewas homotype" Indeed, great responsible in large measure for the wide acceptance of The excellent works of Newman (47) and the generalization embodied in Ul loa's words and for Stewart and Newman (56) make it unnecessary the conversion of Ulloa's words into an "adage." (p. 22) to devote too much space to this question. Dis- Morton's influence lasted for more than half crepancies on this point have a long history. Antonio de Ulloa's statement toward the end of 3 Most of these arc well-known general works and therefore do not appear in the References. For additional the eighteenth century that "If we have seen information see J. Comas, Manual of Physical Anthro- one American, we may be said to have seen all, pology, (Springfield, 1960), pp. 81-86 and 588-600.

22 Ak,d't ,45

a century, until Hrdlicka arose as the new had been definitely discredited. While the great champion of the homogeneity of the Amerin- majority of anthropologists are of this opinion, dian. In summarizing his arguments Hrdlickawe must not overlook the fact that some dis- said in 1912(32):"The conclusions are thattinguished investigators differ. Ashley Montagu the American natives represent in the main a (44), for example, has written: single stem or strain of people, onehomotype." The American Indians exhibit a certain basic homogeneity He reaffirmed this thesis in 1928(33) inan but at the same time are evidently characterized by an attempt to refute those who supported the racial equally certain diversity of types. Owing to the lack of plurality of the American Indian: the necessary data itisimpossible to say quite how many diverse types there may be. General impressions We find that the various differences presented by the based on sporadically measured and photographed indi- Indians are often more apparent than real; that actual viduals from various groups provide an insufficient basis and important differences arc in no case of sufficient upon which to erect a satisfactory account of the Ameri- weight to permit of any radicaldissociation on that can Indian. (p. 465) basis. (p. 481) This position had the able support of SirSome years later the same author does affirm Arthur Keith(36): (45)that the Eskimos of the Arctic coast of North America form part of the group of Arctic Certainlythe American Indian differsin appearance from tribe to tribe and from region to region, but un- Mongoloids while the other American Indians derneath these local differences there is a fundamental constitute another group composed of "an unde- similarity.This, too,is in faveur of descent from a termined number of ethnic groups of North, single, small, ancestral community. (p. 218) Middle, Central and South America." Nevertheless, little by little, the physical varia- Coon, in his important volume on human bility of the American Indian had shown itself races(18)devotes barely two pages to the racial to be an undeniable fact, leading to differing characteristics of the American Indians. He says: and even contradictory descriptions and systema- The American Indians are more uniform racially than tizations. Laughlin(38)presents the situation any other group of people occupying an equally vast very clearly and objectively: arca. In fact, they are more uniform than many peoples Much progress has been made since the early days of who occupy an arca a tenth as large. All of this indicates that a relatively small number of peoples crossed the anthropology in America when it was assumed that all Bering Strait during thelast part of the Wisconsin Indians were essentially alike. The diversity of the New glaciation, and that their descendants gradually filled the World populations has been well established. The sig- uninhabited regions of the New World. They are Mon- nificance of the diversity in terms of the evolution or development of types there or of the importation of goloid in general and despite some of their peculiarities preformed types from the Old World remains to be in blood groups do not necessarily merit classification as a subspecies of their own. . The American Indians clarified. (p. v) differ from the Asiatic Mongoloids mainly in that they Stewart and Newman (56) expressed the same have less facial flatness, particularly in the nasal skeleton, opinion at the conclusion of their article: and a more variable skin color. There is no valid evidence that the Indians were derived from more than one source This review of opinions regarding Indian variability has or that they came into the New World by a route other revealed thefact that the principle of Indian raitil than the Bering Strait. (p. 152) unity rests almost solely upon the outer appearance of living Indians. In so far as the Indians exhibit in com- In spite of these sporadic casesand quite mon such physical characters as straight, black hair, possibly there are other adherents of this theory copper-colored skin, dark brown eyes, high cheek bones, scarity beard and a relatively long trunk, they can be I believe it can be said that the beginning of 'a7 said to be uniform. ... That the Indians, on the other the second half of the twentieth century coin- hand, are quite variable within this racial pattern, and cides with an end to the myth of the "Indian especially when comparisons are made inmeasurable homotype" and an explicit recognition by the dimensions, also has been shown. (p. 33) great majority of anthropologists of the exis- All this would lead one to think that fromtence of somatic and osteological variability and 1951 on, the theory of the American homotype heterogeneity among the aboriginal groups of

23 America. It is then necessary to establish theto localize "the sectors and zones where the origin and the causes of this variability, bearingeffects of mixture and hybridization occurred," in mind that it in no way exceeds the limits ofalthough he rejects every explanation for what characterization of Homo sapiens. he called the "environmentalist creed." Have we sufficient and adequate information, Second, there is the theory of somatic varia- posterior to 1951, to be able to determine whether bility as principally the consequence of environ- the differences of physical types among themental influences, which Stewart and Newman Amerindians are due to prehistoric immigrationexplained as follows: from the Old World or to adaptive evolution to Interpretations of this sort were made largely by Ameri- the new habitat? This is what we mwt try to canists who had no classificationsto justify, and ac- analyze. cordingly were more willing to admit that anthropometric determinationswerenot alwaysstable in changing environments. As a group, these Americanists did not Cause or causes of biological variability of deny the migrationist postulates of the classifiers, but American Indians seemed to believe that hereditarian and environmental explanations could be harmoniously blended in overall At first glance it seems easy to distinguish two interpretations. (p. 31) explanations for this phenomenon. First, there is the proposal of those who accept But what seems essential to me is a somewhat the immigration of diverse human types, eachmore careful analysis of Newman's own view of which represents one of the existing Amerin- (54), which he expresses thus: dian "races." Stewart and Newman (56) de- The adaptive responses of bodily form to environment scribe it very clearly, attributing it primarily toin warm-blooded animals have led, largely in the last century, to the formulation of several ecological rules. those who have been concerned with American . Extensive testing of these ruks on human materials, racial taxonomy: however, has not been performed. For this reason,I have examined the applicability of two of the best vali- ...theclassifiersof Indiantypes operated with a dated of these rulesBergmann's and Allen'sto the strongly hereditarian bias, out of keeping with the main body forms of Ncw Wodd aborigines.2 The principle stream of biological thought. If explanations were given behind both rules is that the maximum retention of body for their dassthcations, they were usually to the effect heat in cold climates occurs when the radiating skin that each Indian "race" represented a separate migration surface is small relative to body mass. Since this ratio from the Old World. . ImpliAy, such explanations can bc achieved by larger body size, Bergmann's rule disavowed the possibilitythat physical changes could holds that within a widc-ranging species, the subspecies have occurred among New World peoples. (p. 29) incolder climates attaingreatersizethan those in However, they themselves indicate that the warmer climates. Allen's rule holds that in addition the various authors place different degrees of empha- cold climate subspecies have reduced extremities and sis on the hereditary aspects of the prehistoric appendages, thus further reducing the body surface. In warmer climates, following Bergmann's rule, easier dis- immigrants, on cross-breeding with each other, sipation of body heat goes with the low body mass body and on the environmental influence of the new surface ratio achieved by smaller body size (p. 324) habitat to explain the presence and existence of Newman presents with the greatest objectivits distinct Amerindian "races"; and they evaluate a series of facts, some of which I wish to men- the greater or lesser importance thatsome of the tion because his interpretation of them seems to most conspicuous "polyracialists"for example, me in certain cases to be doubtful, erroneous, Dixon, Griffith Taylor, and Hootonconcede to and even contradictory with respect to the final each of these factors (p. 30). Even Imbelloni, comment. one of the most devoted defenders of a compli- 1. He states (p. 312): "In mammals and birds cated polyracialism, refers to only seven distinct migratory waves and on the other hand describes 2 j. A. Allen, The influence of physical conditions in and localizes eleven Amerindian "races," which the genesis of species, Radical Revicle 1:108-140, 1877. implies agreement that in the new habitatnew C. Bergmann, Ueber die Verhiiltnisse der Wiirmebko- nomic der Thicre zu ihrer Grösse, Glitiinger Suidien racial types were formed. Heeven tries (34) 3:595-708, 1847.

30 there are a number of exceptions to these rules: their shorter stature, then, is attributable solely It) to 30 per cent for Bergmann's rule, calculated to their short legs. This reduction of extremity only from subspecies in the most contrastinglength is in accordance with Allen's rule, and climates of the species range (Rensch, 1938: probably [italics mine] represents an adaptation 282)." Actually, the percentage of exceptions tofostering body-heat retention." As to the eastern Bergmann's rule among birds and mammals isEskimos, whose stature and sitting height are larger thanthis. Rensch (51) mentions "forless than those of the Indians who live further palearctic and nearctic birds I calculated 20-30% to the south, he attempts to explain this by of exceptions on the average" and "for pale- saying "Possibly [italics mine] the use of heavy arctic and nearctic mammals 30-40%." tailored clothing in combination with factors of 2. He says (p. 313): "I am aware that inuncertain food supply and periodic undernutri- certain parts of the Old World [Bergmann's and tion may cancel out the selective adv antage of Allen's rules] do not seem applicable. Upon larger bodies in colder climates, but his cannot superficial examination, the rules do not appear be demonstrated." operative in Africa south of the Sahara. Yet, in Continuing to note exceptions to the sup- Europe and the Near and Middle East and in posed applicability of Bergmann's rule to man in East Asia and Malaysia, there seem to be north-America, he mentions groups of small stature south body size clines conforming to Bergmann's "surrounded by taller groups," such as the Yuki rule. The explanation of these discrepancies in of northern California, the Lillouet of southern Africa and perhaps elsewhereis not yet ap- British Columbia, and the Yahgan and Alakaluf parent." To admit that Bergmann's rule is notof the Magellan archipelago. With respect to the operative in Africa south of the Sahara indicateslast two groups he further indicates that the the impossibility of generalizing such a rule in sitting height has not been calculated, "but other support of human variability. Furthermore, itsmeasurements indicate they are not particularly application seems doubtful in Europe, the Nearshortlegged." and Middle East, eastern Asia, and Malaysia. Neel and Schull (46) say: "In the simplest How does one explain, for example, the differ- terms we may conceive of stature asbeing the ences of stature and body proportions indiffer- cumulative effect of a number of genes whose ent regions of Europe, and among theVeddas actions are similar and whose effects are addi- and Brahmans of Bengal, the Sikhs of the Pun-tive" (p. 107). "We may surmise, therefore, that jab, the Tapiros and Papuans? under the environmental conditions in which 3. In support of his thesis Newman includesthis study was conducted, the primary cause of n;ne maps showing the distribution in Americ. variation in stature is genetic" (p. 110). of stature,sitting height (living), head size Barnicot (9), referring to geographical varia- (living), cranial module (skeleton), upper face tions in stature, says: size (skeleton), morphological facial index (liv- There is a great deal of information about stature varia- ing), upper facial index (skeleton), and nasal tion throughout the world, but not infrequently itis index (living and skeleton); but he adds (p. based on samples which are either very small or were 315): "The categories used in these maps are selected in a way which may render them unrepresenta- tive of the general population. ..On the whole the arbitrary, and in some areas the data are inade- pattern of stature variation throughout theworld shows quate, but probably[italics mine] the maps no very striking regularities. Bothtall and short peoples represent reasonable approaches to the real dis- are to be found in most of themajor regions. (pp. 203-- tributions." 204) Then he notes that the small stature of the And after specifying different cases of geographi- Eskimos constitutes an exception to Bergmann's cal distribution of stature, he concludes: rule but that, as "the Western Eskimo are not This distribution of stature has been interpreted as an inferior in sitting height to the tallest Indians, example of a dine with adaptive significance in relation

25 31 to climate (Bergmann's rule). A substantial negative build is influenced by both hereditary and direct correlation between body-weight and mean annual tem- environmental factors." And "without denying perature has been demonstrated for various regions of the world. that the New World was peopled by successive migrations or infiltrations of physically differing Harrison (27) also writes, in dealing withpeoples it is very likely that the Americanraces environmental interaction: of the classifiers are at least partly the products Differences in stature are inherited, but they are also of adaptive changes that took place in the New environme.ntally determined, since grrwth is profoundly World." This, generally speaking,seems accept- affected by nutritional state, and probably by climatic able; I have earlier made observationson this factors as well. Further, the nature of the variation pro- duced by both types of factor tend to he the same. matter, with particular reference to the charac- (pp. 144-145) teristics that are considered hereditary (stature, body proportions, and cranial and facial indices). We can see from this how opinions differ with The problem to be resolved should be to deter- regard to the problem Newman presents. mine preciselyquantitatively and qualitatively 4. Farther on Newman writes (p. 323): "Ofthe influence that heredity and environment the remaining standard dimensions, only [italics (in their broadest sense) have exercisedon the mine} head form and relative head height show present somaticvariabilityofthe American distributional patterns not readily interpreted asaborigines. adaptive ones. Indeed, the earlier and marginal However, inhis Summary (pp. 324-325) distribution of long heads and the apparently Newman generalizes about the applicability of late arrival of low heads seems best explained by Bergmann's and Allen's rules for the formation migrations of people differing in these regards. of the Amerindian "races" without regardto But since the diagnostic criteria of most racial all the reservations and exceptions made in the classifications of New World aborigines are prin- course of his article and cited here. And in a cipally the body size and proportion traits shownlater work (49) he reaffirms his opinion: "All here to be adaptive, itis most curious that if these data should make it clear why anthropolo- explanations of these classifications are attempted gists have paid serious attention to the applica- at all, they are in terms of a separate migration tion of the ecological rules toman. If anything, from Asia to account for each race." The "head these rules seem to be more closely operative in form and relative head height" are hereditaryman than in other species of homeotherms" characteristics because head breadth (dominant) (p. 104). and head length (recessive) enter into their This is important, since other anthropologists determination. The same is true with respect to not only accept such a generalization but tend face height (dominant) and face breadth (re-to amplify it; Stewart (55), for example, says: cessive), broad nose (dominant) andnarrow "Thus Marshall Newnan has demonstrated for nose (recessive)characteristics to which New-the hemisphere at large that many elements of man also refers in the course of his argument. the Indian phenotype are primarily adaptivere- On the other hand, the most discussed racial sponses to environment and are distributed in classification of the American aborigines (14,accordance with Bergmann's and Allen'seco- 47)that of Imbelloni, whouses as his basislogical rules" (p. 262). And,as expected, he theclassifications of Biasutti,Eickstedt, and goes on to state: Schwidetzky takes into account cranial, facial, When the first Asiatics crossed Bering Strait into Amer- and nasal indices, all of them hereditary char- ica they enterid a huge cul-de-sac offering every variety acteristics,in additiontostature. Therefore, of environment and no forerunners to mix with. Are- Newman's statement does not exactlyreflect constructionof what happenedthereafter takesinto the real facts. account that the resulting population at the time of 5. Finally, Newman states discovery constituted a major isolate that was homoge- (pp. 323-324): neous, both phenotypkally and genotypically.(p. 269; "From the foregoing, it seems clear that body italics mine)

26 I wish, however, topresent arguments that Other authors confirma divergence of opin- establish categorically thetrue scope of this ions about the applicability and importance of "geographical and climatic determinism." I have Bergmann's and Allen's rulesto the biological already said above that accordingto Rensch thevariability of the American Indian. Roberts(53), percentage of exceptions among birds and mam- in an important contribution to the subject,says: mals to Bergmann's and Allen's rules isvery high. The weight/temperature relationships here demon- strated suggest that Bergmann's ruleisapplicableto In an interesting paper on thesame problem, man. Clearer definition of "body size" is, however, neces- C. G. Wilber(61)says that "On the basis of sary. Defined by reference to stature, although from the our present knowledge the rules of Bergmann series here considered Bergmann's rule mightseem to be applicable, this suggestion is refuted by and Allen appear to be of historicalor descrip- more extensive material [italics mine]. Defined by weight, it isnot only tive interest only and certainlyare not valid applicable but needs restatement to incorporate, with generalizations for animals in the cold" (p. 332). the postulated variation in size among subspecies, similar His Summary is as follows: variation within the subspecies. (p. 551) This brief and rapid survey does not postulate that Ashley Montagu (44,45)holds that such climate is without effect on man. At another time the zoological rules "are tosome extent also appli- ecological effect of this variable of man will be discussed. cable to man," but adds: This presentation attempted to show ina sketchy fashion the following: The application ri Bergmann's and Allen's rules toman 1) The rules of Bergmann and Allen find littlesup- have been seriously and cogently questioned (Wilber), port as causal agents in modern studies of temperature on the ground that the ;nadequate data has been im- regulation in homeotherms. properly interpreted, and that in any eventman has 2) The various formal examples often cited in favor never responded to his environment in a passive manner, of these ecological generalizations do notsupport the but has always done everything within hispower to case of the climatic determinists. One is forced to con- control and shape the environment to his requirements. clude that the rules just do not apply causally to animals. But while this is undoubtedly true, it should beremem- 3) In man the ecological forces supposedto be acting bered that ecological rules are generalizationsto which are not doing so; Eskimos were not cold, the skinny exceptions can be found in every group, but that by and aboriginal Australians were. large they do apply to most populations ofa species. The studies of Newman and of Roberts 4) The rules of Bergmann and Allen haveno causal on New and Old World human populations lend role in the formation of racial diflerences inman. Such strong support to use of these rules on the part of same anthropologists is the view that ccolog :al rules apply toman as well as a source of misinformation and confusion [italics mine]. to other animals. 5) Some human groups havemet the demands of Weiner (58), for his part, states that Berg- severe climate by technological and behavioral adjust- mann's and Allen's rules are applicable to animal ments; the Eskimos are an example. Others have de- veloped specific heat-conserving functional changes with populations in general and continues: "That no gross morphological changes; the Australian aborig- human body-size and shape tend to follow these ines are an example. rules has been demonstrated in several studies. The mean body-weight of populations in hot Gam, on the other hand, concludesas follows regions is demonstrably lower than that in a detailed criticism refuting Wilber (22): tem- perate and cooler climates" (p. 455). But farther I know of no anthropologist so rash as to claim that on in the same work, on examining genetic and temperature and the radiant heat load art exclusive or nongenetic factors in climatic adjustments, he even major causes of the differences between geographical races, or that the past-century formulations of Bergmann points out: and Allen completely solve the problems ofrace forma- Twin studies indicate that variations in body-shape, size, tion in man. I know of none who adopt the Lamarckian fat deposition, growth pattern, skeletal and physiological approach that Dr. Wilber so gleefully demolishes. But, maturation are all determined by genetic constitution to when Wilber asserts that the "rules of Bergmann and a larger extent than by purely environmental factors. Allen have no causal role in the formation of racialCertain of the population differences rest undoubtedly differences in man," I doubt very much that he intended on distinctive genotypes or multifactorial recombinations, such a sweeping and untestable counter-generalization. e.g. nose-shape, or the ratio of limb length to trunk

27 length since such characters remain unaffected on change to ecological and environmental conditions have of environment. (p. 460) greatly interested numerous biologists and physi- Baker (1), also referring to the racial differ- cal anthropologists. A human Adaptability Sec- ences in heat tolerance, says: "These results fur-tion is part of the International Biological Pro- ther suggested that the differences found weregram (1BP), and several international meetings not a function of transient environmental effects have been held (Burg Wartenstein, 1964; War- and may be mostly genetic in origin" (p. 303). saw, Kyoto, and New Delhi, 1965) to discuss Later(2) he arrivesatthe conclusion thatsuch subjects as "Human adaptability and its "However, it is not enough to find evidence of methodology," "Human adaptability to environ- climatic adaptation. There remains the muchmental conditions and physical stress," and a larger question of how climatic selection wouldproposed regional study of high-altitude adapta- operate on man's genetic structure to produce tion. A detailed presentation of the topics of the these adaptations" (p. 4). Human Adaptability Section appear in the Guide In general terms I believe that Dobzhansky of the Human Adaptability Proposals;a the re- (21) best synthesizes the question. sults of the meetings have been published (8, The environment thus instigates, foments, conditions and 41, 63). circumscribes evolutionary changes; but it does not decide In November 1967, 60 scientists from 12 na- exactly which changes, if any, will occur. (p. 408) tions participated in the Conference on Man at The rules of geographic variation used to be a happy High Altitudes, sponsored jointly by the U.S. hunting ground for partisans of Lamarckism and selec- National Committee for the IBP and the World tionism, abounding in data intTrpretable astheir pre- and Pan American Health Organizations. They dilections decreed. Nowadays these disputes may, I hope, agreed that research on high-altitude peoples be bypassed. The rules attest in any case that the environ- mentisimportantas an instigatorofevolutionary could also be applied profitably to populations changes. At the same time, it must be emphasized that living at sea level and to their medical problems, what has been observed arcrules indeed, not laws. and recommended that coordinated studies be (p. 412) made on "problems of growth, ageing, nutrition, Exceptions to the rules do occur, as Rensch, who has fertility, natural selection and epidemiology." contributed more than anyone else to their study, has As for investigations completed or in progress duly stressed. And while these exceptions do not exactly prove the rules. they arc in some ways as valuable as in America on human adaptability to different therules themselves. The lessonto be derived from conditions of heat, humidity, altitude, and lati- them is that, although the environment may guide the tude, a number of monographs have appeared evolution of living things, it does not prescribe just what especially on peoples of the arctic regions (Eski- change must occur. (p. 413) mos) and of high altitudes (Quechuas and Waddington (57) offers an explanation of how Aymara* Continuing the research program so this genetic-environment interaction is effected. successfully initiated by Carlos Monge, Hurtado, and other investigators at the Institute of Andean We have, infact, found evidence for the existence of a"feedback" between the conditions of the environ- Biology in Peru in 1928, Paul T. Baker and ment and the phenotypic effects of gene mutations. The collaborator!: are responsible for the most recent "feedback" circuit is the simple one as follows: (1) en- studies on acclimation and adaptability to dif- producedevelopmentalmodifica- vironmentalstresses ferent ecological environments. tions; (2) the same stresses produce a natural selective pressure which tendstoaccumulate genotypes which Despite these advances, the problem of human respond to the stresses with co-ordinated adaptive modi- variability and adaptation to different climatic fications from the unstressedcourseof development; conditionsthat is, an evaluation of the interde- (3) genes newly arising by mutation will operate in an epigenetic system in which the production of such co- pendence of nature and nurtureis still unre- ordinated adaptive modifications has been made easy. solved. Besides Dobzhansky, whose opinionis (p. 399) Published by the Central Office of the IBP (7, Mary- Recently the problems of human adaptability lebone Road, London, N.W. 1).

28 34 quoted above, Neel and Schull (46) had already races or breeding isolates of many authors. . Conse- quently, though the actual number of races in the world stated: exists apart from observers, the number recognized de- It is therefore practically impossible when one is dealing pends..upon the aim of the observers. (p. 90) with human populations to crcatc situations which throw a sharply critical light on thc relativeimportance of The concepts of geographical, local, and micro- heredity and environment. races, expounded by Garn first in 1961 and more recently in 1965, are thus comprehensible. How- Efforts to establish principal biological sub- ever, before examining these possiblebiological divisions of Indians subdivisions or races of the American Indian, I Once the variability of the living aboriginal think it advisable to repeat what I said at the populationsthe presence of different subspe-beginningthat certain contemporary anthro- cific polytypical forms, regardles-s; of origin and pologists, especially Montagu and Coon, take causesis recognized and accepted, allefforts the view that the existing variability and bio- at a systematization orclassification require a logical differences among the diverse groups of prior definition of the concept of race. aborigines are not sufficient to justify a racial This is not the place or time to analyze such subdivision. disputable and controversial themes as the non- Coon (18), however, in referring to a work existence of human races according to Living-of Osman Hill on "The Soft Anatomy of a stone (40) and Brace (17), orthe "ethnicNorth American Indian," says:'`...they are groups" of Ashley Montagu (44, 45) as substi-Mongoloids of a particular kind, just as they tutes for racial groups, Of the skepticism ofBarni- would be Caucasoids of a particular kind had cot (10) about the possibility ofdefining the the New World been peopled by a small band human "races" with the necessary precision. Theof Upper Paleolithic Europeans." Here I wish classic description of the "human race" based on to point out that the possible presenceof Cauca- the typologist approach is of historical interestsoid elements of European origin had been spe- only. Let us stick to the modern populationist cifically noted in 1928 by Cottevieille-Giraudet.4 view and accept, for our purposes, any of theApparently certain culturalfeatures citedby fundamentally similar definitions of Dobzhansky Greenman (24) as persisting on the eastern (20), Laughlin (39), Garn (23), Mayr (43),coast of North America tend toconfirm this Bielicki (13), and others. I should like, how- supposition. ever,to quote here some considerations pre- The classicai subdivisions of mankind from sented by Laughlin (39), because of their clarity the serological point of view proposed by Otten- and conciseness. berg, Snyder, Wiener, and principally Boyd (16) Race does not refer to an arbitrarily selected series of mention solely an Amerindian group, to which individuals, even though they may be similar in ap- they accord a certain homogeneitythis despite pearance, i.e., of the same typc. Races may beconti- the fact that Mourant had noted evident sero- nental or local, homogeneous or heterogeneous, large or small, ancient or recent,distinctive in appearance or logical differences between different populations nondistinctive, sharply bounded or imperceptibly bounded of American aborigines. and possess a high or low degree of genetic relationship During the last few years, the multiple investi- between members. The term race can bc used atdif- gations of Henckel, Layrisse,Lisker,Loria, ferent levels or abstraction from a continental race down Salzano, to a local race or even their tribalsubdivisions. Local Matson, Neel, Robinson, Reynafarje, races are composed of collections offamily lines which Sandoval, Sutton, Swanson, Zepeda, and others constitute breeding isolates within thc larger population. have made available greatly increased data on No arbitrary standard of magnitude exists for the size or number of differences which mustexist for a group 4 See my own Manualof Physical Anthropology to be termed a race....For summary purposes and (Springfield,Illinois,1960,pp.633-635) and "The major contrast such high levels of abstraction are suit- Upper Palaeolithic and theNov World. Comments" able. However, for research purposes itis necessary to (Current Anthropology, Vol.5, No. 4, 1964, pp. 321- compare the smaller, constituent subdivisions,the local 322).

29 35 diverse antigens, hemoglobins,transferrins, and logically and serologically typical traits. Earlier, haptoglobins that demonstrate an obviouspheno- however (p. 120), he alludes to the factthat typic and genotypic variability in theseaborig- some anthropologistsfind the morphological, ines. This confirms the present conception re- serological, and biochemical differences between garding the evolutionary process of thehuman Mongoloids and American Indians insufficient species and the formation of races according tofor separating them into two geographical races populationist and dynamic criteria. and join them into a single, polytypic geo- In this respect, I cite the latestconclusions graphical race. I agree with him that there are of Matson and collaborators (42) onSouth slight grounds for this racial hypothesis. America: 2. As for local races, Gan writes: "In con- which are geographi- Yet as pertaining to the present study, it seems that a trast to geographical races sensible position of equanimity, based on the available cally delimited population collections, local races blood group data, would permit of an hypothesis that correspond more nearly to the breeding popula- the American Indians arc not completely Mongoloid, tions themselves. Whether isolated by distante, that thepresent Polynesian populations area racial by geographical barriers or by social prohibitions, mosaic and that migrants from both west and easthave local races are totally or largely endogamous, and contributed genes to present panmictic potpourri which is Polynesia. (p. 188) the very small amount of gene-flowordinarily and related local races" Since this variability of blood groups observed comes from contiguous among Indian populationsdoes not coincide(p. 16). with other morphological variations, one is led 3. What Dobzhansky defined as microgeo- graphical races and what Lasker prefers to call to think that Garn's conclusion(23) is correct: breeding populations are what Garn calls micro- As withclassificationsbased on morphologicaltraits races (p. 18); inthese he makes manifest cer- rather than on the populations themselves,artificial local "serological races" add nothing to human taxonomy. tain differences in the composition of a The major use of blood groups inclassificationis in race. He adds: "Micro-races,though not isolated the comparison and analysis of natural populations, and geographically or by extensive cultural prohibi- in the study of natural selection in contemporary races. tions, still differ from each other in numerous (P. 51) ways." I shall now review in some detail theracial 4. With regard to the value and usefulnessof taxonomy proposed by Garn (23)with reference these concepts, Gam explains(p. 22): "Geo- to geographical, local, andmicro-races, begin- graphical races, local races and micro-races offer ning with his definitions and continuing withopportunities for very different investigations his subdivision of the aboriginal populations ofin relation to race. One is not more real or more America. fundamental than the other, but each provides 1. The concept of geographical race, first used tne answer todifferent questions and the solu- by Rensch, is defined by Garn as "a geograph- tion to different problems of ongoingevolution ically-delimited collection of similar races," and in man." he adds that "the existence of geographical races 5. In referring specifically to the populations is due, of course, to the great geographical bar- the Amerindian riers, chief among them oceans, that formerlyof the New Worldthat is, geographical race---Garn mentions a series of limited the expansion and migration of local local races that "in strict contrast with geo- races and protectedthem from introduction of different genes" (p. 14). Garn's geographicalgraphical races are true evolutionary units. As have race is equivalent to the conceptof continental populations such localracesevolveor races. On this basis hedivides mankind intoevolved separately." He recognizes that "with nine geographical races, only one of whichthesuch a diversity of local races, it is clearly impos- Amerindian--he attributes to the New Worldsible to make a listing of all of them" but says (p. 128). This he describes with some morpho- that "it is possible to call attention to some local

:30 36 races that exemplify particular taxonomic, de-include in a single local race all the aboriginal scriptive, or evolutionary problems" (p. 140). populations of South America, from the Goajiro And before proposing a subdivision of the living in the north to the Araucanians in the south aboriginal populations, he contends: "In pre-(latitude 10° N to 40° S), living at altitudes Columbian America, there were hundreds ofranging from sea level to 4,000 meters. And this such local races, each with its own language. applies as well to the North American local race, We still recognize the Penobscot, the Pima, theranging from the Athabaskan and Algonquian Papago and so on. Other local races, in the in the north to the Mayas to the south (latitude Americas, as in Europe and Asia, constitute a60° N to 18' N). number of isolated or semi-isolated populations as is true for the several Apache and NavahoDiscussion groups now." An appraisal of new techniques for interpret- Only after these admonitions does Gam listing the physical anthropology of the American five Amerindian groups of local races: North Indian was on the agenda of the Fourth Summer American, Central, Circum-Caribbean, South Seminar in Physical Anthropology, held in New American, and Fuegian. His descriptionof York in September 1949. I should like to start biological characteristics (pp. 144-146) is quiteby quoting from the report of that discussion poor, and in the case of two of these local races (62). he refers exclusively to cultural traits. This in spite of the fact that he qualifies them as "true Washburn suggested that much of the confusion preva- evolutionary units." lent in the American Indian field today is a result of too many' varying techniques being used on the same Incidentally, with reference to the origin of materials, each yielding different results and hence some- these local races Gam takes a firm position what different interpretations. He indicated that some (pp. 128-129): "Once, local differentiation in re-evaluation of the severalmorphologic, metric and the Americas was attributed to successive waves genetic techniques was in order if the best results were of migrations. Today such diversity is generally toll? obtained from the available data. (p. 33) accepted as the result of natural selection acting This carefully weighed opinion expressed 19 on generally small population isolates, some of years ago still constitutes, to my way of thinking, whom may have a respectable antiquity of as one of the principal reasons, though not the only much as 20,000 years, as shown by radio-carbon one, why even now attempts at a racial taxonomy dating." 5 of the American aborigines yield such vague, Ihave emphasized the concepts on which imprecise, and even contradictory results. Garn bases his racial taxonomy, particularly with I agree with Baker (4) that the concept of reference to the region under discussion, because race has two usesas a pedagogic device for I consider his concepts proper and his principles teaching human variation and as a research tool valid. However, I disagree on the subject of the for investigating biological variation, and that, five local races he proposes, since mere observa- as he explicitly recognizes: tion of some of the aboriginal populations in- habiting the areas in which he locates them Indeed, racial classification systems arc, at best, interim structures for dealing with genetic and phenotypic dis- (especially demonstrates their great biological tances, and should be replaced by e.uantitative systems. somatic and serologic) heterogeneity. It may be hoped that the comparrtive method will be According to Garn's own definition,Ifind replaced by the more accurate m:thod of mechanism it difficult to understand how it is possible to analysis. However, neither of thesc hopes are . to materialize in the near future and for decades race is 5 Current data indicate that man's antiquity in America likely to remain a useful scientific concept. As such, it dates back 35,000 to 40,000years. See Krieger, in appears that the race concept will remain in human prehistoric Man in the New World (J. D. Jennings and biology for many decades even thoughitwill, un- E. Norbeck. eds..University of Chicago Press,1964, doubtedly, be a constantly changing informational con- p. struct. (p. 25) 317 I have expressed a similar point of view on biological heterogeneity as a consequence of the various occasions, in discussing the usefulnessdiverse origin of the immigrating contingents and importance of anthropometry and ostcome- who peopled the New World some 40,000 years try in any attempt to determine the variabilityago and also as the result of a process of adapta- between different population groups. I reiteratetion to different environmental and ecological this belief and in its support cite the well-docu-conditions. At the present time, this is a point mented opinion of Hunt (34). of controversy among investigators. In all likeli- As remote racialhistory has ceased to be the chiefhood the biological differentiation of the Amer- excuse forfield anthropometry, microevolutionespe- indian is due to the joint action of the two. cially the adaptive value of racial features in different However, more data are necessary before it can environmentshas become the core of recent studies. be determined which is the more important. Schemes of measurement are being revised in terms of 4. The most widespread and best-known sub- the 'hereditabilityof somatic dimensions and factors of physical growth. Work on demography, new techniques divisions of the American Indian in local races of mapping, models of gene-flow, and racial physiology we owe to Garn. lf, as he says, these "are true is proceeding rapidly. The unraveling of microevolution evolutionary units. As populations, such local is a sufficient challenge to maintain the vitality of anthro- races evolve or have evolved separately," then pometry for a long time to come. In particular, it offers many opportunities for the collaboration of physical and it is necessary to assemble a series of biological cultural anthropologists. (p. 82) characteristics that will make it possible to dis- tinguish the local races proposed for America. I have deliberately included inthis paper many quotationstoo many, the reader may 5. The Amerindian groups of local races pro- thinkfrom different workers, so as to presentposed by Garn are not precisely defined, and he the most recent and contradictory opinions on appears to recognize this himself when he writes the topic and thus document as objectively as (p. 121): "Such differences in taxonomic opinion possible my own view, contained in the follow-are both legitimate and salutary. They point out ing provisional conclusions: problem areas that need resolution. Areas of agreement on the other hand, may reflect prob- 1.I consider acceptable the proposal of various anthropologists for uniting all the aborigineslems long settled, or they may reflect a virtual of the New World under the denomination lack of information." Amerindian Geographical Race. What is absolutely indispensable in the imme- 2. That biological variations create perceptible diate future is the organization on a hemisphere- differences between American Indians in diverse wide basis of intensive, methodical, uniform regions of the continent and thus necessitate a biologicalinvestigations(somatic,serologic, taxonomy of local races, in accord with popula-physiological, psychosomatic, and so on) of the tionist and dynamic criteria,has been fully various aboriginal populations that permit subse- proved and substantiated by many investigations quent comparative studies and finally the estab- in various fields of human biology. lishing of local races with a clear and indisput- 3. Attempts have been made to explain thisable differential biological base.

REFERENCES I. BAKER, P. T. Racial differences in heat toler- 4. BAKER, P. T. The biological race concept as a ances. Amer. I.Phys. Anthropol. n.s. 16:287-305, research tool. Amer. f. Phys. Anthropol. n.s. 27:21 1958. 26, 1967. 2. BAKER, P. T. Climate, culture and evolution. 5. BAKER,P.T.,E.R.BUSKIRK, E.PICON- Human Biol, 32:3-16, 1960. REATEGUI, J. KOLLIAS, and R. B. MAZESS. Regulaci6n 3. BAKER,P.T. EcoLgical andphysiological de la temperatura corporalen indios quechuas adaptation in indigenous South Americans.In P. T. nativos de la altura. Anales del Instituto de Biologia Baker and J.S. Weiner (eds.),The Biology of Andina 5:286-298, 1966. Human Adaptability. Oxford,Oxford University 6. BAKER, P. T., E. R. Busxunt, J. KOLLIAS, and Press,1966.541 pp. R. B. MAZESS. Temperature regulation at high aid-

32 38 tude: Quechua Indians and U.S. whites during total among Indian groups of the Southwest. Amer. body cold exposure. Human Biol. 39(2) :155-169, Phys. Anthropol., n.s. 20:499-508, 1962. 1967. 27. HARRISON, G. A. Environmental interaction. 7. BAKER, P. T., and M. A. LITTLE. Bone density In 0..A. Harrison, J. S. Weiner, J. M. Tanner, and with age, altitude, sex and race factors in Peruvians. N. A. Barnicot, Human Biology. Oxford, 1964, pp. Human Biol. 37(2) :122-136, 1965. 144-149. 8. BAKER, P. T., and J.S. WEINER (eds.). The 28, HIERNAUX, J. Le concept de race en anthro- Biology of Human Adaptability. Oxford, Oxford pologic physique.Fl Congres International des University Press, 1966. 541 pp. Sciences Anthropologiques et Ethnologiques 1:471- 9. BAmicor, N. A. The size and shape of the 477. Paris, 1962. body. In G. A. Harrison, J. S. Weiner, J. M. Tanner 29. HIERNAUX,J.Heredity:;-.ndenvironment: and N. A. Barnicot. Human Biology. Oxford, 1964, Their influence on human morphology. A compari- pp. 197-216. son of two independent lines of study. Amer. 1. 10. BARNICOT, N. A. Le probleme de la race dans Phys. Anthropol. n.s. 21:575-589, 1963. l'etatactuel des connaissances scientifiques. Rev. 30. HIERNAUX, J. Applicabilite du concept de race Int. Sciences Sociales,17 (1) :90-92, 1965. l'espece humaine. Rev. Int. Sciences Sociales 17(1): 11. BENOIST,1.Microraces et isolats. Rev. Mt. 117-119, 1965. Sciences Sociales 17 (1):93-95, 1965. 31. HIERNAUX, J. Problemes de definitions sur la 12. BIELICKI, T. La genetique des populations et race. Rev. Int. Sciences Sociales 17(1):120-i 22, 1965. la formation des races. Rev. Mt. Sciences Sociales 32. HRDLICKA, A. The problems of the unity or 17 (1) :96-98, 1965. pluta:ity and the probable place of origin of the 13. BIELtext, T. Typologistes contre population- American aborigines. Amer. Anthropol. 14:11, 1912. nistes et theorie genetique. Rev. hit. Sciences Sociales 33. HRDLICKA, A. The origin and antiquity of the 17(1):99-101, 1965. American Indian. Rev. ed. Smithsonian Report for 14. BIRDSELL,J.B. The problem of the early 1923. Washington, 1928, p. 481. (Publication 2778) peopling of the Americas as viewed from Asia. In 34. HUNT, E. E. Anthropometry, genetics and W. S.Laughlin(ed.), Papers on the Physical racial history. Amer. Anthropol. 61:64-87, 1959. Anthropology of the American Indian. New York, 35. IMBELLONI, J. Nouveaux apportsla classifica- Viking Fund, 1951, pp. 1-68a. tion de Thomme am&icain. Miscelanea Paul Rivet 15. BOYD, W. C. Blood groups of American Octogenario Dicata 1:107-136, 1958. Indians. American 1. Phys. Anthropol. 25:215-235, 36. KEITH, A. A New Theory of Human Evolu- 1939. tion. London, Watts, 1948, p. 218. 16. Bovo, W. C. Genetics and the human race. 37. LASKER, G. W. The Evolution of Man. New Science 140(3571):1057-1064, 1963. York, Holt, Rinehart and Winston, 1961. 240 pp. 17. BRACE, C. L. On the race concept, and Com- 38. LAUGHLIN, W. S. Introduction.In W.S. ments. Current Anthropol. 5(4) :313-320, 1964. Laughlin(ed.), Papers on the Physical Anthro- 18. COON, C. S. The Living Races of Man. New pology of the American Indian. New York, Viking York, Knopf, 1965. 344 pp. Fund, 1951, pp. v-vii. 19. DOBZHANSKY, T. Human diversity and adap- 39. LAUGHLIN, W. S. Races of mankind: Con- tation. Cold Spring Harbor Symp. Quant. Biol. XV: tinental and local. Anthropological Papers of the 385-400, 1950. University of Alaska 8(2) :89-99, 1960. 20. DOEZHANSKY, T.Evolution,Genetics, and 40. LIVINGSTONE, F. B. On the non-existence of Man. New York, Wiley, 1957. 398 pp. humanraces.Current Anthropol. 3(3):279-281, 21. DOSZHANSKY, T. Evolution and environment. 1962. In The Evolution of Life. Chicago, University of 41. MALHOTRA, M. S. (ed.) Human Adaptability Chicago Press, 1960, pp. 403-428. to Environments and Physical Fitness (Delhi Sym- 22. GARN, S. M. A comment on Wilber's "Origin posium 1965). Madras, India, Defence Institute of of Human Types." Human Biol. 30:337-39, 1958. Physiology and Allied Sciences, 1966. 23. GARN, S. M. Human Races. 2nd ed. Spring- 42. MATSON, G. A., H. E. SUTTON, R. ETCHEVERRY, field, Illinois, Charles C Thomas, 1965. 155 pp. B. J. SWANSON, and A. ROBINSON. Distribution of 24. GREENMAN, E. F. The Upper Palaeolithic and hereditary blood groups among Indians in South the New World. Current Anthropol. 4(1) :41-91, America, with inferences concerning genetic con- 1963. nections between Polynesia and America. Amer. 1. 25. HAMMEL, H. T. Effect of race on response to Phys. Anthropol. n.s. 27:157-194, 1967. cold. Fed. Proc. 22(3), 1963. InYearbook of Physi- 43. MAYR, E. Les races dans l'evolution animale. cal Anthropology 11:208-213, 1965. Rev. Mt. Sciences Sociales 17 (1) :126-127, 19,"*. 26. HANNA, B. L. The biologicalrelationships 44. MoNTAou, M.F.A. An Introductionto

33 39 Physical Anthropology. 3d ed. Springfield, Illinois, 54. SALZANO, F. M. The blood groups of South Charles C Thomas, 1960. 771 pp. American Indians. Amer. 1. Phys. Anthropol. n.s. 45. MONTAGU, M. F. A. The Science of Man. 15 :555-579, 1957. New York, Odyssey, 1964. 158 pp. 55. STEWART, T. D. A physical anthropologises 46. NEEL, J.V. and W. J.&Hun,. Human view of the peopling of the New World. South- Heredity. Chicago, University of Chicago Press, western J. Anthropol. 16(3) :259-273, 1960. 1958. 361 pp. 56. STEWART, T. D., and M. T. NEWMAN. An 47. NEWMAN, M. T. The sequences of Indian historical resume of the concept of differences in physical types in South America. In W. S. Laughlin Indian types. Amer. Anthropol. 53:19-36, 1951. (ed.), Papers on the Physical Anthropology of the 57. WADDINGTON, C. H. Evolutionary adaptation. American Indian. New York, The Viking Fund, In The Evolution of Life. Chicago, University of 1951, pp. 69-97. Chicago Press, 1960, pp. 381-402. 48. NEWMAN, M. T. The application of ecological 58. WEINER, J. S. Climatic adaptation. In G. A. rules to the racial anthropology of the aboriginal Harrison, J. S. Weiner, J. M. Tanner, and N. A. 144- 0'0) New World. Amer. Anthropol. 55:311-327, 1953. Barnicot, Human Biology. Oxford, 1964, 59. WEINER, J. S. Acclimatation et differences de 49. NEWMAN, M. T. Adaptation of man to cold climat: !curs effets sur les differences raciales. Rev. climates. Evolution 10:101-105, 1956. Int. Sciences Sociales 17 (1): 159-162, 1965. 50. NEWMAN, M. T. Geographic and microgeo- 60. WIERCINSKI, A. The racial analysis of human graphic races.Current Anthropol. 4(2) :189-192, populations in relation to their ethnogenesis. Cur- 1963. Comments 4(2) :192-207, 1963. rent Anthropol. 3( 1) :2-46, 1962. 51. RENSCH, B. Some problems of geographical 61. WILBER, C. G. Physiological regulations and variationandspeciesformation. Proc. Linnean the origins of human types. Human Biol. 29:329- Society of London, 150th Session, 4:275-285, 1938. 336, 1957. 52. RENSCH, B. The laws of evolution. In The 62. Yearbook of Physical Anthropolzgy, 4. New Evolution of Life. Chicago, University of Chicago York, Viking Fund, 1949, p. 33. Press, 1960, pp. 95-116. 63. YOSHIMURA, H., andJ.S. WEINER(eds.) 53. ROBERTS, D. F. Body weight, race and climate. Human Adaptability and its Methodology. Tokyo, Amer. I. Phys. Anthropol. n.s. 11:533-558, 1953. Japan Society for the Promotion of Sciences, 1966.

34 40 -1r

BIOLOGICAL SUBDIVISIONS OF THE INDIAN ON THE BASIS OF GENETIC TRAITS

Miguel Layrisse

Improved techniques for the identification ofdifferentiate the American Indians so far studied new traits in the erythrocytes and in chemical from other subdivisions of mankind. However, components of the plasma have brought about aintertribal differences suggest that Indian popu- new era in the field of population genetics.Sev- lations do not constitute a homogeneous gene eral of these traits have shown a distribution re- pool as is sometimes suggested in the anthropo- strictedto a single subdivision of mankind, logical literature. attaining thereby the rank of gene markers; Characteristically, Indian populations through- others are common to virtually all populations, out the Americas have been organized in terri- although with variable frequencies; still others torial and tribal groups displaying a certain are fixed and/or of very lowfrequency. It is now esprit de corps based on a distinctive cultural possible to set up a more comprehensive "mo-and linguistic homogeneity. The tribes live or saic" of gene frequenciesin populations, to have lived in a state of semi-isolation, separated complement the traditional division of mankind from their neighbors by natural and cultural based predominantly on somatological charac-barriers that have i'..nctioned as more or less teristics. effective impediments to gene flow. Larger tribes Several attempts have been made at classifying may be divided into smallersubtribes made up mankind on the basis of genetic traits (2, 3, 30). of several villages. These villages are frequently Originally classifications were made by means of quite independent of each other, with a strong blood groups; later, other blood traits were alsotendency to local endogamy. For instance, in taken into consideration. In a recent classification three recently studied villages of the Warao Boyd ( 3) distinguishes 13 different human races Indians of the Orinoco Delta, more than 90 per comprising 7 major groups, one of which is the cent of the marriages recorded werecontracted "American group" of a single racethe "Ameri-between individuals of the same local group. can Indian." Boyd sJggests that "it may eventu-This high incidence of local endogamy occurred ally be possible to distinguish between North in an almost complete absence of geographical American and South American Indians serologi- barriers to communication. Similarly high de-

cally...." grees of village autonomy and localendogamy Considered as one population, American In- were observed, although underdifferent cultural ians in general are characterized by a high fre- and environmental conditions, by Salzano, Neel, quency of blood groups 0, M, R2, Fyn, Dia, andand Maybury-Lewis (28) among the Xavante ABH secretor, and by a low frequency or ab-and by Salzano (26) among the Caingang. sence of A2, B, Ro, r, Lua, K, Lea, and abnormal Analysis of the peculiar socio-political charac- hemoglobins (3, 21, 22). These characteristicsteristics of the Amerindian is needed so that we

35 41 may learn how these subdivisions affect the fre-Maya groups living in Central America; Sal- quency of genetic traits. In this connection atten- zano (27) in Brazilian tribes, Matson et al. (19) tion must be drawn to (a) the intertribal andin Peruvian tribes; and Matson et al. (20) in (b) the intratribal distribution of genetic traits. Chilean tribes. COrdova, Lisker, and Loria(7) did not find such divergence in Mexican Indian Intertribal distribution of blood traits tribes, but their study was conducted in a large population. One of the most striking examples of inter- tribal divergence in frequencies of blood traits Intratribal distribution of genetic traits is provided by two tribes living in the Amazon Territory of Venezuela: the Makiritare and the The study of 10 villages of the Yanomama Yanomama. The Yanomama occupy an exten-tribe, carried out in 1966, provided excellent sive area in southern Venezuela and northern information on how gene frequencies may vary Brazil. The Makiritare inhabit adjacent terri- between tribal villages in the same area separated tories, mainly to the north-northeast. There are by relatively short distances (2). Five hundred considerable differences in culture between the and sixty-eight individuals, 72 per cent of the two tribes. The Yanomama are a Pa leo-Indian village populations, were examined for7blood- society of bunters and gatherers with a rudi-group systems, 7 serum protein traits, and7 mentary agriculture(2, 5, 15, 14, 32).The Maki- erythrocyte enzyme groups. The variation in ritare have what basically can be identified as a gene frequencies was greater than in any other neo-Indian culture based on slash-and-burn agri- tribe examined so far under similar circum- culture(14).Although there is evidence ofstances. Genes with low frequenciesMS, NS, interbreeding between the Makiritare and the112, Lpa, and Agawere found to have been Yanomama, the actual gene flow between themlost in several villages, while genes with high can be considered extremely limited. The com- frequenciesPGM and Sewere fixed in some parisons of blood-group frequencies between thevillages. Genes with intermediate frequencies, Makiritare and two Yanomama subtribes (Sa- such as R1, Ms, Ns, Fya, and Jka, showed a very nema and Waica) showed significant differences wide range of frequencies. in practically all the blood-group systems. S, of The Warao villages of the Orinoco Delta the MNS system, is high in the Makiritare (63provide another illustration of gene frequency per cent) but only 19 per cent in the Waica and variation within the same tribe. Three villages 9 per cent in the Sanema. RI, of the Rh system, were studied: Jobure, Sacupana, and Winikina, is about twice as high in both Yanomama sub-with populations of 190, 160, and 300 respec- tribes as in the Makiritare. The Yanomama sub- tively(31).Three previous studies of these tribes showed the lowest frequency of 112 ever tribes had shown a very low M (less than 15 found in Indians-6 per cent in the Waica and per cent) with a relatively high S (40 per cent), 3 per cent in the Sanema. The frequency of Fya traveling mostly with N, which is uncommon is 83 per cent in the Makiritare and 58 per cent in Indians; very high Ri; and negative or very in the Waika and Sanema. Dia is 16 per cent low frequency of D?. The village of Sacupana in the Makiritare, 0 per cent in the Waica, and was the eccentric tribe; its frequencies of the 3 per cent in the Sanema. Since the Sanema MNSs and Diego systems were completely dif- live in close geographical proximity to the north- ferent from those found in the other two villages ern Makiritare, the Diego frequency can pos- and from those previously reported. The fre- sibly be accounted for by intertribal admixture. quency of gene M was 90 per cent and of gene S This is by no means an isolated occurrence. only 13 per cent, and this gene was mostly For example, differencesthough less marked traveling with gene M. It was also found that in blood groups have been reported by Matson these tribes had 3 per cent of D?. The genealogy and Swanson(17, 18)between Maya and non- of the Sacupana village gave a clue to the ex-

36 42 planation of this gene distribution. Twenty-one group as representative of the population at out of the 48 living members representing the large would very likely produce a distorted pic- second generation come from a polygynous ture of the total gene pool, family in which the father happens to be MSs Blood tests in Indians are more meaningful if and his three sororal wives Ms. All the Diego- accompanied by other anthropologica1 and medi- positive cases found among the Warao of thecal studies. Demographyincluding a history of Sacupana villages came from one family, which the tribe and formation of the villages, and a we were able to trace through three generations. careful genealogyprovides valuable informa- It is interesting to observe that the members of tion that could serve for at least a partial under- this family also carried the transferrin Dem)standing of the variation of the gene pool be- which is absent in all Warao individuals ex- tween villages. The gene pool of a village among amined but present at various frequencies inthe Warao, for instance, isthe result of the Cariban tribes. Since the western and south- fusion of two, three, or four extended families eastern Warao territory is surrounded by Cari- that have split off from neighboring villages. ban tribes, itis likely that both Dia and Debi This pattern is very similar in many respects to genes were introduced into the Warao popula- the "fission-fusion" model observed by Neel tion by Carib in early times. and Salzano (23) among the Xavantes and by The three Xavante villages studied by Ger-Chagnon (5) among the Yanomama. It is not showitz el al. (9) showed a variation of 1to unlikely, therefore, that variants of these pat- 15 per cent for the gene SN and of 24 to 39 per terns operate in many tribes. cent for the gene 112; the other gene frequencies It is evident that in the formation of the new had comparable results. village, as previously described, there are circum- stances that may induce variation of the gene Comments frequencyinthe following generationsthe number of individuals from each family who Analysis of the frequency of blood groups and contribute to the breeding population, sororal other blood traits has shown that the socio-marriage, polygny, and others. Consequently, political unit used in cultural anthropology does random genetic drift would seem a likelier ex- not always correspond to the biological unit; it planation of the variation of gene frequencies in appears that the population of each village rather Indian populations than othergenetic forces than the whole tribe represents, in some cases, such as selection and gene flow. It is also prob- an independent gene pool. Therefore, itis not able that this dispersive process may operate surprising that independent studies carried outsimilarly in the formation of hybrid populations, in a tribe may show great divergence in gene explaining, in some cases, gene frequencies equal frequencies because they are performed in dif- to or even higher than those observed in the ferent villages. Such possible variations should original populations (14,16). be kept in mind when it is necessary to decide, Even though we have considered random ge- in trying to collect information from the litera- netic drift as the most important genetic process ture, whether the sampling isinadequate oroperating in small Indian populations, the In- really representative of the population under dians in their habitat must be exposed to selec- study. According to our experience in the study tive forces that maintain the polymorphism. So of two large tribes, the gene frequency obtained far, comprehensive stu dies on mor tali ty, fertility, by sampling the less closely related individuals and diseases in Indians are scarce; however, the from several villages of a tribe fits very well with new orientation in the study of Indians,which the frequency obtained by summing up the includes such information, will eventually make genes counted separately in villages in whichit possible to elucidate the role of selection in more than 70 per cent of the population wasthe variation of gene frequenciesinIndian examined. But taking the test results of one local populations. 37 43 firstsub- tomake a the bepossible Indians: might American Mayan 3. It Middle and like ofthe Totonacs Ishould division including ofChibchan variation, group group differ- this of asubdivision northern thesouthern theonly inmind might and frequen. Bearing thepossibility that families, However,isthe finally populations traits affiliations. these high discuss ofblood Hokan between tobe to Indian Dr.or exist general 10 per theAmerican thefrequency found to tendin of of characteristics.some ence which with amean somewhat here ofDia, low(from reconcile tocies 18 percent, and otheranthropologicaldescribed 4 to group, nt)in with previously Americanists (from of 3per c has classifi- thenorthernmean JuanComas byprominentofthese cent)in with a attempts none traits.Gan cent, ofthe However, blood 0 to10 per American Indians. of American group. South classify distribution South southern ofthe fitsthe all andFue.the ofmicrndifferen- cations included Thegeographyformation variations instance, It is 4. the ofthe vil- (8),for theCircum.Caribbeanpopulation. favors inspite and except popu- continent But subtribes to Indians homogeneous there are populations. in one area cultural, tiated between bepossible guian such alarge frequencies itmight thatin morphological, in gene tribe, thatcharacterize known marked a ofthesame genetic with toattempt lages peculiaritiescommon or lations differences.difficult of genetic with a tribes serologicalit isvery aborigines identify awhole ofother be and ofthe thefol- tribe as fromthat would Eventhough traits, the it fre- subdivisiongenetic formu- differentForexample, high basisof canbe heritage tribes. relatively first comprehensiveonthe of the considerations clusters investigate Indians, America available: be to theAndean accom- taxonomic can interesting among is not lowing theinformationtheEskimoNorth ofA, (24),which from that other quency othernon-Indian lated likely the by byNewman of basisof I. Itis very from noticed presence onthe geneticallyarecharacterizedofthe bythe and They panied connection, Indians distinguished andabsencetraits possible Chilean a/. Indians.geneB blood the the a American ofthe other genes; between byMatsonthat ofthe American traits, mentioned thepresencefrequency from genetic as thepossibilitythe Dil. In greatly Polynesians, and by gene differ arethe study; donot tribes in arecent distinguished they (20) andWarao, anthropological American ofheavy andby Indians. North because theYanornama Dia ortogether, Thevarious partly classified ofthe gene separately 2. analyze, been of tribes. to Theyhave absence beclassified, verydifficult thepresence could neighboring hesita- admixture. by however, traits, fromthe present the non-Indian characterized ofD?; that our on group tribes. asdifferent probable Indians in one lowfrequency in some highly American inthe Aand a observed show avery It is the overcome gene are subdividing willbe markers ofAlberta which tionin traits of gene microdifferentiationsBlackfeet of A,(6), genetic and ofNorth of theincrease inwhich TheBlood frequencythe rest basis with models (60 percent) from the pos- future be com- high group of near ofmathematicaltribecan this exploration and the use in a distinguishes An Yaqui,ofand found Indians. Navajo, thevariables American between thepresence(1,all sibleconnectionexhibiting interest puted. tribes beof great would LacandonBO-1 theYanomamaSci. transferrin amongNat.Acad. REFERENCES differentiation Proc. 26). genetic Venezuela. tribal ofsouthern Blutgruppen. infor- Indians 1967, und Unpublished C.Anthropologic1956. R.LISIKER. L. 57:1252-1259,W. 37:993, T.and M. 3.BOYD, I.ARENDS, N.CHAGNON,NEEL, Wochensehrift 1964. J. Kiinische mation, T.,G.BREWER,M.LAYRISSE, Intra- 2.ARENDS, GERSHOWITZ,and L.WEITICAMP. H. 38 CALLANGO, R.TASMAN, SHREFFLER, 4. BOYD, W. C. Genetics and the human race. 18. MATSON, G. A., and J. SWANSON. Distribution Definition of race on the basis of gene frequencies of hereditarybloodantigens among American supplements definition from morphological char-Indians in Middle America: Lacandon and other acters. Science 140:1057-1064, 1963. Maya. Amer. J. Phys. Anthropol. 63:1292-1322, 1961. 5. CHAGNON, N. A. Yanomamo warfare, social 19. MATSON, G. A., H. E. SUTTON, J. SWANSON, organization and marriage alliance. Doctoral thesis. and A. ROBINSON. Distribution of hereditary blood Ann Arbor (Michigan), 1966. groups among Indians in South America. Am. 1. 6. CHWON, B. and M. LEWIS. The ABO, MNSs, P, Phys. Anthropol. 24:325, 1966. Rh, Lutheran, Lewis, Duffy, and Kidd blood groups 20. MATSON, G. A., H. E. SUTTON, R. ETCHEVERRY, and the secretor status of the Blackfoot Indians of J.SWANSON, and A. RomsoN. Distributionof Alberta, Canada. Amer. 1. Phys. Anthropol. 11:369- hereditary blood groups among Indians in South 384, i953. America. Am. 1. Phys. Anthropol. 27:157-194, 1967. 7. CORDOVA, M. S.,R. LISKER, and A. LORIA. 21. MOURANT, A. E. The Distributionof the Studies on several genetic hematological traits of the Human Blood Groups. Oxford, Blackwell, 1954, p. Mexican populations.XII. Distribution of blood 136. group antigens in twelve Indian tribes. Amer. 1. 22. NEEL, J. V., and F. M. SALZANO. A prospectus Phys. Ant hropol. 26:55-66, 1967. for genetic studies on the American Indian. Cold 8. GARN, S. M. Human Races. 2nd ed. Springfield, Spring Harbor Symp. Quant. Biol. 29:85-98, 1964. Illinois, Charles C Thomas, 1965. 155 p. 23. NEEL, J. V., and F. M. SALZANO. Further studies on the Xavante Indians. X. Some hypotheses- 9. GERSHOWITZ, H.,P.C.JUNQUEIRA,F.M. SALZANO, and J. V. NEFL. Blood groups and ABM- generalizations resulting from these studies. Amer. Le' secretor types in the Somi5es Lopez and Sin 1. Human Genetics 19:554, 1967. 24. NEWMAN, M. T. A trial formulation present- Marcos Xavantes. Amer. f.Human Genetics 19:502, 1967. ing evidence from physical anthropology for migra- 10. GREENBERG, J. In J. H. Steward, and L. C. tion from Mexico to South America. InR. H. Faron, (eds.), Native People of South America. Thompson (ed.), Migration of the New World New York, McGraw-Hill, 1959, p. 22. Culture History. Tucson, University of Arizona, 1958. (Social Science Bulletin 27) 11. LAYRISSE, M., Z. LAYRISSE, and J. WILBERT. Blood group antigen tests of the Waica Indians of 25. PARKER, W. C., and A. G. BEARN. Haptoglobin and transferrin gene frequencies in a Navajo popu- Venezuela. SouthwesternI.Anthropol.18:87-93, 1962. lation. A new transferrin variation. Science 134:106- 108, 1961. 12, LAYRISSE, M., Z. LAYRISSE, and J. WILBERT. 26. SALZANO, F. M. Studies on the Caingang The blood groups of northern continental Caribs. Indians. I. Demography. Human Biol. 33:110-130, Human Biol. 35:140-166, 1963. 1961. 13. LAYRISSE, M., Z. LAYRISSE, and J. WILBERT. 27. SALZANO, F. M. Blood groups of Indians from Blood group antigenstudiesoffour Chibchan Santa Catarina, Brazil. Amer. J. Phys. Anthropol. tribes. Amer. Anthropol. 65:36-55, 1963. 22:91-106, 1964. 14. LAYRISSE, M., and J. WILBERT. Indian Societies 28. SALZANO, F. M., J. V. NEEL, and D. MAYBURY- of Venezuela: Their Blood Types. Caracas, Sucre, LEWIS. Further studies on the Xavante Indians. 1966, p. 72. I. Demographic data on two additional villages: 15. LAYRISSE, M., C. PEDREIRA, and J. WILBERT. Genetic structure of the tribe. Amer.I. Human Blood groups in Rankokamekra. To be published- Genetics 19:463-489, 1967. 16. LISKER, R., A. LORIA, and M. S. CORDOVA. 29. SUTTON, H. E. In E. Giblett, The plasma Studies on several genetic hematological traits of the transferrin. Progress in Med. Genetics 2:34, 1962. Mexican population. VIII. Hemoglobin S, glucose-6- 30, WIENER, A. S. Blood grouping tests in anthro- phosphate dehydrogenase deficiency and other char- pology. Amer. J. Phys. Anthropol. 6:236, 1948. acteristics in a malarial region. Amer. j. Human 31. WILBERT, J., and M. LAYRISSE. The Waran Genetics 17:179-187, 1964. Indians. In preparation. 17. MATSON, G. A., and J. SWANSON. Distribution 32. ZERRIES, 0. Waika. Die Kulturgeschiechtliche of hereditary blood antigens among Maya andnon- Stellung der Waika-lndianer des oberen Orinoco im Maya Indians in Mexico. Amer /. Phys. Anthropol. Rahmen der tieilkerkunde Sudamerikas. Munchen, 17:49-74, 1959. 1964.

39 45 DISCUSSION

Moderator: Dr. Comas' paper emphasized the As Dr. Comas pointed out, to determine which variability existing in the American Indianon of these characteristics stems from nurture and the basis of morphological characteristics. Dr. which from nature makes the problem more Layrisse has now emphasized the variability on complex. the basis of genetic traits, but has also directed However, if we accept that the original popu- our attention towards an aspect of the Indipnlation of America derived from a small group thatis increasingly occupying the thinking of of hunters-and-gatherers from eastern Siberia those of us who work with the Indian: thatwho crossed the Bering Strait during the latter within a relatively undisturbed tribe, there isa part ofthe Wisconsin glaciation and whose great deal of genetic variability from one partdescendants gradually became masters of an of the tribal distribution to the next. The geneti- uninhabited continent, we can understand bet- cist has tended in the past often to think ofa ter the great influence of environmental factors tribe as a kind of homogeneous beanbag, butupon the 'American Indian. There isstillthe it now becomes clear that thisis not the case question of contact with other racial groups and that there may be very significant differences during the pre-Columbian era, especially with between, say, one part of the Yanomama terri- Caucasoids of European origin as mentioned by tory and another. Dr. Comas. The discussion of these twopapers will be The great progress made lately in the study of opened by Dr. Arends. blood groups, serum proteingroups, and enzyme polymorphism nullifies the idea expressedsome Tulin Arends years ago: "Type one Indian and you have typed them all; so much are they alike in their blood The physicalvariabilityofthe American groups."2 The view that serology has not added Indian was recognized even by the conquista- anything new to human taxonomy2 isalso no dors. In 1530, Federman described a tribe of longer tenable. On the contrary, we who work dwarff in western Venezuela "not taller than in this field believe that the amount of informa- five to six handbreadth" and found the Guay-tion already accumulated wit', play an important queri in the eastern part of the country to be rolein the reinterpretation of the American it peopleasblackascoal."' Anthropometric Indian taxonomy and is laying thc foundations studies have accumulated a great many data on for a future molecular anthropology. Boyd has stature, cranial and cephalic indexes, body pro- portions, and so forth, which multiply these said: "The genetic method has, in short, made contributions to anthropology that morphologic differences. If to the somatic characteristicsare added.he many inheritedbiochemical and methods could hardly have made, and ispro- hematological variations known today thedi- versity of the American Indian becomes infinite. 2 Stewart, T. D. A physical anthropologist's view of :he ). of the New World. Southwest. 1, Anthropol. 16.)9 1960. 1 Federman, N. Historia indiana. Trans!.J.Fricde. ..: S. M.: Human Races. 2na ed. Springfield, Madrid, Aro, 1 .;harles C Thomas, 1965.

40

46 viding a new and fundamental basis for the the dynamics of indian populations, could ex- study of race."4 plain some odd findings in the prevalence of genetic traits. Important factors in the study of the American populations Common characteristics in the American Indian To establishbiological subdivisionsofthe From a genetic standpoint, most of the Ameri- American Indians based on the study of theircan Indians share certaincharacteristicsthat genetic chiracteristics, the best-known methods differentiate them from other racialgroups populatioa genetics should be used, with(Table 1), as Dr. Layrisse has already men- special emphasis on the following points: tioned. We may say that the authen South 1. Itis advisable to study as many genetkAmerican Indians are all of group 0; usually traits as possible, in order to avoid biased inter-lack r and R° rne complexes of the Rh system; pretations and rule out gene drift, especiallyare negative for the main factor of Kell, Lewis, when dealing with small populations subjected and Lutheran blood group systems; have only to epidemics, wars, displacements, and otherHb-A; and are not deficient in glucose-6-phos- events that might have modified the frequency phate dehydrogenase.8 It could be added that of some traits. they are equally negative for, or have a very low 2. Indian villages should be studied inten- frequency of, the atypical pseudocholinesterase sively from a historical and genealogical point gene Eia.9)10 of view, for the purpose of evaluating the so- called "founder effect" or pursuing a foreign the Xavante Indians. X. Some hypothesis-generalizations resulting from these studies. Am. 1. Hum. Genet. 19: gene introduced through a nonapparent ad- 554-575,1967. mixture. Mourant, A. E. The Distribution of the Human 3. The "fission-fusion" phenomenon, which Blood Groups. Oxford, Blackwell, 1954. s Arends, T. Haetroglobinopathies, thalassaemia and according to Neel et al.,5, 6 is not infrequent in g1ucose-6-phosphate hhydrogenase deficiencyinLatin America and the West Indies. New Zealand Med. 1. 4 Boyd, W. C.: Genetics and thc human race. Science 65;831-844,1966. 140:1057-1064,1963. Gutschc, B. B., E. M. Scott, R. C. Wright. Hereditary 5 Ned, J. V. F. M. Salzano, P. C. Junqueira, F. Keitter, deficiencyofpseudocholinesterase in Eskimos, Nature and D. Maybury-Lewis. Studies or thc Xavante Indians 215:322-323,1967. of the Brazilian Mato Grosso. Am. 1. Human Genet. 10 Motulsky, A. G. and A. Morrow. Atypical cholines- 16:52-140,1964. terase gene Eia: rarity in Negroes and most Orientals. Neel, J. V., and F. M. Salzano. Furthertudies on Science 159:202-203,1968. Le lima n1)-A bsenee Arcwas, T 0.4 .1Vvies,31\ Jorizurv44 tipt-tx.fr, Pt4000nSr at TABLE I. Characteristics common to American Indians ca.c.y WIi Ikea 1)-c 4 h ia se) Sotkiit AVM riCab '4tiN4rNe-A. AtIN0 Ob-Ct_ ict67.

CHARACTERISTICS COMmON TO ALL GENETIC SYSTEM SUBJECTS EXCEPTIONS

ABO Group 0 Navajo, Pueblo, Blood, Shoshone, Blackfoot (U.S.A.) Rh r and R° absent Apache, Pawnee (U.S.A.); Tarasco, Chontal (Mexico); Itza Maya (British Honduras) Kell kk Lewis Le(a) Lutheran Lu (a) Dieguerio (U.S.A.); Brazilian(Brazil) Hemoglobin Flb-A Nahua, Chiapas (Mexico); Oyana, Arawak (French Guiana) G-6PD Absence of deficients Ciana, Carib (Dutch Guiana)

41 Where published reports give different char- greatly upon the condition of the sample ob- acteristics from those in Table I, the reason istained. In addition to the exceptions mentioned mainly the small size of the sample studied (in by Dr. Layrisse, there are tribes that have given some cases fewer than 50 individuals) or ad- frequencies different from those mentioned even mixture with other racial groups. Nevertheless, though the sample was larger than 100. Again, there are some tribes for which the results alsohere are many populations that could be very disagree although they are unmixed and a large usefully studied with the available genetic, bio- enough sample was examined. These popula-chemical, and hematological battery of tests. tions deserve to be thoroughly studied for a good explanation of this exception, and tilisDifferential characteristics of some Indian tribes could be the so-called heterc of American Finally, there are some genetic markers that Indians. (The same may be sd., of the figuresidentify certain tribes (Table 3). The antigen in Table 2.) Di(a+ ) of the Diego system, for instance,is Quantitative differences in the genetic systems a Mongoloid characteristic. The same may be of the American Indian said of the transferrin Doit. The absence of those markers in many tribes could be an indication In certain genetic characteristics the American of gene drift, fission-fusion, or founder effect. Indian is distinguished only by a quantitative Furthermore, the transferrin system has another frequency widely different from that of other variant (Tf-B11_1) that distinguishes the North racial groups (Tab It 2). The interpretation here and Central American from the South American is really difficult b,:cause different samples from Indians.12 the same population have sometimes given dif- Human alloalbuminemia, or the presence of ferent results. For example, when the hapto- albumins will electrophoretic miglation differ- globin system of two Makiiitare samples wasent from the no-mal, expressed by either one studied," results for Hp' as dissimilar as 0.610 and 0.342 were found. These results depend ia Arends, T.. G. Brewer, N. Chagnon, M. L. Gal- lango, H. Gerashowitz, M. Layrisse, J. Ned, D. Shreffler, 11 Arends, T. Los grupos s6ricos humanos determina- R. Tashian, and L. Wietkamp. Intratribal genetic differ- Hes por métodos electroforéticos: su distribuci6n en Sur entianon among the Yanomama Indians of Southern Am&ica. Acta Cient. renez. SuppL 3:162-178, 1967. Ventzuela. Proc. Nat. Acad. Sci. 571252-1259, 1967.

TABLE 2. Quantitative differences in some genetic systems of AmericanIndians

GENETIC SYSTEM CHARACTERISTIC EXCEPTIONS

MNSs Gene M '5 0.7 Jicague (Honduras) Rh Gene complex R2 > 0.35 Seneca, Pima (U.S.A.); "league (Honduras); Miskito, Sumo (Nicaragua); Mayal Mazatel (Mexico); Ticuna (Peru): Quechua (Ecuador); Xavante (Similes Lopes, Brazil); Waica (Venezuela) Duffy Gene Fyn 5: 0.6 Guavmi (Panama); Jicaque (Honduras); Mazatel (Mexico); Xavante (Brazil) ABH Secretor Gene Se Xavpnte (Brazil)

Haptoglobin Gene Hp' 0.5 Yupa (Venezuela); North Athabascan, Tlingit, Navajo (U,S.A.) Gammaglobulin Gene Gni° ,= 1.0 Xavante (Brazil) Inv Phenotype Inv(a+) 5: 0.6 Gc Gene Ge1 > 0.65 Xavante (Brazil)

42 48 TABLE 3. Genetic markers in some Indian tribes

GENETIC SYSTEM CHARACTERISTIC TRIBES IN WHICH IT HAS BEEN FOUND

Diego Di(a+) North American, Central American, South American Transferrin Tf Navajo (U.S.A.); Yaqui, Cora, Chinantec, Lacandon. Tzotzil, Zapotec (Mexico) Quidie (Guatemala) Tf Irapa, Macoita, Paraujano, 111,64 Shaparu, Monk), Piarea, Warao, Guahibo, Guayqueri (Venezuela); Paez (Colombia); Quechua (Peru)

Albumin Naskapi Naskapi, Montagnais (Canada); Sioux, Athabascan (U.S.A.)

Mexico Zapotec (Mexico) Maku Maku (Brazil) Warao \Vara() (Venezuela)

(homozygous form) or two components (het- 3. Some genetic markersfor example, Di erozygous form), is a system that could be very (a+) group of the Diego system, transferrin useful in the American Indian taxonomy. This Dem and B01, and albumin Naskapi, Mexico, system, which at present is the subject of inten- Maku, and Waraohave been found in only sive study, has made it evident that certainsome Indian tribes. Thecharacterization of each tribes in Canada and the United States have an of them and their differentiation from other albumin variant faster than normal (albumin traits of the same system should be determined Naskapi). In Mexico a tribe was found to have by the best biochemical and physico-chemical an albuminslowerthannormal(albumin methods available. Mexico). In Brazil a faster variant (albumin 4. The finding of some genetic traits (trans- Maku) and in' Venezuela a slow variant (al- ferrinDchl,Diego group Di(a+ ))among bumin Warao) different from those previously tribes is positive proof of present or remote rela- mentioned were found. tionship. Their absence in other tribes could be explained by gene drift, founder effect, or fision- Summary fu s ion. I. Some characteristics are shared by all the 5. The more genetic systems are discovered, Indians of the American hemisphere. Popula-the more urgent it is for them to be studied in tions that seem to be an exception should be American Indianr before these populations dis- investigated by the best serologic, immunologic, appear through cultural andeconomic pressures electrophoretic, and chromatographic methods or through epidmic,endemic, or sporadic at hand in order that characteristics reportedin pathological processes. the past, when modern methods and reagents were not available, may be rechecked. Ruben Lisker 2. The differentiation of populations by fre- quency gradients of genetic traitsshould be the The data presented today by Dr. Layrisse subject of careful studies, about their formation, clearly show that present-day Amerindian popu- cultural level, and kinship organization. Sincelations do not have a homogeneous genetic these populations have been under physical, constitution. However, the task ofsubdividing pathologic, and war pressures, it will always be them on the basis of genetic traitsisnot an very difficult to evaluatehow much these agents easy one, especially if to taxonomyis added an have influenced the distribution of the inherited interest in interpreting or understandingthe variations. reasontor the differences encountered.A gen-

43 49 era! cause of this difficulty is that the Amer-TAI1LE 1. Frequency of Diego antigen in18 Indian indians as a group have characteristic gene tribes and 9 coastal mestizo groups of Mexico frequencies for many traits that do not allow further subclassification. A good example would PHENOTYPE FREQUENCY GROUP RANGE (%) (70) be blood group 0 and the Rh-negative pheno- type. A reasonably "pure" Indian population Indiansa should have close to 100 per cent of group 0 5 tribes 2-8 9 tribes 12-19 individuals and no Rh-negative subjects; devia- 3 tribes 20-24 tion fromti,ispattern would be generally 1tribe 40 interpreted as the result of non-Indian admix- Mestizos ture. WeSt Coa.tit Even so, some g..netic traits do show enough Ometepec 28.2 variability in apparemly unmixed Indiangroups Cuaiin'cuilana 26.3 that attempts at taxoucmy are possible. The San Pedro M. 23.4 Pochutia 31.3 difficulty hereand I should like to stress this East Coast pointis what the classification means. Ifeel Saladcro 4.1 sure that this considerati3n explains, at least in Veracruz 32.8 part, Dr. Layrisse's understandable hesitation in Paraiso 32 A subdividing Amerindians. El Carmen 38.6 I want to consider briefly some examples of Tamiahua x .5 the difficulties I have mentioned. The Diego More than 50 individuals studied in ailbut two antigen appeared to be an excellerrt marker of tribes. Indian groups, especially suitaFte for taxononii- b Mare than 90 individuals studied in each location. cal purposes since it is not present in other ethnic out of the iiine hybrid groups studied on the groups (except some Mongoloids) and showscoasts of Mexico have higher Diego frequencies enough intertribal variability to allow for sub- than most of the pure Indian tribes. Before divisions.1 However, it soon became apparentlearning the meaning of this phenomenon,one thatthe correlation ofitsdistribution withwould hesitate to use this marker to subdivide either geographicalor cultural critena was not Amerindians. Incidentally,I would saythat too good,2 and furthermore, although itsab- although these populations show less variability sence in Negroes and Caucasians is accepted,than those Dr. Layrisse has studied, there is still some hybrid populations in which they certainlyno pattern to Diego frequency because individ- contribute to the gene pool have higher Diegouals with the same linguistic affiliation show frequencies than the original Indian popula- very marked differences in frequency. tions.2. 4. 5 This is clearlyseen in Ta lie 1; seven An additional difficulty stems from the fact that in the study of Indian tribes withsome 1 Layrisse, M., arid J. Wilbert. El antigen° dd sistema degree of non-Indian admixture, the variation sanguinco Diego. Caracas, Sucre. in the gene frequencies of 'ome of the traits 2 Comas, J. Significado de. la presencia del antigeno does not follow the expected trend The follow- Diego entre lo.s Amerindios. Anal. Antropol. (Mexico) 2:89, 1965. ing examples will clarify this statement. In the 2 List:el-, R., A. Loria anti M. S. Cordova. Studieson investigation of Mexican Indian groups, four several genetic hematological traits of the M:xicanpopu- gens in Nahuas, Yaquis, Tarahumaras, Tarascos and lation. VIII. Hemoglobin S. glucose-6-phoiphate dehy- Mixtecos. linm. Biol. 35:359, 1963. drogenase deficiency, and other characteristIcs ina ma- .5 Cordova, M. S., R. Lisker and A. Loria: Studies on larial region. Am. 1. Hum. Genet. 17:179, I 965. several genetic hematological traits of the Mexican popu- 4 Rodriguez, H., E. Rodriguez, A. Loria, and R. Lisker: lation.Xli. Distributionof blood group antigens in Studies on several genetic hematologicaltraits of the twelve Indiantribes. Am. 1. Phys. Anthropol. 2655, Mexican population. V. Distribution of blood group anti- 1967. /

44 50 had evidence of Negro admixture, as judged by the slow variant known as albumin Mexico 10 the presence in them of a few individuals with was present in southern North American In- the sickle-celltrait and/or redcell glucose-6- dians and in some Mexican Indian groups and phosphate dehydrogenase (G-6-PD) deficiency.° mestizos. The only information from South These groups would be expected to have higher America relates to two groups of Peruvian In- frequencies of the M gene and the cDe chromo- dians, another from Brazil, and a fourth from some and lower of the Fya gene than the other Brazil and Venezuela (the Yanomama), where tribes. As can be seen in Table 2, neither of no variantswere found exceptinthelast these expectations was confirmed.5 It would be group.n This pattern has already at least three also logical to expect the Hpl gene frequenciesexceptionst the Huasteco in northeast Mexico of the Indian tribes with evidence of Negroand the Chol in southeast Mexico have a fast admixture to be higher than in the I-est. Table variant that has electrophoretic mobility similar 2 shows that thisis not the case, and that in to the Naskapi type and may be the same, and fact the Cora have the lowest Hpi gene fre-the variant recently encountered in the Yano- quency in our material.' mama is fast bu different from the Naskapi. The recently described polymorphism of the It is also interesting to remark that whereas albuminlocus seemedtohave taxonomical the presence of albumin variants in Eskimos value, since in all the initial reports it followed might lead to an attempt to group them to- an orderly geographical distribution.° Albumin gether witil other Indian groups, itis evident, -,-ariants are rare in all groups so far studiedas stated by Dr. Layrisse, that the only Ameri- CACt pt the Amerindians, and it appeared that a can aboriginal population that clearly deviates fa,l- variant known as albumin Naskapi 9 was from the rest of the Amerindians in the distribu- prevalent only in Eskimos, Canadian Indians, tion of several genetic traits is the Eskimos. and northern North American Indians, while It seems to me that, regardless of the rather erratic distribution of some traits in the Amer- 6 Lisker, R., C. Zárate, and A. Loria. Studies on several indians, and despite the fact that the precise genetic hematological zrait., M Hcar2 .IX.Aho.-:7-rnal reason for this is unknown, itis important to hemoglobins and erytlm,(N 0,1!..-ose-6-phcmpi,:irt t!ehy- colitinue the characterization of the genetic con- drogenasedeficiencyinsevv...! trib:..-, Blood iolou of the Arnerindians, quite apart from 27:324, 1966. 7 Lisker, R., G. Zaratc, and E. Rodrigu,.;'.. the ciu.tstion whether or not it can be used at several genetic hematological traits of the Mexican t+,,7,!- this1,ointfor meaningfulst,bclassification. lation. XIV. Serum polymorphisms in severalIndia, tribes. Am. I. Phys. Anthropol. 27:27, 1967. Mciartio. 1, 81,:.-nherg, and R. Lisker. Albumin 8 Melartin, L. Albumin polymorphism in man. Studies Mexico: -aiE alarr3 i5: on albumin variants in North American native popula- 288, 1967. tions. Acta Path. Microbiol. Stand. Suppl. 191, 1967. 11 Weitkamp, L.,J. Chagnon,I.Saave, F. Salzano, 9 Mdartin, L., and B. Blumberg. Albumin Naskapi, a and J. Gall. Serum albumin variantsin American and new variant of serum albumin. Science 153:1664, 1966. New Guinea indigenes. Clin. Res. 16:298, 1968.

TABLE 2. Comparative frequencies of genes M, Fya, Hp1, and chromosome cDe+cde in groups having some cases with Hb.S and bor 0-6-PD deficiency with other tribes investigated for these markers

NO. CASES GENES CHROMOSOME Hp1 GROUP NO. Hb.S G-6PD Fya cDe cde GENE

Huastcco 235 o 3 0.51 Nahua 141 o 4 0.77 0.64 0.04 0.44 Chontal 101 2 2 0.69 0.67 0.11 0.57 Cora 96 1 0 0.76 0.99 0.06 0.40 Others (range) 0.650.79 0.50 1.00 0.03 0.12 0.440.65

45 51 offer at least two important reasons: one, that inthefirst. The third stage would he that this will provide the geneticist of tomorrow with reached by tribal groups in Mexico. The genetic a good baseline for understandingpopulation structure would also be different then; it would dynamics in the future; the other, that longi- be more likely to be the one that Sewall Wright tudinal study of the correlation between the calls the isolation-by-distance model. This means genotype and the biological development of a that the genetic diffeeentiation of these groups population, for which atleast some Indian would follow a different pattern from the other groups are quite adequate, mightsignificantly two, with less variability. In a general way, I contribute to our understanding of the reasons believe the genetic and demographic data are for the variability observed today. substantiating this. But, of course, in order to be stoi: we need more data on different groups, General Discussion in different ecological situations. I do not thir,k that genetic data per se would Bier:Iwouldliketo know whether the albumin variants are antigenically identical. bevery important for thedifferentiationof Lisker: When I say there are two iticrikital Amerindians unless they were correlated with albumins this kind of data. For 'instance, the selection I am only saying that they have the sameacting on groups in the second stage is probably eleetrophoretic mobilky. Nobody has done an quite different from that acting on the other amino acid sequence, so that is all we know.two. This was clearly indicated by Dr. Dubos Whether or not itis the same mutation is un- inhis book Man Adapting. The nutrition of known. these groups would become somewhat more Bier: Before doing the sequence, wouldn't itlimited; they would rely on just a few crops be easier to make an Ouchterlony plate to see that can provide enough .calories but make for whether or not they have common determi-an unbalanced diet.I could give many other nants? such examples to show that the situation be- Lisker: It has been done, and no differences tween groups inthesethree stagesisquite have been found. different and that if data from populations at Salzano: I think that the studies on Vene- thesedifferent culturalstagesaremixed,a zuelan Indians in different stages of cultural meaningful picture is probably impossible. evolution, and a!so the researches of Dr. Lisker Waterlow: Dr. Neel, isit really proved that in Mexico, are very important since a patternall these thingsalbumin variants, haptoglobin appeared among the Brazilian groups that can variants,red-cell variantsare genetically de- be tested. Three stages can be visualized. The termined or is that an assumpdon? first would be that of the hunting-gathering Moderator: I believe the speakers could take tribes, who have a kind of fission fusion model. care of this point better thanI could, but very That is, the pattern is one of exchange between briefly the answer is yes, it is proved. However, groups carried out mainly by more or less large I should qualify this a bit. The genetic basis subgroups of the tribes. In what I would call cannot be determined from a simplepopulation the second stage, the groups would rely moresurvey Family studies arerequired. And in on agriculture hut woukl not yet have a critical everyinstance in which avariant has been size. These vs-uuld conform inore to what Sewall describedtoday, there have been svidies that Wright calls the island model: -thatis,their show it is tranyrt;tted accordi3-3- !.o del;: !- copulations would be small and separated by laws within familylinf:c. thi...kirrj large distances. A prier', more variation would evidence is necessary befort pK ot perhaps be expected in the second groups than variant with any assurance. THE AMERICAN INDIAN IN THEINTERNATIONAL BIOLOGICAL PROGRAM

James V. Neel

It will be Administrative background four other countries may do so soon. the purpose of this presentation to attfmipt to 1960 As is a matter of common knowledge, in point out the manner in which wriPt\ f-f types Unions the International Council of Scientific of studies of the Americat within (1CSU) took the initiative in the promulgationthe framework of tne ITU' of an International Biological Proram As e. section concenicd with humanadaptalAity. In obrctives of this inally eovisioned b\ fact, the Indian provides the Americaswith a 'dt oi- program wt rethe world ( unique resource in IBP-orientedstudies and at ganic production on the land, in fresh 'voters,the same time presents interdigitatingmedical and uses and in the seas, and the potentialmes and humane challenges that areworthy of the and of new as well as existing natural resources; consideration of this Organization. 12) human adaptability to changingconditions. consideration of ICSU in At all the various kvels of A Planning Committee appointed by human adaptability, it has been clearthere are turn established a numberof :Aibcommittees to two interrelated components:the genetic, which consider various aspects of the program and to sets the limit of theorganism's ability to re- elaborate a more extended protocol. These van- spond, and the physiological, whichdetermines combined their efforts into subcumm the range of response available to theorganism, ind recommendations, a stries of sogw:.tions given its genetic set. At one extreme inthe spec- which the Pianning C.:Ayanitte,.: coordinated and be brought to bear on submitted to It:.;U in Novemix; I963. ICSU, trum of interests that can human adaptability is environmentalphysiology, ;IAC.Irecom- after approving these sligg:.:ions in mendations, then called ti,,ori the vAdOliS cuun- and at the other is population geneticsand tries having representation in it throughnationo,J isveen thereis room for many aspects of societies to formulate national programs.The modern medicine and anthropology. iimetable for IBP, several times revised, consist. Prom the earliest considerations of mostof of a "planning phase," whichofficially termi- the various committees andsubcommittees con . nated in mid-1967, and an "operationalphase," cerned with human adaptability,ithas h.:er to continue untilmid-1972. apparent that the survivingprimitive pope,"a- It has been clearly understood fromthe outset dons afford an opportunity to studyphysiologi- that the various national committees werein nocal and genetic adaptationspotentia:lyquite way bound bythis international formulationit different from those of highly civilizr.1. gri ups. was designedmerelytoprovide convenientParenthetically, although our cultwal oricrita guidelines. Thus far ten countries inNorth, tion is such that we tend to view man:, oftliese Central, and South America haveestablished surviving primitive groups as representing ex- committees to formulate national programs,and tremes of adaptability, thefact is !.1:at they haw 3 lived under their conditions far longer thanwe of these problems has reached sucha magnitude under ours. Viewed from their vantage point,as to call for concerted action on ahitherto we are the ones pressing for new limits of adap- unknown scale. But I submit that behind these tability, and it is only an article of faith thatwe immediate problems lurks a far larger issue. will adapt to our man-made environmentas Now that man so largely controls both theen- successfully as they have done to theirs. We must vironment and his reproduction, is thereany note in passing that some of the adaptations of ng-range objective more important than that these groups are complex and ingenious,ar u.h,l..rstand the genetic potentialities of the that the term primitive as commonly tificci and the species, the manner in which something of a misnomer, amounting to dpri this potential interacts with an environment that literate with a relatively simple technology." is increasingly man-made, and the rate of genetic In late 1962 the World Health Organization change in response to a changing environment? convened in Gene\a Scientific Group on Re-You will recognize thatIhave only restated search:il'oi ilationGenetics of Primitivein a different context the need to understand Groups. its purpose was to consider studies ofthe genetic aspects of human adaptability. The populations of unusual genetic interest, withpar- Amerkan Indian offers prime research material ticular reference to the surviving hunting andfor this objective. The principal lines of attack food-gathering groups of the world, to advisecan be conveniently summarized under three on themostsuitablepopulationsforsuch headings (see 12). studies, and to draw up appropriate research 1. Population structure and natural selection designs. The pertinence of this to IBP-relatedin primitive man. One of the basic tenets of developments is obvious. In the preliminary in- biology is of course a belief in the role of natural ventory of these societies published in the reportselection in shaping each species to its ecological of the Scientific Group, various South American niche. As biologists we believe that natural selec- tribes figure prominently (26). Thisreport, inci- tion has guided human evolution and willcon- dentally, was updated in thesummer of 1967 tinue to guide the evolution necessary fora suc- and a revised version has just been issuedas cessful genetic adjustment toa changing world. WHO Technical Report 387. It is clear that modern man evolved under cir- A recurrent consideration in the consultations cumstances approximating the conditions found of both IBP-related and WHO-relatedgroups in primitive cultures. Otherwise stated, there is has been the rate at which these primitivegroups no evidence that the developments of the past are disappearing. It can be said without exag- few thousand years have yet resulted inany very geration that ours is probably the last generation significant modifications of man's geneticpo- privileged to study suchgroups in a relatively tential. It follows thatone way of understanding pristine state. If there are important scientific the genetic impact ofour chunging environment lessons to be learnedand the WHOdocument is from the baseline supplied by detailed knowl- would seem to develop thatcase adequately edge of primitive man in his primitiveecosys- then there is an urgency to this problemnot shared by many other areas of research. tem. The simple fact is that at present we know virtually nothing about how selection works in The dimensions of Indian-oriented studies man. What few insights we have are largely These are days in which those ofus who are based on the results of artificial selection with concerned with human biology viewour species experimental animals and deal with traits, such as beset by a growing number of pressing prob- as extremes of egg production or milk yield, lems of our own making: the population in-considerably more important tous than to the crease, trace chemicals in foodstuffs, air pollu- animal species involved. tion, water pollution, the so-calledstress diseases, What the geneticist terms "populationstruc- cultural alienation, to mention onlya few. Each ture" places limits on the effectiveness of natural

48 54 or artificial selection. Thus, hand inhand withcians. Although the precise nature of the data efforts to understand selective pressures must gocollected depends upon local circumstances, un- efforts to understand the genetic structure of der optimal conditions we assemble as complete populations. To the geneticist, this means pat-demographic and genealogical data as possible; terns of mating, fertility, and survival; amountconduct physical, dental, and anthropological of inbreeding;migrationpatterns; mutationexaminations; and obtain blood, urine, saliva, rates; and so on. Although some data on these and stool specimens for a variety of genetic and subjects are to be found in the voluminous an-other determinations. The effort to understand thropological literature,inthe main what isbetter man's interaction with his environment available simply lacks the precision and depthcarries far beyond the identification of genetic necessary to the precise statistical treatmentsof traitswe are fortunate to have the collaboration the geneticist. of such groups as the Communicable Disease Even with all the facilities of modern scienceCenter of the U.S. Public Health Service, the and the conveniences of our culture, such com-Virus Institute at IVIC, and many specialized prehensive studies remain extremely difficult in laboratories at the University of Michigan. civilized countries. How much more difficult This is not the time for a detailed review of are they, then, in the remote primitive popula- the findings on either the Xavante or the Yano- tionsproblems of logistics and communication mama, which in any event must be regarded as being what they are. Even so, from our ownpreliminary. Not at all surprisingly, both the experience of the past several years and that ofpopulation structure of these people and the several other groups with kindred interests, it biological pressures on them are qualitatively is clear that a variety of types of data relevantand quantitativly different from those of civi- to population structure can be collected. lized groups. These are populations in which In1962, aided by a grant from this Organiza- infertilityisrare,practically all women con- tion, Dr. Francisco Salzano of the University oftributing to the next generation; in which effec- Rio Grande do Sul, Brazil, and I, with several tive fertility seems intermediate, and infant and collaborators, undertook apilot study on thechildhood mortality also intermediate; and in Xavante of the Brazilian Mato Grosso (13). which the variance of male reproductive indices These and further studies on the Xavante, in is significantly greater than of female (15). A 1963 and 1964, raised a number of rather basicsurprising amount of genetic microdifferentia- questions for population genetics (14), but it tion between villages has been encountered (1), was felt that the Xavante were neither a large originating, we believe, in the fission-fusion pat- enough group nor sufficiently undisturbed for a tern that dominates village proliferation and critical approach to these questions. Accordingly, results when a new village comes into being in after a careful consideration of the possibilities,a very non-random sampling of the tribal gene a second study was undertaken, concerned with pool. We reason from this that if new tribes the Yanomama Indians of Venezuela and Brazil originate in general as new villages do, then and involving as principal colleaguesin addi- chance may have played a greater role in much tion to Dr. SalzanoDrs. Miguel Layrisse and measurable human genetic variation than here- Tulio Arends of the Venezuelan Institute fortofore seemed likely. The roles of traumatic, Scientific Research (IVIC) and Drs. William J. infectious, and nutritional disease in determin- Schull and Napoleon Chagnon of the University ing survival are far different than in the next of Michigan. We have now been in the field on step on the cultural ladderthat is, in relatively three different occasions. The composition of the densely populated agricultural societies. team varies from year to year, hut ideallyit We do not pretend to understand as yet the includes an ethnologist, a linguist, a physical full genetic implications of all that we see. One anthropologist, a geneticist, and several physi- reason for our lack of understanding is that a

49 55 substantial portion of the mathematical theory each may be unusual in one or several respects. of population geneticsiscurrently being re- Only from the perspective of numerous studies worked. A second reason is that the data are so in depth can we begin with any assurance to complexthatprecisemathematical formula-factor out the significant common denominators tions do not appear imminent, and we are beingthat have guided human evolution. Already forcedto involved computer simulation pro- parallel studies are in progress on such groups as grams to explore the genetic consequences of the Cayapti of Brazil by Salzano and colleagues the differences between these societies and our and the Kashinawa of Peru and Brazil by Johns- own. However, even at this stage it is very clearton and colleagues, as well as in other parts of that it will be possible to define objectives and the world, and more are to be hoped for. test hypotheses in a way that will justify the 2. The rate of human evolution, i.e., genetic effort expended. diversification.Despite the many unanswered I mentioned above that a substantial portion questions discussed earlier inthis session by of the mathematical theory of population ge-Professors Griffin, Cruxent, and Laughlin, the netics is being rt worked. One of the most pro-date and place of arrival of the Indi=th in the vocative developments in the biomedicine of the Americas is more accurately known than the past twenty years has been the realization of the corresponding information for any other major extent of man's concealed genetic variability, as subdivision of mankind. In view of what can be revealedby modern biochemicaltechniques. deduced concerning population density in north- Many of the newly recognized variant typesern Europe 20,000 years ago, we may surmise occur in populations in such frequencies that we that the total number of immigrants across the speak of the underlying systems as genetic pu!:,- Bering Bridge, whether they came in one or in morphisms. The maintenance of suchpoly- several discrete waves, was not large. The ge- morphisms is thought to require selective pres- netic characteristics of this original group will sures of one kind or another; not only is theof course be forever conjectural, but let us as- nature of these forces unknown, however, butsume that thetribalpopulation(s) concerned under the previous formulations the mortality had the rangt. of variability we see in such a and fertility structure of our population scarcely group as the Yanomama. (I am assuming in seems adequate to accommodate the necessary this discussion that the contribution to the gene selection, and new formulations are being con-pool of individuals reaching the Americas other sidered (7,9,25). Understanding how all thisthan by the Bering Bridge is negligible.) Once variabilityis maintainedis one of the great here, extension throughout the Americas seems challenges to the human biologist (see discus- to have been rapid. In this extension, the diver- sions in 8 and 16). Now, the genetic polymor- sity of habitats explored and occupied represents phisms are well represented in all the primitive most of the extremes of the habitable portions of populations studied to date. This is where they the earth.If we make the assumption of a came into being. Accordingly, it seemsself- reasonably homogeneous beginning, then the evident that, despite the inherent difficulties, we diversity of types of Indians at the time of the must make every effort to conduct studies in rediscovery of this continent five centuries ago depth of population structure and the relation- constitutes an unusual opportunty to measure ship atthis cultural level between the poly- the rate of human evolution. morphisms and specific biological attributes. Much of the early thinking on this topic drew There is an urgent scientific need for similar on morphological characteristics, and these, as studies, while the opportunity exists, on as many Professor Comas hasjust demonstrated, still as possible of the other unaccultUrated groups of remain valid and pertinent. However, in recent Indians. None of the surviving primitive Indian years the newly discovered polymorphisms men- communities can safely be labeled "typical" tioned earlier have provided a powerful battery

,50 of objective indicators of genetic distance. Dr. contexts far greater interest than the data on Layrisse has just provided us with an overview mixed, an effort has been made to recognize of how great the range is with regard to a threelevels of miscegenation: (1)essentially variety of specific systems. In theory, for any none; (2) known, but probably accountingfor given genetic system the position of each Indian less than 5 per cent of the gene pool; and (3) population with respect to all the others can begreater than 5 per cent. The tabulation contains defined by a point in a space that has one lessdata onthe following polymorphic systems dimension than there are alleles in the system, (these being the ones on which there seemed the precise position in this space being deter-enough data to warrant a tabulation): ABO, mined by the frequency of each allele. FurtherRh, MN, Kidd, Duffy, Diego, P, Lewis, hapto- in theory, when many different genetic systems globins, transferrins, and Gm. Gene frequencies are considered simultaneously,the position ofhave been computed for each study with com- each population vis-a-vis the others can be plot- puter programs written in Fortran andavailable ted in a space that now has as many dimensions on request. as there are genetic systems.Previously, the com- In closing this section, let me agree with those putations when a considerable number of sys-who feel there has already been quite enough tems was involved were aslaborious as this of surveying populations for gene frequencies in description sounds (6, 22). But the advent of thethe vague hope that someday the information modern computers has dramatically altered the will be valuable (see discussion in 2). In the situation, and now a number of groups are de-case of the Indian, however,such surveys bear veloping computer-based measurements of popu- on a clearly defined problem onwhich consider- lation distance (see 3). able progress may be expected in the near future. Although considerable progress has been made, 1 The response of genetic systems to a rapidly there is still a great deal to be done in the waychanging environment. As I mentioned earlier, of characterizing the genetic attributes of thewe recognize that our species isall over the relativelypure Indian groups, as a basisfor world subjected to a rapidly changing environ- exploring the evolutionary problem discussed ment. Unfortunately, only in the last few years above. Incidentally, the HA/IBP/IUBS empha-have biology and medicine found themselves in sizedthe need for continuing effortsatthe a position to begin todocument with any pre- genetic characterization of all the human strains, cision what biological adjustments really take so that this type ofinvestigation of the Indian place. We have probably lost forever the oppor- fits readily into a larger framework. The pub- tunityto study what occurred in Caucasian lished data on the Indian are widely scattered, populations. We are about to lose the opportu- no survey having beenattempted since those ofnity to document what happens in Negro and Mourant (10, 11) and Salzano (18). Dr. Rich- Mongoloid (including Indian) populations. ard Post, Dr. William Schull, and I have now The transition of surviving primitive Negro completed a compilation of all publications prior or Indian groups willbe telescoped into a far to November 1967 that deal withthe prim- ipal shorter time period than was true for Caucasian genetic polymorphisms known to occur in the groups. Thus, some Indian groupswill be called Indian.1 The rate of discovery of new poly- upon to move from the StoneAge to the Atomic morphisms being what it is, any such compila-Age in a few hundred years, whereas the transi- tion is soon dated, but for the specialist it at tion for most Caucasian groups took thousands. least provides a nidus for updating. The 271 It seems reascliable to argue that here is an tabulation is based attest papers on which the opportunity to study changesinpopulation Since the to the strong interest in this subject. structureand adjustments to new stresses that data on unmixed Indian groups have inmost we must not lose. ' Sec Appendix of this volume. As for the previous two sections, so here the

5157 intellectual challenge isto convert an easily Some moral issues enunciated but somewhat diffuse series of prob- It has seemoi appropriate, ina presentation lems into speciiic issues susceptible of precise to an audience of scientists, to stress research study. A number of exact studies of population opportunities. But as in the 1960'swe increas- structure intransitional groups have already ingly recognize the issues created by scientific been completed or are in progress, notably those inquiry divorced from ethical and humanitarian of Salzano and colleagues on the Caingangf considerations, it also seems appropriateto con- Brazil (19, 20), of Layrissand colleagues on sider briefly what these st:zdies, and especially the Warao of Venezuela, of Covarrubias and the type mentioned in the last section, might colleagues on the Pewenche of Chile, and of mean to the Indian. We have no accurate cen- Cann and colleagues on certain Quiche speakers sus of the relatively pure Indians left in the of Guatamala. There are also studies in the Americas, or of the persons of mixed but "sub- United States on groups ina more advanced stantial" Indian ancestry. Estimates of the order stage of the transition, namely by Spuhler and of 16,000,000 (17, 20, 24) have been made for colleagues on the Navajo (21) and Hackenberg the former, while the latter come to easily on the Papago (5). sev- eral times that figure. We are talking about large Also in progress are studies of specific medical numbers of people. issues that arise in transitional groups, as the Who among us can read the history of the latter poi tion of this session willattest. The relations between the early settlers of his country work of Roche and colleagues in particular will and the Indians without deep shame for the illustrate just how rapid and subtlemay be the barbarism heaped upon a people whowere changes introduced by minimal acculturation. If driven to defend the land they occupied? The genetic factors are important in the suscepti- world is watching my countryas it agonizes bility to the thyroid hyperplasia and diabetes o.er the Negro problemit might equally well mellitus now so common incertain Indian be watching the Americas for signs ofa belated groups, then, because of the recency, frequency, moral resurgence with respect to the Indians. and intensity of the phenomena. these groups How satisfied are any of us with the official may be unusually iavorable material for the programs of our governments for the health, study of these genetic factors. economic advancement, and education of the Some unpublished studies of Drs. Warren Indians? How can we translate the results of Eveland, William Oliver, and myselfmay helpour sciti,tific investigations into concrete act;on point to some of the nuances of this transition. programs, programs that must be carefully re- Six E. coli strains were isolated by Dr. Evelandlated to other government measures? Itis a from each of 77 Yanomama stools, each stooldubious favor to lower infant mortalityamong from a different subject. An effort is inprogress the Indians without parallel economic measures to type each of the resulting 462 strains, using to ensure food for the extra mouths. Nor does some 140 different typing sera. Thus far ap-it seem likely that Indians' accident-proneness proximately 44 per cent of the strains failto (see references in 23), so easy to attribute un- type out. By contrast, experience indicates thatcritically to their heritage of violence and lack only a very small fraction of E. coil strains iso-of familiarity with our gadgets, will yield to lated in the United States would failto type. education until the frustrations that contribute What remains to be determined is whether this to accident-proneness are relieved. is a unique finding ina long-isolated population In a world that seems to be groping for per- or whether similar studies en other such groups spective, the Indian provides a reference point will reveal the same pattern, with the inference from which to view the fantastic distupt'---, of a predictable shift in the intestinal flora with that modern men, intrinsically still little differ acculturation. ent in all essential biological attributes from

52 58 Indians, havc brought about. There arethose data are grossly inadequate and wewho call who wul take umbrage at mycharacterization ourselves scientists must adhere tothenull of us representatives of Westernculture as "in- hypothesis, the more so since its variousalterna- by those trinsically still. .Indians." I am aware of the tive ,. can be so conveniently misused so-called intelligence tests that purport toshow who would evade their socialresponsibilities. the inferior intellectual qcalities of theAmerican Be this last digression as it may,each time I Indian, ;ust as I am aware of similar resultswith return from the fieldthere is a period of culture the American Negro. These results canbe and shock as I realize how greatly in ashort period have been used as an excuse forless-then-equal of time we have contrived todisrupt our eco- schoolin. But in both instances it is a matterof system, and howprofound is our ignorance of a culturallydeprived and alienated group--per- the long-range results of this disruption.Now haps also subjected to early nutritionaldefi- in this time of greatly intensified concern over ciencies, whose role in impakedintellectual per- these problems, studies in depth of theIndian, recognize- formance we are just beginning to within or without the framework of IBP,will standard- being judged by tests designed by and surey contr:bute not only to his well-beingbut 4). By ized on a very different group (see also eventually, the these remarks I do not mean todismiss the also to our own pe;spective and, toward which we are possibilityofintellectualdiffereuces between necessary adjustments ethnic groups, but only to insist that to datethe evolving.

REFERENCES The ABO 1. ARENDS, T., G. BREWER, N.CHAGNON, M. 11. MOURA NT, A. E., and A. C. Konic. J. NEEL, Blood Group:: Comprehrnsive Tablesand Maps of GALLANGO, H. GERSHOWITZ, M. LAYRISSE, Charles C D. SHREFFLER, R. TASH IA N, and L.WE ITKAMP. World Distribution. Springfield, Illinois, (Royal Anthropological Intratribal genetic differentiation amongthe Yano- Thomas,1958.276 pp. mama Indians ofSouthern Venezuela. Proc. Nat. Institute Occasional Publication 13) 12. NEEL, J. V., and F. M. SALZANO. A prospectus Acad. Sci. 57:1252-1259, 1967. for genetic studies of the AmericanIndian. Cold 'k.G.,13.H. BOWMAN, and F. D. 2. BEARN, 1964. KITCI-HN. Genetic and biochemicalconsiderations of Spring Harbor Symp. Quant. Biol. 27:85-98, Cold Spring 13. NEEL, J. V., F. M. SALZANO, P. C.JUNQUE IRA, the serum group-specific component, &tidies on the Harbor Symp. Quant. Biol. 29:435-442, 1964. F. KEITER, and D. MA Y BUR Y-LEWIS. Mato Grosso. 3. CAVALLI-SFORZA, L. L., and A. W. F.EDWARDS. Xavante Indians of the Brazilian Analysis of human evolution. In S. J. Geerts(ed.), Amer. I. Hum. Genet. 16:52-140, 1964. Genetics Today, vol.3. Oxford, Pergamon Press, 14. NEEL, J.V., and F. M. SALZA NO. Further studies on the Xavante Indians. X. Somehypoth- 1965, pp. 923-932. studies 4. FiscHER, J. L. Letter to the editor.Perspectives eses-generalizationsresulting from these Amer. I. Hum. Genet. 19:554-575,1967. in Biol. & Med. 11:325-328, 1968. demog- 5. HACKEN BERG, R. A. An anthropological studyof 15. NEEL, J. V., and N. A. CHAGNON. The demographic transition. Milbank Memorial Fund raphy of two tribes of primitive, relatively un- Acad. Quart. 44:470-493, 1966. acculturated American Indians. Proc. Nat. 6. HANNA, B. L. The biologicalrelationships Sci. A:680-689, 1968. among Indian groups ofthe Southwest. Amer. J. 16. NEEL, J.V., and W. J. Salm.. On some Per- Phys. Anth..op. 20:499-508, 1962. trends in vnderstanding the genetics of man. L. Continuously distributedfactors spectives in Biol. & Med. 11:565-602, 1968. 7. KING, J. indlgena de affecting fitness. Genetics 55:183-492, 1967, 17. ROSE NBLAT,A. La poblai:ion 8. LEWONTIN, R. C. Population genetics.Ann. America. Buenos Aires, Institución Espaiola,1945. of South Rev. Genetics 1:37-70, 1967. 18. SALZANO, F. M. The blood groups 15:555- 9. MILKMAN, R. D. Heterosis as a major causeof American Indians. Amer. J. Phys. Anthrop. heterozygosity in nature. Genet:es 55:493-495, 1967. 579, 1957. 10. MOURANT, A. E. The Distribution of the Hu- 19. SALZA NO, F. M. Studies on the Caingang man Blood Groups.Springfield, Illinois, Charles C Irodians. I. Demography. Human Biol. 23:110-130, Thomas, 1954, 438 pp. 1961.

53 59 20. SALZANO, F. M. Some geneticaspects of the Accidental deaths among British demography of American Indians. Columbia Indians. In J. Sutter (ed.), Canad. Med. Assoc.J. 94:228, 1966. Human Displacements, Entretiensde Monaco en 24. STEWARD, J. Sciences Humaines, Premiere H., and L. C. FARON. Native Session. Monaco, Im- Peoples of South America.New York, McGraw- primerie Nationale de Monaco,1962. Hill, 1959. 21. SPUHLER,J.N., and C. KLUCKHOHN. In- 25. SVED, J. A., T. E. REED, and W.F. BODMER. breeding coefficients of the Ramah Navahopopula- The number of balanced polymorphisms tion. Human Biol. 25:295-317, 1953. that can be maintained ina natural population. Genetics 55: 22. SANGHVI, L. D. Comparison ofgenetical and 469-481, 1967. morphological methodfor a study of biological 26. World HealthOrganization.Researchon difference. Amer.I.Phys. Anihrop.11:385-404, Population Genetics of Primitive 1953. Groups. Gen ev a, World Health Organization, 1964.26 pp. (Techni- 23. SCHMITT, N., L. W. HOLE,and W. S. BARCLAY. cal Report Series, 279)

54 DISCUSSION

Dubos: Before we open the general discussion. determine the hign mortality to a much greater Dr. Neel, I should like to ask you to take over extent thnn any innate susceptibility. again the problem raised by Dr. Water lo v. a Now, how to pin down that kinJ of impres- while ago. He asked about the evidence for the sion? It is very difficult. One practical approach genetic character of some of the differences in is simply to provide the Indian, when he first blood groups and you answered. But I should encounter:: tuberculosis or meLsIes or pertussis, like to ask if you could outline for us what kind with the same kind of medical care and nutrition of genetic evidence you can hope to obtain with that corresponding Caucasian pepulations are regard to those adaptive processes of ,Jhich you given and see what the difference is. have spoken, especially with regard to the prob- As you know, Dr. Dubos, there is a consider- lems of suscep:ibility to disease, which will be able literature, basal largely on stAies in twins, discussed this afternoon. In other words, how on disease susceptibilities in man. The.-e is also possIle ;sit in practice to obtain the kind ofa large literature of studiesin experimental information that would determine whether theanimals, the results of which are sometimes change is genetic or simply phenotypic? transferred to man. This literature leaves much Moderator: I am afraid that an answer to that to be desired; there is a need for precise investi- question might anticipate a good deal of ourgations, as with standardized challenges under discussion forthis afternoon.ftisrelatively controlled conditions. simple to obtain evidence for the genetic nature Waterlow: What is worrying me, Dr. Ned of the kinds of traits that Dr. Layrisse, Dr.and I am very ignorant of this subjectis that Arends, and Dr. Lisker have presented. Itis the markers you are studying may in a sense be clearly mucA more difficult to obtain evichnce of no physiological importance, just as the color for the genetic basis of such attributes as disease of one's iris might be said to be of no importance resistance. physiologically. We shall hear from Dr. Nutels this afternoon I am assuming that the variety of polymorphic aly ut some of the medical problems ai Indian proteins, the isoe7.zymes such as the phospha- groups in transition. We have recently had the tases,dehydrogenases, andsoon,arealso experience of being in the middle of a measles genetically detcrmined. I do not know whether epidemic in a virgin population of Indians. It this is proved yet. Such proteins are likely on was a very instructive and sobering experience. physiological grounds to be more important in But one of :he strong impressions we came relation to adaptability. What I am not clear away withand Dr. Centerwali will presentabout is how far your program of the kind of some of the datais that there may have beenthings you study is complete in itself. It rather a great deal of uncritical thinking in the past looks asif geneticists had their own list of about the constitutionalsusceptibility of theproteins that they callmarkers, whereas we Indian to this, that, and the other disease. We biochemistsarethinking of rather different feel that the secondary epidemiological charac- types of proteins. teristics of populations living at this level may Moderator: Dr. Waterlow, you have asked a

55 very perceptive question, on which, again, we simply physiological adjustment to lifeat high could spend a great deal of time. Probably the altitudes. most important development in human genetics Now, one of the better opportunitiesto get in the past decade has been the realization of at a question like that exists in Ethiopia. In the the tremendous amount of genetically deter-highlands and lowlands of Ethiopiaare groups mined biochemical diversity in our species.1, who on the basis of the geneticr ,arkers we for one, think that one of the outstanding chal- have been discussing appear to bevery similar. lenges in human genetics is to understand theAnd yet one group lives ata very considerable significance of this diversity. What is it all there altitude and one group not far abovesea level. for? It is so great that we are having difficultyThere is thus an experiment ofnature, as it in seeing how it can all be maintained at oncewere, in which a group has been subdivided and by selective pressures. The problem is to sort outthe resultiug subgroups placed in &verseen- which oi these polymorphisinsare like vestigial vironments. organs and which are meaningful to the species. The situation does notseem to be quite so Now, to come more directly to how we getsimple in the Americas. We have not yet found at this. It is quite clear that those polymorph-the "control" for Andean man. Once thereare isms arose in primitive man. And we feel that such controls it is possible to investigate whether many dues to their function may come from they are physically more similar to high-altitude primitive man. One of the things we attempt to try.ri than other Indian groupsthat is, whether do in our field work is collect allour data in there has been self-selection of groups able to family relationships. We can then analyze familylive at high altitudes. There is also the hope of material to see whether cercain polymorphisms finding marital exchanges between the high- are associated with increased fertility or with and low-altitude groups. decreased infant mortality. This isa long, slow Dubos: Would you elaborateon the state- business. T'arallel studies on civilized populations ments that were made last year about the ana- are also needei. The numbers necessary to estab- tomica' differences in the heart betweenman lish a significant associaton are tremendous.born at a high altitude andman born at our And, in fact, we almost have to think ina altitude? higher order of magnitude than our past studies Moderator:I would prefer that those who when we try to come to grips with these issues. made the statements were here to elaborateon To sum up, althongh we are interested in them. As I recall it, what we were shownwas these traits as "markers," for which purposewe evidence that changes appear very earlyon in need not understand their physiological signifi- the heart and lungs, But I didnot see any cance, we are also just as interested in under- convincing eidence that these early changes standing what they do. were genetic in nature. As far as I am concerned, Dubos: Do you see any possibility of deter- itisa wide-open question whether thisis mining whether adaptation to high altitude isa genetic or an environmental adaptation. genetically determined ora purely phenotypic Griffin: I have a suggestion on the general adaptation? I think this is relevant toour dis- problem Dr. Neel is working on. Therewas a cussion last year. somewhat comparable dispersal, bya quite dif- Moderator: There is a great deal of interestferent group, into Australia at about the same now in adaptation to high altitude, not only in time, according to radiocarbon dates. Studies of the Andes but also in the Himalayas and other the Australian aborigines, which werea much parts of the world. And there is, as you know, less diversified and much smaller population, a running debate on how much of the adapta-might be important with regard to evolution tion we seefor example, the barrel chest of the among "isolated" human groups. Andean Indianis genetic and how much is Moderator: Yes, I agree that the Auitralian

56 aborigine does represent the group most neatly and it is a very different worid, the world of comparable to the Indian. However, the disrup- primitive manyou nevertheless quickly recog- tion and decimation of the Australian aborigine nize all the familiar types of our own culture. is even further along than that of the American You do not have to be very long in an Indian Indian.Itis extremely difficult now to find village before you get the same feeling for the groups of any size that have notbeen seriously strong men and the weak men and the operators disturbed by being displaced into very unfavor-and so on that you encounter in our own cul- able country or in which there has not been a ture. forcedamalomation ofpreviouslyseparate So I would say that at least part of the tran- groups. scription is probably much the same at all cul- Cohen: Following up Dr. Water low's and Dr. tural levels. Whetho-, on the other hand, there Neel's remarks, I would like to ask Dr. Neelare sabtle differences in the genetic readout, whether geneticists\ are being prepared for the who can say? fact that, at least in molecular biological or bio- McDermott: ig the lines of Dr. Dubos' chemical terms, the concept of the regulation of question, Dr. Neel, arcthe people who are readouts at the translational level as well as at conducting the studies of Ifswer-animal behavior the transcriptional level is beginning to emerge. dso making ths.se batteries of observations on What brings this to mind is your allusion to the the various genetic markers? I was thinking, primitive Indians' often being relieved of the for example, that the question of adaptability to burden of contamination carried by our popula- high and low altitude might conceivably be tion. They are living with quite different intes- approached from that standnoint. Is there work tinal flora, parasites, and perhaps even types of in this field, attempting to relate these genetic natural foodstuffs to which our population istraits to the behavioral pattern of the animal? not exposed, but I do not know whether in fact Moderator: We know more about man's their environment can necessarily be spoken of genetic constitution than about that of any other ac less contaminated. The primitive population mammal at the present time. However, what would be less contam:nated by synthetic andmight be called the comparative polymorph- non-natural factors; relative to our own situa- ology of the primates is under very ac ive investi- tion, however, it may be more contaminated. gation. A great deal of effort is now going into This comes toaffect the whole question ofexploring the extent to which the higher pri- separation of genetic potential because of the mates have polymorphisms comparable to those possibility that transcription may be regulated of man. It is too early !or any generalizations, at the level of translation. There may be allbut itis a fact that some polymorphisms en- kinds of factors in terms of the phenotypiccountered in man are found in the higher pri- result that could emerge by very subtle devices. mates. On the other hand, the primates show I am wondering whether even the familial tests polymorphisms that are not found in man. No that the geneticists traditionally use would get pattern has yet emerged. I assume that in due around this problem. time it will be possible to try again to make the Moderator:Iwould remind you that the kinds of associations you refer to. But, again, I geneticist has had quite a hand in molecularcome hack to the problem of numbers. Itis biology. going to be very difficult to get large numbers Obviously, there is at present no answer toon any of the higher primates. It is much easier your question about differences in transcription, with man. qualitative or quantitative, at that cultural level Dubos: There can be no doubt that Dr. Neel as compared with our own. I think all of ushas not only touched on but developed for us who have done field work have emerged with the foremost problems of man's adapting to a feeling that here in a very different worldthe new conditions he is creating. All sorts of 5763 statements are continuously being made, such we obviously are not going to solve it here. But "Modern man has become adapted to crowding" I hope at least that Dr. Neel will conclude with or "... cannotbecome adapted to crowding"; a general statement on this issue. "Modern man has become adapted to pollution" Moderator: I merely want to agree that we .. cannot or ". adapt to pollution." And, in are not going to solve the problem this morning fact, these problems, which are so crucial to the and w agree with one other thing Dr. Dubos whole development or survival of our civiliza- tion, have not as ytt been formulated in suffi-has impliedthat there are grkat difficulties ;r1 ciently clear terms for us to determine whetherclear formulations in this field. How do we set we are dealing only with genetic adaptability, up specific questions to which we can get what only with phenotypic characteristics, or whether would be called scientifically hard answers? I we are simply playing on the polymorphic rich- think we are in accord ;hat this is an important ness of the human race. area. And I think we are in accord -sith respect The problem is so large and so important that to the methodology: we have a long way to gL.

58 SURVEY OF THE UNACCULTURATEDINDIANS OF CENTRAL AND SOUTH AMERICA'

Francisco M. Salzano

Acculturation and biological change context, is that related to thebiological changes that occur and sometimes are decisive for the Before trying to make the survey mentioned direction and speed of acculturative modifica- in the tide itis important to indicate the prob- lems that arise when an attempt is made to tions. Ribeiro (20) recognizes for Brazilian Indians specific group as"acculturated"or definea three stages on the road to acculturation. The "unacculturated." The term acculturation is first would be represented by the isolated tribes generally employed by American anthropolo- those living in regions not yet occupied by the gists, while their British colleagues prefer the word culture contact and the Cuban scholar Brazilian society and having only accidental and civilization. Some simply F. Ortiz the term transculturation, which em- rare contacts with our avoid such contacts, while those living near phasizes the reciprocal character of most contact pioneer settlements may have suffered violence situations. Though much criticized and modi- In fied, the most widely med definition is that ofand be frankly hostile to "foreign" people. either case, they can obtain all they need from Rcdfield, Lindon, and Herskovits (19): "Accul- the region wherc they live and from their work; turation comprehends those phenomena which they are therefore able to maintain complete result when groups of individua's having dif- cultural autonomy. The second stage wouldbe ferent cultures come into cot; tinhous first-hand that of populations having intermittent contacts contact, with subsequent changes inthe original with civilization. They live in regionsalready cultural patterns of either or both groups." reached by the expanding frontiers ofthe na- A number of basic problems exist in the inter- tional society; though they maintain a certain pretation and analysis of these situations; they cultural autonomy, they already have needsthat are reviewedby, among others, Beals (1) and can only be satisfiedthrough economic relations Oliveira (16). For instance, we have sometimes with civilized people. The third stage is repre- conflicting views of culturaliqs and sociologists, sented by groups in permanent contactwith and in some specific cases the intrinsic nature of more numerous and moredifferentiated mem- the process and its consequences are difficult to losttheir identify. A new dimension, which so far as Ibers of the civilization. They have cultural autonomy to a large degree, being en- has not been properly analy2ed in this am aware tirely dependent on others for the supplyof products. They stillpreserve I Research supported in part by the Rockefeller Foun- many tools and dation, Consdho NaCior,;11de Pestwisas, Conselho de some of theirtraditional customs, but already Pesquisas da Universidade Federal do Rio Grandedo show profound changes due to ecologic, eco- Sul,U.S.PublicHealthService.andWenner-Gren Foundation for Anthropological Research. nomic, and cultural pressures.

59 65 -.4 ) What concomitant modifications occur in thesonal communications of many colleagues. The biology of these groups? Only very limited rapidity of change in these poputations and also start has been made on quantifying the observa- the discovery of new groups indicate that such dons made in groups at these three stages, and a list should be considered as merely an approxi- much more is needed to obtain a complete pic- mation of the situation in the several areas stir- ture (12, 13). One important point that should veyed.Moreover, the publications examined be emphasized is the speed of these changes.sometimes do not give basic details about the Probably the first characteristics to be affected degree of acculturation of the group in question. are the epidemiological ones, and here manyEven with these reservations, the list may per- groups that show all the other traits typical of haps serve to give a rough idea of the opportuni- untouched populations may present patterns re- ties that still exist for the study of populations sembling those of groups at other stages of at this cultural level. In all cases the figures con- acculturation. A typical example is provided by cerning the country's total and its Indian popu- the Txukahamae, a Cayap6 group of central lations were checked in the must recent official Brazil (14). They have a completely autono- publications. The references indicated are those mous life and in every instance are representa- examined in addition to these sources. Only tive of an isolated population, but they already relatively recent estimates were included;for have a high inci'lence of tuberculin reactors. reviews concerning the older ones, see Rosenblat Since the Amerindian populations apparently (21) and Steward (23). had no contact with tuberclebacilliinpre- Columbian times,the questionarises. FromThe survey what groups did they get the infection? Their CENTRAL AMERICA contacts with neo-Brazilians are recent and in- termittent, and itis reasonable to suppose that Politicalsubdivisions:Bahamas,Belize,Bermuda. CostaRica. Cuba. Dominican Republic,ElSalvador. the source of infection was the Gorotire, another Guadeloupe (and depLndencics), Gu:uemala, Haiti, Hon- Cayapb group from whom they separated at the duras. Martinique, Mexico, Netherlands Antilles, Nica- beginning ot this century but with whom they ragua, Panama, Puerto hico, Virgin Islands, Wcst Indies. still have frequent interchange. The diffusion of Total population: 77 million. this Pathological agent in some ways parallels Countries with significant number of Indians: Belize. Costa Rica, El Salvador. Guatemala, Honduras, Mexico. the cAusion of culturaltraits. The speed of Nicaragua, Panama. these epidemiological changes is such that itis Indianpopulation:8,000,000(10percentofthe improbable that the study of present groups, no area's total population). matter how untouched, would give us exact Unacculturated Indians: Perhaps 1,000,000(1.3per duplicates of past populations. But investigation cent of total population).

of the biological and nonbiological factors that BELIZE influence the genetic and physiological adapta- Total population: 100,000. tion of these groups to new diseases and infec- Indian population: 8,000-10,000. tious agents has intrinsic interest and can eluci- InformationabouttheIndians: Mainly Maya and date important problems of human biology. Chekchi, who came from Mexico and Guatemala. Unacculturated Indians: Probably none. Reference: 9. Source of the data and their limitation In the next section Ishall try to present a COSTA RICA survey of the relatively unacculturated Indians Total population: 1,500,000. of Central and South America. Since I have Indian population: 7,200. Information about the Indians: Most important grcups first-hand contact only with Brazilian Indians, appear to be Bribri (about 3,500), living in Talamanca information concerning other countries has had Range in south, and Boruca (about 1,000) in southwest to be limited to printed material and the per- on the Pacific side. Remaining Indians belong to groups

60 66 of Chorotegamangue (Nicova Peninsula) and Guatuso NICARAGUA (northern and western bordeis). Total population; 1 ,631.,000 ( 966). Unacculturated Indians: Probably none. Indian population: 80,000. Refermices: 9, 24. Information about the Indians. Most important groups El. SALVADOR arc Miskito (with a krge component of Negro mixture), Sumu, Rama, and Ulva, Total population: 3,000,000. Unacculturated Indiars: Stor,:2 (24) mentions the Ulva Indian population: 100,000. tribe bettk e.,,r) the Mico and Siquia rivers, and the Sumu Information about the Indians: Most important group (3,000-4,00) in 1948). is Pipil (about 80,000) of Nahuatl origin, concentrated References: 9, IS. 22, 24, in various towns in Sonsonatc Department. Also smaller groups of Nahua and Cakchiquel distributed over several PANAMA depart men ts. Unacculturated Indians: Probably none. Total population: 1,31-.A60. References: 9, 24. Indian population: 65,000-100. 000. Liformation about the Indians: Main tribes are Guaytni GUATEMALA (about 36,000), Cuna (some 22,000), and Choc6 (6,000). Two other small groups are Bocota (2(10) and Teribe Total population: 4,438,000 (1965). Indian population: 2,500,000. (400), Guaymi are found in Chiriqui, Bocas del Toro, Information about the Indians: Main groups in center and Veraguas provinces; Cuna mainly in San Bias Pri.v- and west: Maya, Cakchiquel, Tzutujil, Uspanteca, Achi. ince, both mainland and some islands: Chou; in Dari&I West: Mam, Aguacateca, jacaltec, Kanjobal, Chuj, Ixil. Province. Unacadturated Indians: Some Choc6 groups of Pan- North: Kcehki, Pocomchi, Pocomam, Lacand6n, Mopan. amanian-Colombian border (:). East: Chortf, Lacand6n Chol, Chontal. Southeast: Xinca. Reference's: 9. 24, 26, Unacculturated Indians: Some groups livinginin- accessible places could provide information on thc influ- SOUTH AMERICA ence of a non-European culture on genetic and other traits. Political subdivision: Argentina, Bolivia, Brazil, Chile, References: 9, 18. Cokmbia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Surinam, Uruguay, Venezuela. HONDURAS Total population: 175 nUllion. Total popula'i,?n: 2,400,000. Countries withsignificant number of Indians:All Indian population: 110,000. except Uruguay. Information ab,1 the 'ndians: Main groups in west: Indian population: 10 million (6 per cent of the area's Chorotega, Miquirano, Guajiro. On coast: Moreno. Other: total population) Opatoro,Jicaque,Paya, Zambo, Miskito, Sumu. Thc Unaccultw.ated Indians: Perhaps 980,000 (0.6 per cent Miskito nave mixed with Negro groups. of the total population). Unacculturated Indians: Probably none. ARGENTINA References: 9, 18, 24. Total population: 22,254,102 (1965). MEXICO Indian population: 130,000. Total population: 40,913,000 (1965). Information about the Indians: Main groups in Chaco Indian population: Estimates vary widely. A conserva- region:Mataeo, Chulupi, Chorote, Chiriguano, Pilag tive number would be 5,000,000. (totaling about 30,000). Andean region: Quechua and Information about the Indians: Main groups in north: Colla (some 18,000). South: Araucanian. Other regions: Kikapti, Tcpchuano, Nahuatl, Panic, Huastec, Otomi. Caingui, Mocobi, Toba. Northern Pacific: Cora. Huichol, Yaqui, Seri, Papago. Unacculturated Indians: Perhaps some Chaco groups. Center: Mazahua, Matlatzinca, Tarasco, Mazatec, Mixtec, References: 8, 9. Popoloco,Totonac. Zoque, some of previous groups. Southern Pacific: Chol, Tzeltal Tzotzil, Tojolabal, Maya, BOLIVIA Lacand6n, Amuzgo, Tlapancc, Chatirio, Chinantec, Cho- Total population: 3,800,000. cho, Chontal, Huave, Mixe, Triche, Zapotec. Indian population: 2,450,000. Unacculturated Indians: LacandOn (only about 200). Information about the Indians: Main groups in Andean Some groups living in inaccessible places could provide region: Quechua (about 1.500,000) and Aymara (850,- information on influence of a non-European culture on 000). Northeastern region: The forest-dwelling popula- genetic and other traits. tions (approximately 87,000) are divided into about 63 References: 4. 9. 18, 25. tribes belonging to 15 linguistic families. Sonic of them arc as follows: Beni Depanment: Sirion6, Baurc, lora, COLOMBIA Chacobo, Tacana, Chimane, Movima, Yan,Sirineire, Total population: 11,787,000. Cayubaba, Itoma; Departmentf Santa Cruz: Guarayo, Indian popdatio-,: 300,000. Borosese, Chamacoco, Zainuco, Lengua, Tapieti, Otuqui, Information about the Indians: Main groups in Cauca Guatose, Yuracare; Pando Department: Ara( na, Toro- region: Guambiano, Pgez, Guanaco, Yanacona,Coc.)- norna, Pacaguara, Machicanga; Department of La Paz: nuco(approximately80,000).Narifio:Quillacinga. Chama. Chimane, Guarayo, Leco. Soutilcastern region Kwaiker, Inga, Kamsi (about 60,000). Gn..;i-a: Guajiro (BolivianChaco):TarijaDepartment;Chiriguano, (50,000). Magdalena: Kogi, Ika, Chimila, Yu,o (15,000). Mataco,Toba,Chulupi,Guaycuru,Chanese(about Antioquia: Cuna (2,000). 14,500); Quiluisaca Department: Chorote, Chiriguano Unacculturated Indians: Perhaps 100,000. (some 3,800); Cochabamba Department: Sirion6. Yura- References: 2, 9. care (2,000). General total for the region: approximately 50,000 Indians. IR Unaccultw-ated Indians: Almost allforest-dwellers. References: 9, 28. Tovi? population: 5,084,000. India:i population: 2,000,000. Information about the Indians: Main groups in coastal BRAZIL region and cast:Colorado,Jivaro(13,000), Yumbo, Auca. Sic:ra: Quechua groups: Quindivana, Saguatoa, Total population: 90,193,000. Salasate, Poat), Ilumane, Peguche, Quinchuqui, Quilotoa, Indian population: 100,600. Zunibagua, Pilaloe,Siquisilie, Puruhay, Caiiaris,Sara- Information about the Indians Tribes or groups classi- guro. fied by Ribeiro (20) and others as isolated or in inter- Unacculturated Indians: Perhaps 25 ),006 (50,000 in mittent contact(excluding those with fewer than 50 the juggle). individuals): Agavotokueng, Apalai (100-250). Araras, References: 9. 15, 17, 2". Asurini(250-500), Atruahi, Baraw5na, Boca Negra,

Canoeiros, Cayabi (250-500), Diore (500-1,000), FRRNCII GUIANA Gaviiks (1,500-2,000), Guaji (100-250), Ipewi, Javai (250-500).Kabixi,Kalapalo(100-250),Kamayuri Total population: 36,000. (100-250), Katawiin, Kren-Akoro (2,500-3,000), Indian popdation: 1,200. Kuben-Kran-Kegn (250-500) Kuben-Kragnotire (400), Information about the Indians:Incoastalregion: Kuikuro(100-250),Makii, Mandawgka, Maopityan, Galibi (600), Arawak (100), Palikour (100). In interior: Makir; tareorMayongong (500-1,000),Mehiniku Wayapi (130), Mereyo (60), Waiyana (200). (50-100),Mentuktire(560-1,000),1udjetirc(50- Unacculturated Indians: About 900 (Galibi, Palikour, Waiyana). 100),Nahuktui(50-100), Nambicuara(500-1,000), Pakainovas (500-1,000), Parakanin (250-500),Pari- Pejo ences: 3. kot6, Pianokot6, (250-500),Purubori (50-100), Sikiâna, Tapaytina, Tine.) (2,000-3,000), GUYANA Txicio, Txukahamae(200), Urubus-Kaapor Total pop;ilation: 647,000. k500-1,000), Waimiri, Waiwai (100-250), Wauri (100- Indian population: 27,840. 250), Xavante (2,000-3,000), Xikrin (250-500), Ya- Information abdut the Indians: Main groups in coastal baina, Yanomama (2,000). For location, sec references region: Arawak Carib, Warrau, Akawaio (about 10,000). 11. ind 19. The former presents many othcr names, but In interior: Carib, Akawaio, Patamona, Arckuna, Makusi. it is not always clear whether tribes or tribal subdivis.ons Wapisiana, Waiwai. are meant. Unaccuiturated Indians: Perhaps 2,000. Unacculturated Indians: Pertaps about 40,000. References: 3,Iniorroillon noten Arte4d9. References: 11, 20. T4(1111.efft et)V.a'Spew.

PARAGUAY

GIME Total population: 2,030,000. Indian population: 37,570. Total population: 8,567,000 (1965). Inform.:tion about the Indians: They speak languages Indian populatior. 130,000. of 6 linguistic families and live in i7 locations. Main Information aboia the Indians: Main groups in North- groups:Chiripi,Mbia,Tabiterii,Guayaki,Guarayti, ernregion: Quechua and Aymara. Southernregion: Chiriguano. Tapicti, Maki (about 600), Chulupi (more Araucanian (Mapuchcs). than10,000),Chorote(approximately100), Lengua Unacculturated Probably none. (5,000), Sanapana (3,400), Guana (400), Angaitc (400), References: 9. Toba (1,400), Chamacoco, Morotoco. Unacvulturated Indians: Perhaps 10,000. has to do with the identification of theaccultura- References: 9; data supplied by Department of Indian tion process per se; the other is the properplace- Affairs. ment of the groupstudied into "acculturated" or "unacculturated" categories.Because of the PERU lack of detailed studies in most of the tribes Total population: 11,800.000. surveyed here, and of reviews placing them in Indian population: 4,800,000. Informatior about the Indians: Main groups in North- the appropriate category, the numbers obtained Aguz,runa,Jivare,Huambisa, Andoa,Shapra, have a very relative value. They should be Curaraye, Chayhuita, Jebero. Northeast: Bora,Ocayna, viewed as indicating orders of magnitude only. Orejon,Pano,Yagua,Mayoruna,Charna,Huitoto, With these reservations in mind it is now pos- Ticuna, Cocoma, Capanahua, Cashibo, Shipibo. Center: sible to examine the numbers arrived at.The Campa, Amuesha, Amahuaca, Marinahua,Cashinaua. Southeast: Sirinayri, Machigiienga, Piro, Mashco,Hua- population of Centi al America was estimated at rayo. Carneiro (5) also mentionsfollowing tribes from 77 million and the number of unacculturated montaiia: Yaminahua, Remo, Marobo and Iiiapari. Indians at about one million (1.3 per cent). In Unacculturated Indians: Perhaps 450,000. South America the proportionis even lower: References: 5. 9, 27, 28. for a total of 175 million I calculate approxi-

SURINAM mately 980,000 (0.6 per cent). But the fact that have about two million largely Total population: 360,000. we may still Indian population: 4,500. "untouched" Indians points to the magnitude Information about the Indians: Main groups in coastal of the task, if we want to obtain truly compara- region: Galibi(2,400), Arawai:(1,500). Ininterior: tive data in a number of populations living in Waiyana (2(A)), Trio or Tiri6 (00). different ecosystems. As a matter of fact, we Unacculturated Indians: Perhaps 2,000. simply do not have experienced persons to un- Reference: 3. dertake the study of even, say, 10 per cent of this VENEZU ELA number in the immediate future. 7 otal population: 8,880.000. Withacculturationchangesoccurringso Ihdian population: 62.700. quickly, we iiced to work fast to establish on a Information about the Indians: Layrisse and wilhert firm basis the biological and nonbiological forces (9)listthefollowing:Guajiro(8,429).Paraujano (1,348). \Alp (2,057), Bari (1,000), Yaruro (1,427), acting at this cultural level and determining .he Guahillo (5.397), Piaroa (1,886). Panarc (412), Yaba- fate of these populations.This informationis rana (64), Makiritarc(1,200), Piapoco (99), Curipaco basic to our understanding of human evolution, (212), Puinave (240), Bare (645), Yanomama (20,000), but it is also important in relation to practical Sap6 (100), Pemon (2.700), Warao (11,700),Carilia problems. The acculturation process is generally (2,776), unknown (1,000). painful and distressing. In fact, a large propor- Unacculturaoed Indians: Perhaps 30,000. References: 6, 7, 10, 29, 30. tion of the groups affected by it si:nply disappear physically. Data on populations living at dif- Discussion ferent cultural levels can provide clues to an As was mentioned preyiously the estimates easier and more humane way toward techno- obtained present at least two types of error. One logical advance.

REFERENCES 1. BEALS, R. Acculturation. In A.L. Kroeher 3. BuTT, A. J. The Guianas. Bull. Intern. Comm. (ed.), Anthropology Today. Chicago, Universityof Urgent Anthrop. Ethnol. Res. 7:69-90, 1965. Chicago Press, 1953, pp. 621-641. 4. CAMARA, P.Rescate etnogra'fico nacional en 2. BURGOS GUEVARA, H., and G. PESANTEZREINos., Maico.Bull,Intern. Comm. Urgent Anthrop. Plan nacional indigenista de Colombia:1966-1969. Ethnol. Res. 7:99-104, 1965. America lndigena 27:751-781, 1967. 5. CARNEIRO, R. L. Little-known tribes of the

63 69.:1 Peruvian Montana, Bull.intern. Comm. Urgent aniversario,I. Wxico, D. F., Libros oe Mexico, Anthrop. Ethnol. Res. 5:80-85, 1962. 1965, pp. 79-134. 6. ComsION INDIGENISTA NACIONAL (VENEZUELA). 18. PEP ICOT Y GARCIA, L. America In digena. Tomo Nuestro Indio. Caracas, Ministerio de Justicia, 1960. I. El hombre americaro. Los pueblos de Amer;ca. 26 pp. 2nd ed. BarTelona, Salvat. 1962. 1182 pp. 7. COMISION DIGE NISTA NACIONAL. Informe de 19. REDFIELD, R., R LINTON, and M. J. HERSKO- la ComisitSn Indigenista Nacional 'del Ministerio de vas. Memorandum on the study of acculturation. Justicia de Venezuela, correspondiente al-nio de Amer. Anthrop. 38:149-152, 1936. 1964. America Indigena 25:131-145, 1965. 20. RIBEIRO, D. Culturas e linguas indigenas do 8. Focx, N. Utgent ethnographical tasks in the Brasil. Educa4o e Ciencias Sociais (Rio de Janeiro) ArgentireChaco.Bull.Intern.Comm. Urgent 2:5-102, 1957. Anthrop. Ethnol. Res. 5:162-166, 1962. 21. ROSENBLAT,A. La poblaciónindigent:de 9. Guia de lapoblación indigena de America. America. BuenosAires,InstituciónCultural Es- Indigenista 21 :170-266, 1961. panola, 1945, 296 pp. 10. LAYRISSF., M., and J. WILBERT. Indian societies 22, STEWARD,J.H.(ed.) Handbook of South of Venezuela. Their blood group types. Caracas,American Inthans. Bull Bur, Amer. Ethnol. 143 (5 InstitutoCaribedeAntropologiaySociologia, volumes, 1946-1950). Fundación La Salle de Ciencias Naturales, 1966. 23. FrEWARD, J.H. The native population of 318 pp. (Monograph 13) SouthAmerica. Handbook of South American 1 I MA LCHER, J. M. G. indios. Crau de integracilo Indians 5:655-668, 1949. na comunidade nacional. Grupc 'lnguistico. Locali- 24. STONE, D. Urgent tasks of research concerning za0o.RiodeJaneiro,Conselho Nacionalde the cultnres and languages of Central American Proteciio aosIndios,1964. 266 pp. (Nova sArie, Indian tribes. Bull. intern. Comm. Urgent Anthrop. Publicaciio 1) Ethnol. Res. 5:65-69, 1962. 12. NEEL, J. V., and F. M. SALZANO. A prospectus 25. SWANTON, J. R. The :ndian tribes of North for genetic studies on the Americau Indian. In P. T. America. Bull. Bur. Amer. Ethnol. 145:1-726, 1952. J.S. Weiner (eds.), The Biology of Baker and 26. TORRES DE ARAUZ, R. Los grupos humanos de Human Adaptability. Oxford, Clarendon Press, 1966, Pananmi. In Homtnal? a luan Comas en su 65 pp. 245-274. aniversario,I.Mexico, D. F., Libros de Mexico, V., and F. M. SAL1ANO. Further 13. NEEL, J. 1965, pp. 135-142. studies on the Xavante Indians. X. Some hypotheses- 27. VILLA Ropts, A. Departamento de Investiga- generalizations resulting from these studies. Amer. clonesAntropológicasdelInstitut('indigenista I. Hum. Genet. 19:554-574, 1967. Interamericano. Programa de actividades, julio de 14. NUTELS, N., M. AYRES, and F. M. SALZANO. Tuberculin reactions, X-ray and bacteriological stud- 1965 a junio de 1966. Anuayio lndigenista 25:87- ies in the Cayapó Indians of Brazil. Tubercle 48: 98, 1965. 195-200, 1967. 28. VILLA RojAs, A. Informe de labores del De- 15. OBE REM, U. 1962. Dringende ethnologische partamento de Investigaciones Antropológicas, 1965- Forsehungsaufgaben in Ost Ekuador. Bull. intern. 1966. Anuario Indigenista 26:7-95, 1966. Comm. Urgent Anthrop. Ethnol. Res. 5:75-79, 1962. 29. WILBERT, J.Dringliche Forschungsaufgaben 16. OLIVEIRA, R. C. DE. 0 Indio e o mundo dos in Venezuela. Bull. Intern. Comm. Urgent Anthrop. brancos. Sk Paulo, Difusk Europela do Livro, Ethnol. Res. 5:86-96, 1962. 1964. 144 pp. 30. ZERRIES, 0. Die völkerkundliche Fol.schungs- 17. ORBE, G. R. Destribalización de indigenas ci situation in Südost-Venezuela. Bull. Intern. Comm. Ecuador. In Homenaje a Juan Comas en su 65 Urgent Anthrop. Ethnol. Res. 5:97-111, 1962. Appendix ALPHABETICAL LIST OF TRIBES CITED 1

Achi (Gua) Chama (13o, Pc) fluambisa (Pe) Ag tvotokueng (Br) Chamacoco (Bo, Par) Huarayo (Pc) Aguacate (Gun) Chanese (13o) Huastec (M) Aguaruna (Pc) Chatitio (M) Huave (M) Akawaio (Gui) Chayhuita (Pe) Huichol (M) Amahuaca (Pc) Chekchi (Be) Huitoto (Pe) Amuesha (Pe) Chimane (Bo) Amuzgo (M) Chimila (Co) Ika (Co) Andoa (Pc) Chinantec (M) Ilumane (E) Angaize (Par) Chiriguano (A, Bo, Par) 1fiapari (Pc) Apalai (Br) Chiripa (Par) lnga (Co) Araana (Bo) Chocho (M) Ipewi (13r) Araras (Br) Choc6 (Pan) Itoma (13o) Araucanian (A) Choi (M) lxil (Gun) Araucanian (Mapuche) (Ch) Chontal (Gua, M) Arawak (Fr G, Guy, S) Chorotc (A, Bo, Par) Jacaltec (Gua) Arckuna (Gui) Cnorotega (H) Javai (Br) Asurini (Br) Chorotegamangue (CR) jcbcro (Pc) Atruahi (Br) Chorti (Gua) Jicaque H) Auca (E) Chulupi (A, Bo, Par) Jivaro (E. Pc) Aytw.ra (Bo, Ch) Chuj (Gua) lora (Bo) Cocoma (Pc) Baraw:ma (Br) Coconuco (Co) Kabixi (Br) Bare (V) Culla (A) Kalapalo (Br) Bari (V) Colorado (E) Kamayura (Br) Cora (M) Baure (Bo) Ka 1115A (Co) Boca Ncgra (Br) Cuna (Pan. Co) Kanjobal (Gun) Bocota (Pan) CuraraycPe) Katawi:in (Br) Bora (Pc) Curipaco (V) Kechki (Gua) 13orosese (Bo) Kikaptic (M) Diore (Br) Boruca (CR) Kogi (Cc) Bribri (CR) Galibi (Fr G, S) Kren-Akoro (Br) Gaviiies (Br) Kuben-Kragnotire (Br) Cainguii (A) Guahibo (V) Kuben-Kran-Kegn (Br) Cakchiqud (ES, Gua) Guaja (Br) Kuikuro (Br) Campa (Pc) Guajiro (H, Co, V) Kwaik, r (Co) Caiiaris (E) Guamhiano (Co) Canociros (Br) Guana (Par) LacandOn (Gua, M) Capanahua (Pc) Guanaco (Co) Lacand6n Choi (Gua) Carib (Guy) Guarayo (Bo) Leco (13o) Carifia (V) Guarayti (Par) Lengua (Bo. Par) Cashibo (Pc) Guatose (Bo) Cashinaua (Pe) Guatuso (CR) Machicanga (Bo) Cayabi (Br) Guayaki (Par) Machigiienga (Pe) Cayubaba (Bo) Guaycuru (Bo) Maka (Par) Chacobo (Bo) Guaymi (Pan) Makiritare (V, Br)

Country abbreviationsas follows: A = Argentina Co = Colombia Gua = Guatemala Pan = Panama Be = Belize CR = Costa Rica Guy = Guyana Par = Paraguay Bo = Bolivia E = Ecuador H = Honduras Pe = Peru Br = Brazil ES = El Salvador M = Máxico S = Surinam Ch = Chile Fr G = French Guiana N = Nicaragua V = Venezuela

65 Mak6 (Br) Patamona (Guy) Tiri6 (Br) Makusi (Guy) Paya (H) Tlaponeco (M) Mam (Gua) Peguche (E) Toba (A, Bo, Par) MandawAka (Br) Pemon (V) Tojolabal (M) Maopityan (Br) Pianokot6 (Br) Toronoma (Bo) Marinahua (Pe) PiapoLo (V) Totonaco (M) Marobo (Pe) Piaroa (V) Triche (M) Mashco (Pc) Pilagi (A) Trio (or Tiri6) (S) Mataco (A, Bo) Pi lapoe (E) Trumiii (Br) Matlatzinca (M) Pipil (ES) Txiciio (Br) Maya (Be, M, Gua) Piro (Pe) Txukahamac (Br) Mayongon#7 (Br) Poato (E) Tzeltal (M) Mayoruna (Pc) Pocomam (Gua) Tzotzil (M) Mazahua (M) Pocomchi (Gua) Tzutujil (Gua) Mazatec (M) Popoloco (M) Mbia (Par) Puinave (V) Ulva (N) Mehináku (Br) Puruborii (Br) Urubus-Kaapor (Br) Mentuktire (Br) Puruhay (E) Uspanteca (Gua) Mereyo (Fr G) Miquirano (H) Quechua (A, Bo, Ch, E) Waimirf (Br) Miskito (11, N) Quillacinga (Co) Waiwai (Br, Guy) Mixé (M) Quilotoa (E) Waiyana (Fr G, S) Mixrc (M) Quinchuqui (E) Wapisiana (Guy) Mocob: (A) Quindivana (E) Warao (V) Mop lin (Gua) Warrau (Guy) Moreno (H) Rama (N) Wauri (Br) Morotoco (Par) Remo (TA) Wayapi (Fr G) Movima (Bo) Mudjet;re (Br) Saguatoa (E) Xavante (Br) Sa:asate (E) Xikrin (Br) Saluma (Br) Nahua (ES) Xinca (Gua) Nahuaktui (Br) Sanapana (Par) Nahuatl (ES, M) Sape (V) Natnbicuara (Br) Saraguro (E) Yaba:na (13r) Seri (M) Yabarana (V) Ocayna (Pc) Shapra (Pe) Yagua (Pc) Opatoro (H) Shipibo (Pc) Yaminahua (Pc) Orejon (Pe) Sikina (Br) Yanacona (Co) Otomi (M) Siquisilie (E) Yanomam6 (Br, V) Otuqui (13o) Sirinayri (Pe) Yaqui (M) Sirineire (Bo) Yari (Bo) Pacaguara (Bo) Sidon() (Bo) Yarero (V) Piez (Co) Sumu (H, N) Yuko (Co) Pakaánovas (Br) Yumbo (E) Palikour (Fr G) Tabitirii (Par) Yupa (V) Pame (M) Tacana (Bo) Yuracare (Bo) Panare (V) Tapaytina (Br) Pano (Pe) Tarieti (Bo, Par) Zarnbo (14) Papago (M) Tarasco (M) ZatiluCo ( 3o) Parakanln (Br) Tepehuano (M) Zapotcc (M) ParLujano (V) Teribt (Pan) Zoque (M) Parikot6 (Br) Ticuna (Pc) Zumbagua (E) DISCUSSION

Moderator: Dr. Salzano, Iwonder whether proper injection of the solution. We have just there is perhaps a decimal point misplaced. You published a paper on these questions in Tubercle. estimate that there are 1,000,000 unacculturated Moderator: To be a little more specific, we do Indians in Central America and 980,000 in all know there is a high frequency of positive skin of South America. But when I go through yourresponse to histoplasmin among many of the treatment of Central America country by coun- Indian groups of the interior, in the absence of try, I simply do not f!nd those numbers. Where any tuberculin reaction. So I do not think that a;.e these 1,000,000 concentrated? in this particular instance cross-reactivityis a SaIzano: As I said, these are very preliminary problem. estimates. They make allowance for the very Cohen: I was using that only as an example. largegroups in Guatamala and Mexico on Unless one knows what the spectrum of sensi- which we do not know enough for mr: to set a tivity of a population is to a test that has been developed for a different population, one has specific figure. But I am sure that there should some questions about the significance. be some groups there that are sufficiently un- Hilleboe: I had the same question on this acculturated to be worth studying. small point, ano I wondered about two things. Cohen: How reliable is the tuberculin test in First, was anything done to see how much this population that you referred to in the case tuberculosis was really present inthe group Df central Brazil? Are you sure the 1.ndians with positive tuberculin tests, and whether or do not have histoplasmosis or some other condi- not there was positive sputum? Second, in thc tionthat wouldcross-react and you would group from whichitwas thought :.hatthe appear to be getting a tuberculin that may infection might have come, what was known unrelated to the exposure to the disease? about the amount of tuberculosis caused by SaIzano: Dr. Nutels will probably speak about mycobacterium tuberculosis and not atypical that. But what I can say is that there are not strains? only problems inrelation to thenonspecific Moderator: That is just about to be discussed reaction but o problems of reading and the by Dr. Nutels.

67 73 MEDICAL PROBLEMS OF NEWLY CONTACTED INDIAN GROUPS

Noel Nutels

Before entering on the topic of this paper, smallpox, venereal diseases, malaria, and other I should like to present a few questions on the diseases(15). meaning of "newly contacted Indian groups": In 1946, the Brazilian Government created To what period of time should this classification the Xingu National Park in an area of 22,000 be objectively applied? Might not a group classi- square kilometers in the upper basin of the fied as newly contacted have had direct or in- Xingu River, one of the largest tributaries of the direct contacts previously ignored? How do we Amazon, at an altitude of 820 feet (see Fig- define a tribe that has had one and only oneure 1). This region in the geographical heart rapid contactfor example, eighty years ago? of the country was considered by zoologists, How would we classify groups of Indians who,botanists, anthropologists, and other specialists living in isolation in their own primitive en- to be representative of primeval Brazil from the vironment, have had rare, intermittent, and brief standpoint of flora, fauna, and human condi- contacts during the past sixty years? I shall not tions. The present population is approximately attempt to answer these questions. My aim in 1,000 Indians distributed among the KamaiurA, this paper is simply to mention facts and report Kalapalo, Kuikuro, Wauri, Iawalapiti, Aueti, on my personal experience, not to interpret them. Trumai, Juruna. Aupatse, Nahukwi, Matipuhy, In considering the deficiencies of such a study Kajabi (since 1952), Suii (since 1960), Txu- it is necessary to take into account the difficultieskarram5i (since 1962), and Tchicão tribes(5). Table 1 shows the age distribution in the groups that prevailinthis enormous and primitive region, lacking in means of transportation and that have been examined. These fifteen tribes are autonomous. Some communication, where a diyersed and almost groups now dispersed are to be found among evtinct population still live as in the Stone Age. them. There are others, not yet contacted, living According to various authors cited by Wag ley in or around the Park. The tribes belong mainly (20), the native population of Brazil at the time tofourlinguisticgroups: Tupian, Cariban, of the discovery is estimated at 1,500.000. Nowa- Arawakan, and Tapuyan. There is also an iso- days this population, scattered over the national lated linguistic group, the Trumai. territory,isreduced to approximately 80,000 Access to the Park is almost exclusively by air, Indians living in tribal conditions. Among the for there are no roads and river transport is causes of thisdecline, infectious diseasesat poor. Only military and other government planes times used deliberately as a means of extermina- may land on the Park airstrips, and special per- tion by so-called civilized menhave perhapsmission is needed by all strangers wishing to been the most efficient(13).Many are theenter. Communications within the Park are documented instances of extermination of the limited by the fact that it has only one airplane, indigenous copulation through grippe, measles, a few canoes, and a radio station.

68 FIGURE 1. Location and distribution of tribes in Xingu National Park, Brazil. Inset shows location of tribes

Since its inception the Park has been directed relations between the Indians of the area and by Orlando Villas BOas, who is famous for his the visitors, mainly anthropologists, ethnologists, practical knowledge of the Brazilian backlands. psychologists, physicians, dentists, nurses, and Assisted by his brother Claudio, also well known others who do research in the Park or assist the in this field, he keeps a very close watch on the Indians in various ways.

TABLE I. Population of Xingu National Park, Brazil,in July 1967, by age group

AGE GROUPS

0-4 5-9 10-14 15-19 20-29 30 AND TRIBE NO. MEMBERS OVER

Kamaiuni 89 16 11 '1 9 18 24 Kajabi 152 32 21 15 13 35 36 Kalapalo 68 14 14 5 11 13 11 Juruna 49 10 8 4 6 11 10 Meinaco 35 5 6 2 8 5 9 Suiti 67 11 7 11 6 18 14 lawalapiti 34 2 8 3 3 6 12 Trumai 22 5 3 2 3 4 5 Aucti 40 9 8 6 3 14 Kuikuro 118 20 19 10 8 38 23 WaurA 62 17 14 1 7 15 8 Txukarram5i 170 36 21 19 16 33 45 Tchicio 53 9 4 5 9 22 4 Total 959 186 144 94 99 221 215

69 Is The first known contact between this area and myself saw were all of a benign nature, so I think that so-called Western civilization occurred in 1884, the gonococcus has vanished from the Xingu area.It when the first Karl von den Steinen expedition seems rather peculiar that I found no sign of it, not even congenital, and thatit had not spread throughout the passed through(18).Three years later Von den Indians or affected their sexual organs. Steinen came again. Expeditions since then in- Leprosy, syphilis, and tuberculosisaretotallynon- clude those of H. Meyer, in 1896; Max Schmidt, existent in the Xingu area. The nonexistence of tuber- in 1900; the Rondon Commission, directed by c-losisis of the greatest significance,since wherever Lieutenant Ramiro Noronha, in 1920; Percy Indians come into direct contact with whitc men, terrible devastation results. Fawcett, also in 1920; Heinrich Hintermann, It may 'also be that measles, scarlet fever, and small- in 1924; the Ro. don Commission, this time di- pox are likewise unknown in the Xingu area, although rected by Captain Vicente Vasconcelos, also in we a re not m) sure about their nonexistence at time 1924; Percy Fawcett again, in 1925 (his sec- of our visit as we are in the case of the previously ond expedition, during which he disappeared); mentioned diseases.Only in the case of smallpox can G. M. Doyott, in 1923; Vincent M. Petrullo, in we assureitsnonexistence, because we found no one displaying smallpox scars. 1931; and the National Service for the Protection of Indians, in 1940(1). In 1946 the white man took firm hold of the In 189E: Kari E. Ranke, a German physicianregion, though not in large numbers. Before and demographer who had participated in theentering the area that later be-ame the Xingu Meyer expedition, reported on the diseases he National Park, they were all given a medical had observed(12).His account is still relevant, examination. Between 1947 and 1950, in my and I feel it is worth quoting in full: capacity as a doctor in the Roncador-Xingu expedition, I imposed a sort of moral and sani- The diseases observed in10 Indian villages among approximately 800 to 1000 Indians arc the foilowinj: tary quarantine on every person entering the many healed fractures and one old haunch luxation but, region. Thus we were able to prevent venereal according to the anamnesis, not congenital; one case of diseases and to piotect the Indian women horn clubfoot, congenital; a very frequent skin disease,de- the "civilized" men who would have competed scribed as "Tinca imbriata,- from the Malayan Archi- with the native males. pelago and the Southern sea; numerous furuncles, most At the time of their first contact with tht. Ron- of them in the glutcal region; two cases of ithocy; one case of apparently parasitical tumor in the liver. which cador-Xingu expedition of 1946, about 25 Kala- had as a consequence an ascites, not very developed; a palo Indians died in an outbreak of grippe (5). kind of rheumatic disease of the joints; numerous cases Another outbreak was recorded in 1950 in the of malaria and malarial "cachexia" in children under 10 same tribe and alsointhe Karr ayunitribe, years old; and a fairly mild enteritis among newborn killing 12 persons this time. In June 1954 thcre infants. The occurrence of Ieucomas and staphylomas was extretaely frequent in the Xingu area. Among the was an epidemic of measles in the upper Xingu Bakairi from Kulisehu, none had been spared. (7). Neither from the recollections of the elders With ihe help of a Bakair: from Paranatinga,the nor from the unditions of any of the tribes were famous Antonio, who up to now has accompanied all we able to gather information about it. Every expeditions to the Xingu arca, I learned the history of Indian who came in contact with the Roncador- their existence. Once, after the second expedition to the Xingu arca, the Bakairi from Kulisehu went to Parana- Xingu expedition at that time was struck by tinga and from there proceeded to Rosario. They were that outbreak. Of the 654 patients, 114 died. wekomed, and, although they had no knowledge of the Among those who received medical care, the Brazilian language, they were immediately baptized and death rate was 9.6 per cent; among those who after a few days' stay they went back carrying a heavy load of gifts. could not be treated in time,it reached 26.8 One of the tribesmen acquired an ophthalmic blen- per cent. norrhagia in Rosario, which spread after his return to Following is a list, on which I shall make the Bakairi village on the Kulisehu and became a terri- no comment, of the diseases observed by us ble epidemic.All the inhabitants were affected; some died, othcrs survived with the loss of an eye or with and other doctors who have been in the area: leucomas. The numerous cases of conjunctivitis that I endemicmalaria (Anopheles darlingi, Plasrno-

70

.4 chum vivax and P. falciparum); Tinea imbrt- Pirquet modified: cutipuncture) and since 1960 cata; gastroenteritis; a purulent lung abscess ofwith purified tuberculin (PPD-Rt 23-1 UT), I unknown etiology; furunculosis; childhoodum- have investigated tuberculosis not only in the bilical hernia; arthritis deformans; helminthia- Park but alsoin other hiclianvillages, both sis; pemphigus foliaceus; gallstones; Blastomy-those contacted long ago and those that have cosis cheloidiana (Paracoccidioides Loboi) (3);just been contacted (Table 2). warts; polymorphous acne; pediculosis; Pityria- In 1952 the Kajabi Indians came to the Park sisalba; gravidicstriation; Pulex penetrans;from the Teles Pires River, in the valley of the conjunctivitis; pigmentary and cellular nevi (6);Tapajcis River, bringing with them Blastorny- leucoma; pinguecula; pterygium; pupillary seclu- cosis cheloidiana (Jorge LObo's disease)(3). sion; melanic pigmentation of the conjunctiva; This group had already been in contact with chalazion; cataract; ocular hypotension; talipes; white men in its origilial area. malarial hematological disturbances (17); de- In 1960 the Villas BOas brothers established formation of the auricular pinna due toa fight- contact with the SuiA tribe. This small group ing game called huca-huca; atrophy of the lower ofapproximately 80 Indians had had only limbs due to the ritual custom of prolongedcon- sporadic encounters, over athree-day period, striction in infancy; scapulo-humeral luxation;with the first Von den Steinen expedition in tegumentory leishmaniasiF(4); reversible pa-1834. At that time Von den Steinen estimated ralysis of the lower limbs due to ritual ingestion their number at 150. No other encounterwas of rnucund (Dolichos pruriens and/or Dolichosrecorded, even by Von den Steinen himself on urens); toxoplasmosis, recently discovered (2); his second expedition in 1887 (19). Thepres- filariasis (2); and various arboviruses, recentlyence of the Sui6 group on the upper Paranajuba discovered (2). River was known because oftheirperiodic Since I believe that the purpose of my par- attacks on other tribes, for which they were ticipation in this meeting is to tell about mygreaty feared. A few days after the Villas Mas personal experience in deali.ig with groups ofcontact had been established, we were able to recently contacted Indians,I shall concentrate include 42 members of the tribe in our tuber- on tuberculosis, which is the inain field of my culin survey (PPD-Rt 23-1 UT). Though at observations. Beginning in1952, at first with that time we did not have much suitable equip- X-ray and sometimes with old tuberculin (Vonment and had had little experience with PPD,

TABLE 2. Reactions to tuberculin test (PPD-Rt23-1 UT) inndian tribes ot Xingu National Park. Brazil. July 1960

TESTS READ

0-4 NiNt 5-9 SIM 10 MM NO.

TRIBE TESTED TOTAL NO. NO. NO.

Karnaiurti 76 76 74 97.4 2 2.6

Kajabi 63 63 60 95.2 1 1.6 2 3.2 Kalapalo 51 51 51 100.0 Juruna 45 45 45 100.0 Meinaco 38 38 36 94.7 2 5.3 7 Suia 42 42 34 81.0 1 2.4 16.6 lawalapiti 37 37 37 100.0 Trumai 17 17 17 100.0 Aucti 15 15 15 100.0 Kuikure, 3 3 3 100.0 Wauri 2 2 1 50.0 I 50.0

Total 389 389 373 95.9 5 1.3 11 2.8

71 77 we found eight positive rea-.:tions-one weak TABLE 4. Reactions to tuberculin test (PPD-Rt23-1 UT) and seven strong-among this population, for by age group among Suizi Indians of Xingu National an infection rate of 19 per cent. We also, on Park, Brazil, August 1966 that occasion, gave the PPD test-again for the TESTS READ first time-to the other tribes of the upper Xingu, finding infection rates of 3.2 per cent 0-4 5-9 10 NiNt MM MM among the Kajabi and 5.3 per cent among the AGE NO. Meinako. The former, as I have said, had had GROUP TESTF.DNO. 7-0 NO. NO. Je contact with civilized men before settling in the 0-4 8 8 100.0 - Park. The Meinako reactors, furthermore, had 5-9 10 9 90.0 - 1 10.0 spent a few months in the city of São Paulo (8). 10-14 13 11 84.6 2 15.4 The other tribes were to-ind to be nonallergic. 15-19 5 4 KO- 1 20.0 20-29 22 17 77.3 1 4.5 4 18.2 Subsequent research with X-rays revealed cases 30 and of lung shadows among the Sui6, and in 1967, over 13 7 53.8 - 6 46.2 for the first time, we were able to obtain from Total 71 56 78.9 1 1.4 14 19.7 them a positive sputum by BK. In spite of all the natural difficulties, we were able to take a culture of bacilli in a Su la solution to the Cen- vealed BK in their sputa. Moreover, we dis- tral Tuberculosis Laboratory inthe State of covered other strongly positive cases presenting Guanabara, where they are being studied by radiologic forms of the type that do not usually Professor Milton Fontes Magark and his staff reveal BK in direct sputum analysis, such as, ( Tables 3, 4, and 5). for example, two ganglionary forms. All were In 1965, 180 Indians of the Txukarram5i treated with INH, PAS, and SM (Table 8). tribe, fleeing the pressure of so-called pioneer In 1967 there came to the knowledge of the Park administrators the fact that a tribe of com- fronts(in this case a group of garimpeirns, pletely isolated Indians-the Tchikk-were be- prospectors of precious stones and metal), sought ing pressed by anothei group of prospectors. refuge in the Park. These Indians, who belong These Indians used to live on the banks of the to the Kayapó (Tapuyan) group, settled in a jatobá River, a subtributary of the Xingu. The place called Porori, on theleft bank of the few contacts they are known to have had con- Xingu River. When observed in 1962, this tribe sisted of attacks against other tribes living in had shown a low rate of tuberculin infection the Park, from whom they attempted to take (Table 6). In 1966, after the Txukarraniii had already settled in the Park,we classified 158 of TABLE 5. Reactions to tuberculin test (PPD-Rt23-1 UT) them in our thoracotuberculin file and examined by age group among Suii Indians of Xingu National their sputa (Table 7) (10). Threepersons re- Park, Brazil, July 1967

TESTS READ TABLE 3. Reactions to tuberculin test (PPD-Rt23-1 UT) among Suii Indians of Xingu NationalP.irk,Brazil, 0-4 5-9 10 June 1960 Nti4 NiNt sukt AGE NO.

GROUP TESTEDNO. 7.0 NO. NO. 470 TESTS READ 0-4 11 11 100.0 ^ 0-4 5-9 10 5-9 7 4 57.1 3 42.9 MM MM MM AGE NO. 10-14 11 8 72.7 3 27.3 GROUP TESTED NO. 15-19 6 4 66.7 2 33.3 NO. 510 NO. % 20-29 18 14 77.8 4 22.2 Adults 21 17 80.9 - 4 19.1 39 and Children 21 17 80.9 1 4.8 3 14.3 Over 14 6 42.9 - - 8 57.1 Total 42 34 80.9 1 2.4 7 16.7 Total 67 47 70.1 - - 20 29.9

72 8 TABLE 6. Reactions to tuberculin test (PPD-Rt23-1 UT) TABLE 8,Findings of tubreculintests,X-rays, and among Txukarramiii Indians of Xingu NationalPark, bacilloscopy in Txukarram5i tribeof Xingu National Brazil. 1962 Park, Brazil, August 1966

TESTs READ TUBERCULINTESTS

0-4 5-9 10 PULMONARY 0-4 5-9 10

Nt.Nt SHADOWS MM M M MM TOTAL AGE NO. 1 1 GROUP TEST ED NO. % NO. NO. (70 Minimum -- Moderate - 2(16) .-, Adul ts 19 17 89.4 1 5.3 1 5.3 Advanced 1* 1 1* 3 Children20 20 100.0 - - Pulmonary Ganglion 1 1 2 4 Pleural ---- Total 39 37 94.8 1 2.6 1 2.6 Total 2 2 6 10 women and utensils. In order to preventindis- No. X-rayed 105 11 32 148 Percentile of criminate and undisciplined contact with mem- pulmonary shadows 1.9 18.2 18.7 6.7 bers of our civilization, the administrators were *Positive sputum. abletoresettlethe whole 54-membertribe within the boundaries of the Park. We were then able to give these newcomers the tuberculin an epidemic would have occurred that would test at the moment of their arrival, with negative have presented differ.it foi ms such as the so- results. called infant form in adults and the rapidly In967 we repeated the research among theevolving acute and miliqry forms, similar to Txukarrarniii and verified, this time, an absence those found among the Senegalese soldiers taken of bacilli in the direct sputum analysis and the to France during the First World War (16). radiological regression of lung shadows in spite Such dramatic epidemics among primitive peo- of an increase in the general rate of infection. ples are often described in medical literature. This result was perhaps due to better equipment Nevertheless, we were able to verify that among and technique employed in the test (Table 9). the SuiA and Txukarram5i the forms were the We were surprised at both the clinico-radio- same as those that often occur among civilized logical and the epidemiological aspects of tuber- white people, whose natural and acquired re- culosis in populations so primitive as the Suia' sistance gives them means to thwart the evolu- and the Txukarramai. It was to be expected, tion of the disease. The X-rays presented here once the disease was found among them, that (Figure 2) leave no doubt, in my view, that

TABLE 7. Reutions to tuberculin test (PPD-Rt23-1 UT) TABLE 9. Reactions to tuberculin test (PPD-Rt23-1 UT) by age group among Txukarramii Indians of Xingu by age group among Txukarram5i Indians of Xingu National Park, Brazil, August 1966. National Park, Brazil, July1967

TESTS READ TESTS READ

0-4 5-9 10 0-4 5-9 10 MM ikiNt MM+ mid MM MM AGE NO. AGE NO. NO. NO. GROUP T EST ED NO. NO. /0 NO. GROUP TESTED NO. (70 %

1 1 0-4 36 31 86.1 1 2.8 4 11.1 0-4 34 32 94.0 3.0 3.0 5-9 25 19 76.0 2 8.0 4 16.0 5-9 20 13 65.0 1 5.0 6 30.0 10-14 15 10 66.7 2 13.3 3 20.0 10-14 19 13 68.4 1 5.3 5 26.3 15-19 16 14 87.5 - 2 12.5 15-19 16 12 75.0 - 4 25.0 1 20-29 28 18 64.3 1 3.6 9 32.1 20-29 33 16 48.5 3.0 16 48.5 30 + 38 23 60.5 5 13.2 10 26.3 30 + 44 26 59.1 2 4.5 16 36.4

Total 158 115 72.8 11 7.0 32 20.2 Total166 112 67.5 6 3.6 48 28,9 tuberculosis among these Indians, in its clinical, TABLE 10. Reactions to tuberculin test (PPD-Rt23-1 radiological, and even epidemiological aspects, UT) by age group among three tribes of Mato Grosso can be equated with that of peoples with a long State, Brazil, September-October 1965 experience with BK. TESTS READ Had these tribes at one time undergone a so- 0-4 5-9 10 called epidemic period of tuberculosis like other MM MM MM peoples? This question is not easy to answer. AGE NO. Isit possible that the BK responsible for the GROUP TESTED NO. % NO. % NO. disease among them is less virulent than that 0-7 584 55995.7 4 0.7 21 3.6 responsible for the disease among whites? This 8-14 634 537F4.7 37 5.8 60 9.5 hypothesis may be answered by the result of 15-49 746 52570.4 71 9.5 15020.1 Professor Magarao's research mentioned above. 50+ 145 9867.6 1913.1 2819.3 In any case, we believe that the Park is an Total 2,1091,71981.5 131 6.225912.3 appropriate environment for planned and con- trolled scientific research on the response of primitive populationstothe introductionof among the Terena (11), Kayua, and Kadiweu tuberculosis and other infectious or noninfec- Indians living in the southern part of Mato tious diseases by white invaders. Grosso State, who have been in close contact I consider it useful to conclude with Table 10,with white civilization for at least a hunderd which shows the rate of tuberculosis infectionyears (9).

REFERENCES 1. BALDUS) H. Bibliografia Critica da Etnologia de 1954 apresentado ao Diretor do S.P.I. (pp. 131- Brasileira. São Paulo, Comissão do IV Centethio da 141). Rio de Janeiro, Servico de Proteção aos Indios, Cidade de S5o Paulo, Serviço de Comemoraciks 1955. Culturais, 1954. 8. NUTELS, N., and L. V. DUARTZ. SUSA-1960: 2. BARuzzi, R. G. Determination de l'Etat de Cadastro tuberculinico na Area indigena. Separata Sante d'un Groupe d'indiens lsoles et des Mesttres da Rev. SNT 5(1 9):259-270. Rio de Janeiro, !961. Prophylatiques Necessaires a leur Preservation, Par- 9. NUTELS, N., J. A. NUNES DE MIRANDA) EJGEL, que Nacional do XingU, Brésil. Memoire présente I. YAMASAKI, and A. FRAGA HAUTEQUEST. A tividades l'Institut de Medicine Tropicale "Prince Leopold". do SUSA em 18 aldeiamentos indigenas do Sul do Anvers (Belgique), 1967. Mato Grosso. Rev. SNT 11(41), 1967. 3. BARUZZI, R. G., C. D'ANDRETTA JR.,S. CAR- 10. NUTELS, N., M. AYRES, and F. M. SALZANO. VALHAL, 0. L. RAMOS, and P. L. PONTES. Ocorrencia Tuberculinreactions,X-rayandbacteriological de blastornicose queloideana entre indios Caiabi.Kudies in the Cayap6 Indians of Brazil. Tubercle Rev. Into. Med. Trop.sãoPaulo 9(3) :135-142. 48(3), 1967. 4. CARNERI, I., N. NUTELS, and J.A. MIRANDA. I I. OLIVEIRA, R. C. DE. 0 processo de assimilacao Epidemia deleishmaniosetegumentarentreos dos Terina. Rio de Janeiro, Museu Nacional, 1960. indios Waurli do Parque Nacional do Xing6. Rev. 12. RANKE, K. E. Beobachtungen Ober Bevolke- Inst. Med. Trop. São Paulo 5(6) :271-272, 1963. rungsstand und Bevölkerunsbewegung bei Indianern 5. GALvAo, E. Cultura c sistema de parentesco das Centra1-Brasiliens.Correspondenz-Blatt der Deut- tribos do Alto Xing6. Antropologia (Rio de Janeiro) schenGesellschaft far Anthropologic', Ethnologic No. 14, 1953. und Urgeschichte XXIX (11), 1898. 6. MATTOS, R. B. Acuidade visual para longe e 13. RISEIRO, D. Convivio c contarninacio. Efeitos frequencia de discrornatopsia em indios brasileiros. dissociativos da depopulação prot ocada porepi- Descricão de alguns aspectos oftalmológicos em demias ern grupos indigenas. Sepa dt cologia indios examinados. Tese apresentada para o concurso (São Paulo) XVIH(1), 1956, de docencia-livre da cadeira de clinica oftalmologica 14. RIREIRO, D. Linguas e culth ,c do da Escola Pau lista de Medicina. São Paulo, 1958. Brasil. Separata de Educagto e Ldric. aais. 7. MOTA, J.L. DA. A epidemia de sarampo no Rio de janeiro, Centro Brasileirode Pescwisas Xingt;. Condensação do relat6rio em 10 de outubro Educacionais, 1957. 15. RIBEIRO, D. Politica indigenista brasileira. Rio 18. STEINEN,K. VON DEN. 0 BrasilCentral. de Janeiro, Ministerio da Agricultura, Servico de fraduck de Catarina Baratz Cannabrava. Ri3 de Informacties Agricolas, 1962. Janeiro, Companhia Edit6ra Nacional, 1942. (Origi- 16. RICH, A. R. Patogenia de la tuberculosis% Ver- nal edition Leipzig, Brockhaus, 1886) sion castellana por el Dr. 0. C. Croxatto. Buenos 19. STEINEN, K. VON DEN. Entre os aborigenes do Aires, Alfa, 1946, p. 651. 17. SILVA, M. P. DA. C'ontribuição para o estudo do Brasil Central. Tradu0o de Egon Schaden. Sepa- sangue perifdrico e d i medula 6ssea em indios do rata renumerada da Revista do Arquivo 1:XXXIV Alto-Xingti. Tese de doc8ncia-1ivre de hemato1ogia LVIII. Sfio Paulo, Departamento de Culiura, 1940. apresentada ao Departamento de Medicina. So 20. WAGLEY, C. Races and Classes in Rural Brazil. Paulo, 1966. Paris, UNESCO, 1952, p. 123.

76 DISCUSSION

Willard R. Centerwall: A Recent Experience with Measles in a "Virgin-Soil" Population'

I have elected to use the time allotted for thenorth latitude, deep within the tropical rain presentation of recent experiences we have had forest of the Orinoco-Amazon River basin of on the effect of both measles and measles vaccine Venezuela andBrazil. The expedition was in a "virgin-soil" population of South America.oriented toward the collection of ethnographic, In January of this year a team of seven scien-genetic, medical, dental, demographic, and epi- tists, representing the United States and Vene- demiological data. Three members of the team zuela, made an expedition into the Yanomama ale present here today: Dr. JamesNeel, the Indian territory. The map shows the approxi- expedition leader, who needs no introduction; mate location of the territory, which covers an Dr. Napoleon Chagnon,theanthropologist area extending from the Equator to 5degrees whose work, study, and life among these In- dians since 19642was so essential to our mis-

N' ift sion; and myself, a pediatrician with interest in 751 t-NoynAreimINT or MfANC.5 and some experience withthediseases and LIVN TH4- y44,0Abtivo rihwIropy or n Otqn, ,v,hyRk nutritional disorders of the tropics. This was the third such expedition led by Dr. Neel into this general region, the first having taken place in 1966 on the Venezuelan side and the second in 1967 on the Brazilian side. The present expedi- tion was in essence a cooperative venture of the University of Michigan and the Venezuelan Institute for Scientific Research, and in particu- lar Drs. Layrisse and Tulio Arends ofthe Institute. Both are present at this session also; thus, if what I say in the next few minutes generates questions, there are several resources available for possible answers. There are believed to be about 10,000 Yano- mama Indians living in more than 100villages scattered throughout the region. They are very

IgNC11, VI4N,g5 primitive forest-dwellers who practice a slash- - APAR, *AkOM1M0 60,6,04my -,m,1110.31A7,69* -MfAiLIS fwfwOOtcea wrc ,,r 4.'14 YA40.4.0001,ihry vmooga I Investigationssupportedinpart byU.S. Atomic Ne'' Energy Commission Grant AT ( 11-1)-1552. 61' 2Chagnon, N. A. Yanomamo Warfare, Social Organi- zation and Marriage Alliances, Ph.D. thesis, The Univer- sity of Michigan (1966).

77

82 and-burn agriculture along with hunting andbe at the mission in connection witha study gathering. Although some mission stations have on thyr Did physiology. been established along the Upper Orinoco and The boy never developed the characteristic its tributaries since early in the past decade, themorbilliform rash, so the differential diagnosis majority of these Indians are as yet uncontacted from any of a variety of "jungle fevers"was and unaffected by the outside world. uncertain; nevertheless, 31 Indians and 9 Bra- The typical Yanomama village is a circular zilians at the mission were vaccinatedat once stockade, with families livingin hammocks with no gamma globulincoverage. It turned out under the shelter of the enclosure. In generalthat the boy indeed did have measles and that the people are healthy and handsome, wellpro- visitors to the mission meanwhile had returned portioned and strong but almost pigmoid into their own villages carrying the virus inits stature. Their cheerful, friendly countenances incubating form with them. Thus starteda full- bely the aggressiveness and hostility for which scale epidemic, which, by the timeour last team they are sometimes known. member had left the field three months later, Because measles antibody studies in 1966 and had spread up and down river alonga hundred 1967 demonstrated that this was a virgin-soil miles of waterways and tributaries and adjacent tribe and because measles was known to have inlandregionsinvolving no fewer than15 penetrated into the nearby regions, practically villages. surrounding the Yanomama territory by 1967 Within a few weeks we had exhaustedour (see map), we brought along withus into South vaccine supply. Observationson these Yano- America 2,000 doses of donated Edmonston-mama Indians of the effect of the wild measles strainattenuatedlivemeaslesvaccine,plus and of the measles vaccine with and without gamma globulin.3Half ofthesedoses we theconcomitant gamma globulin formthe diverted to the mission stations on the Brazilian basis of this report. Since itwas neither con- side, where measles was beginning to penetrate ceived nor designedas a scientific study in itseif the region, and the rem inder went with us. We and since our data-collection responsibilitit!s did had intended to var.-,late and to advise the not permit a prolonged stay in any one missionaries on the ,,rocedure toward the end our observations are of necessity disjointed and of our scientific studies. We did not anticipatefragmentary, but still, we believe, of sufficient encountering mt:asies during our two-to-three-interest and value to warrant presentation. month expedition. It was thus a fortuitousoccur- Most of the Indians and also the several rence that our an ival into the region was almost Brazilians we observed became very ill and pros- simultaneous with the introduction of measlestrate with their measles. In several Indian vil- (we believe for perhaps the first time) into this lages where documentation was possible by di- territory. rect examination, from 24 to 43 per cent of A party of Brazilian workers had, the week the victims had frank bronchopneumonia, with before, come across from the Rio Negro region onset usually during the acute attack and some- in Brazil to the south. They were at the Ocamo times during the recovery phase. This seemed mission, at the junction of the Orinoco and to respond well to antibiotic therapy. We were Ocamo rivers, when a 14-year-old boy among unable to evaluate the possibly adverse effect of them became sick with a high prostrating fever. a respiratory flu-like epidemic, which had imme- He was seen shortly after the onset by Dr. diately preceded and sometimes co-existed with Marcel Roche, Direc, - of the Venezuelan Insti- the measles epidemic. The complication of otitis tute for Scientific Research, who happened tomedia appeared to be practically nonexistent. We were particularlyimpressed withthe 3 Wc arc most gratefultoParke, Davis and Co., Phillips Roxane Laboratories, and Lederle Laboratories devastating effect of measles initsnear-total for their donation of the vaccine. involvement of the Indian community. When

78

83 parents and children were simultaneously in- Arctic 5; in the first of two outbreaks the death volved, there was a drastic breakdown of both rate was 7 per cent, whereas in the second, with the will and the means fornecessary nursing better organization and provision of medical care. We have even seen several instances where care, it was only 2 per cert. three generations of Indians were simultane- The Jse of measles vaccine permittedus to ously ill with measles. see some of its effects on the Indians. In villages The reaction of those not already prostrated where we could observe andmeasum the effects in a fatalistic depression was usuallyone of at their height, we found no real complications panic. Sometimes the well members of thebut considerable reaction, especially in terms of village even abandoned the sick, and normalfebrile response. That thisresponse was asso- community structure and function were lost.ciated with the production of protective measies Village fragments often fled to other villagesantibodies was borne out clinically. Toour for haven, thus increasing the spread of theknowledge, no persons vaccinated priorto the disease. exposure to measles later developed the disease. In three villages with approximately 170cases This protective value of the vaccine showedup of measles there were 29 deaths, a rate of 17.1 in peculiar ways, as for example in thecase of per cent. Most of these were due to associatedan unvaccinated mother who Ezcame very iii pneumonia, complicated by dehydration. Thiswith measles while her previously vaccinated despite some measure of applied, though less and healthy youngver was playing about and than optimum, nursing care. Ina fourth village, pestering her for attention she could not give. the authors were present for two days at theThis is just the reverse of the situationcom- peak period of measles reaction involving nearly monly seen in civilized societies. 40 Indians, a majority of them extremely Searching the literature for reports ofreac- Vigorous and dramatic treatment included fliidtions of other peoples to similar vaccine both administration, cooling measures, aspirin, andwith and without concomitantuse of gamma long-acting antibiotics. No deaths occurred dur- globulin, we observed that these Indians ingen- ing our stay, but we did hear later that subse- eral seem to react witha greater febrile response quently two Indians died (approximately 4per than has been reported in North American Cau- cent mortality). casians,"Europeans,'Icelanders,"Africans"' ° These and other observations supportedour and Micronesians.'" Again, however,as with feeling that difference-, in medical and nursing care play the major role in the measles-mortality Pert, H. F. W., and F. P. Nagler. Measles in the rate among the Indians. In this connection, it Canadian Arctic, 1952. Canad. I. Pub, Health 45:146 .156. 1954. shouldberecalledthatthemortality from " Krugtnan. S., J. P. Giles, A. M. Jacobs. and H. Fried- measles among Caucasians in the United States man. Studies with live attenuated measles vi,us vaccine. only half a century ago was in excess of 3per Am. 1. Dis. Child. 103:353-363, 1962. T Cockburn, W. C.,J. Pecenka. and T. Sundaresan. cent; it has steadily dropped to a present-day WHO-supported comparative studies of attenuated live low of less than 0.1 per cent. Nearly two thirds measles virus vaccines,Bull. Wld. Hlth. Org. 34:223 of this reduction occurred in the period before 231, 1966. " Gudnadotter, M.. and F. L. Black. Response of adults the advent of modern chemotherapy and anti- in Iceland to live attenuated measles vaccine. Bull. Wld. biotics.' Other evidence that the death rate in Hlth. Org. 30:753-762, 1964. highly susceptible groups can be substantially Mey'r, H. M.. Jr., D. D. Hostetler, Jr., B. C. Bern- heim, N. G. Rogers. P. Lambin, A. Chassary, and J. E. reduced by eleinentary organization and medical Smadel. Response of Volta children to live atten hated care emerges from a 1954 study on a measles measles virus vaccine. Bull. Wld. Hlth. Org. 30:760-781. epidemic among the Eskimos of the Canadian 1964. Brown, P.. M. Basnight, and D. C. Gajdusck. Re- sponse to live attenuated measles vaccine in susceptible 4 Langmuir, A. D. Medical importance Of measles. island populations in Micronesia. Am. 1, Epidem. 82: Atn. 1. Dis. Child. 103:224-226, 1962. 115-122, 1965.

79 84 the wild measles, we are unable to define accu- be7ome "virgln-soir populations and have then rately the role of concurrent respiratory infec- experienced an epidemic of measles. The classic tions in enhancing the reaction to the vaccine. example is the epidemic Dr. Peter Panumre- Summing up, although the evidence for ge- ported from the Faroe Islands. A Danish popu- netic influences in mammalian resistance to a lation had been free of measles forsome 65 wide variety of diseasesis of course incontro- years and then was stricken much as an Indian vertible, the present data nevertheless suggest village is stricken. that given a comparable previous disease experi- I was most impressed by Dr. Nutels' remarks ence (other than measles), similar care when ill,about tuberculosis and what I took to be his and a less resigned attitude toward the prospect questioning attitude toward how differently the of death, the death rate from measles in a pre-Indian reacts to it. Now, quite aside from its viously unexposed Indian population would not theoretical interest, this seems tome a matter of be much greater than in a group of "virgin-soil" great practical importance. If we believe that civilized Caucasians whose ancestors had been the Indian is more susceptible to tuberculosis repeatedly exposed to the experience of measles. or measles or pertussis, we are apt to be satisfied There is thus, in our opinion, only very limitedwith less than optimal clinicalresults in the evidence for a greater innate susceptibility ofmanagement of those diseases. But if we think the Indian to this disease. It seems likely that he is no more susceptible than we are, then we a re-evaluation is in order with respect to thewill not settle for anything less than the best. primary susceptibility of the Indian to certainThis is perhaps an important point that has been other diseases of civilization, such as pertussis, lost sight of in the past. smallpox, and tuberculosis. McDermott: I think both of these presenta- tions represent very fine contributions to our General Discussion accumulating knowledge on thissubject.In previous meetings of this committee, several of Moderator: As I am sure you gathered from usamong them Dr. Neel and Iexpressed the that capsule presentation, there were some prob- belief that evidence that a genetic predisposition lems in conducting our-elves like calm and de- played a significant part in measles and tubercu- tached observers in the midst of a measles epi-losis should be very sharply challenged. Indeed, demic. And when it was all over, we realized we did not believe it. that in many respects we had failed to document It seems to me the evidence in both these this experience adequately. studies is in that direction. As Dr. Nutels said, Dr. Nutels in his presentation gave data on itis conceivable that the strain of mycobacteria how the mortality rate in an Indian group can either may not have been tuberculosis or may be lowered by even what I think he would agree be attenuated, but I do not think that is very islessthan adequate medical care. Certainly this was our experiencethat the mortality rate likely.The evidence he presented makes it for Brazil and Venezuela in this epidemic would perfectly credible that these people did indeed run under 10 per cent, which is a half to a third have tuberculosis infection anddisease. Our of what is often reported. It is very difficult to challenge to notions of genetic predisposition say how much lower this rate would go if these arises from our greater knowledge nowadays of were Indians given the kind of care that a how these two particular infections, measles and Caucasian would receive today in Europe or in tuberculosis, are spread. Both can be spread the United States, but we believe it would drop (though this is not necessarily the only way) by down to a few per cent. what is known as the airborne route. When this The other side of the coin, of course,is is possible a whole constellation of circumstances revealed by Caucasian populations thatafter having to do with the culture, and f n particular centuries of exposure to measles have somehow the material culture and way of living, then

80 obtains: So far as measles is concerned, we ilust munity against infection with tubercle bacilli. remember first of allthat it may be a very By this I mean that practically all members of severe disease in any adult. Adults withmeasles a group exposed under the propercircumstances, are just as sick as the adults Dr. Centerwall was irrespective of their diet or way of life or level describing. of civilization or anything else,will become Secondly, the incidence of complications ofinfected. We know of course that infection leads measlesand the complications are what people to disease in only a minority of those infected; die ofcan be very high, depending upon thethe exact percentage is not known. If we had, age at which the infection is acquired. Seventy- let us say, 10 per cent diseased in a society that five per cent of the post-measles complications in was virtually 100 per cent infected, we might New Delhi, for example, where they have largesimply think that 10 per cent is an awful lot of numbers, occur under the age of 18 months. In tuberculosis and hence that this group of people, the particular circumstance of primitive societies this race, is especially predisposed to it, whereas with high sustained fertility, a high proportion in point of fact they may have no more tubercu- of the population exposed is extremely young.lous disease per infection than any other group. As for tuberculosis, I think the evidence is It is simply that the particular circumstances of even clearer, because work over decades has their society allow infection to be more prevalent. proved the need to distinguish between tubercu- Now with both tuberculosis and meerles, and losis infection and tuberculosis disease. It is only certainly with measles in infancy, there is of since the infection has become less prevalent course an additional factor: nutrition. Malnutri- that we have been focusing on it and hence have tion and lack of high-quality protein are char- numbers on which to base greater understanding. acteristically present in the so-called unaccul- Tuberculosisinfectioncan be spread and turated societies. What I am really saying is that probably usually is spread by the airborne route. our knowledge of thepathogenesis of both in- This means several things. One is thr.,t the infec- fections now gives us perfectly credible alterna- tious agentin this case the tubercle bacilli tive explanations for what has been seen without can be suspended in the air and in an enclosed its being necessary to invoke the idea of a sig- space will remain in that air whether a human nificant hereditary predisposition. is there or not; anyone entering the room could Behar: How was the nutritional status of the acquire the infection. How long the air will stay children in that population determined? poisoned is net known, but it is an appriable Centerwall: By simple inspection and physical intervala matter of an hour or two. And the examination. The nutritional status of the 2.hil- actual event of infection takes place presumably dren was remarkably good as a whole. There in a flash. This is relevant to contacts with pre- were a few areas in which there wasactive viously unexposed populations. malaria, and some of the children were sick with In any case, the evidence now is quite con- that. But in general they were fine. There was vincing that given the proper physical circum- no evidence of the proteindeficiency seen in stances for the transfer of tubercle bacilli from India and other tropical areas of the world. one person to another, circumstances such as are Moderator: The next part of our program is frequently seen in unventilated huts in aboriginaldevoted to some special medical problems af- or primitive societies, there is no inherent im- fecting Indian populations.

81 THE PROBLEM OF GALLBLADDER DISEASE AMONG PIMA INDIANS

Thomas A. Burch, Leonard J. Comess, and Peter H. Bennett'

Introduction logic study (5) has confirmed the higher preva- The etiology of gallbladder disease is uncer- lence of "clinical gallbladder disease" among tain, though the problem has excited consider- Pima Indians than was established by identical able speculation and controversy. Diet (11),criteria among the predominantly Caucasian female hormones (10), obesity (16), and dia-community of Framingham, Massachusetts (7) betes (13) have all been said to be associated (Table I). with cholelithiasis. In addition, differences in the The purpose of this paper is to report pre- frequency of gallbladder disease among various liminary data on a partly completed epidemio- racial groups have been noted. On the basis of logic study of gallbladder disease based on chole- autopsy data Caucasians have been reported to cystography in a population sample of Pima have ahigher frequency of gallstonesthan Indians, a Uto-Aztecan tribe living in the des,...rt. Negroes (6), and several studies based on hos- of the southwestern United States. pital admission rates have suggested that Ameri- Methods and materials can Indians have a still higher prevalence of gall- Sample selection bladder disease (9, 12, 15). A recent epidemio- Since 1963, the Clinical Field Studies Unit of Presented by Dr. Burch. the National Institute of Arthritis and Metabolic

TABLE I. Prevalence ot documented gallbladder diseasein Framingham community and among Pima Indians

AGE AT ENTRY FRAMINGHAM GROUP PIMA INDIANS INTO WITH DISEASE WITH DISEASE STUDY NO. NO. (YEARS) AT RISK NO. AT RISK NO.

Males

30-39 832 1 0.1 51 1 2.0

40-49 779 7 0.9 45 1 2.2 50-62 725 23 3.2 57 7 12.3 n

Total 2,336 31 1.3 153 9 5.9 n Females 30-39 1,037 24 2.3 45 16 35.6 n 40-49 963 58 6.0 51 20 39.2 50-62 873 88 10.1 62 21 33.8

Total 2,873 179 5.9 158 57 36.0n

n p < 0.01.

82 Diseases has been performing prospective studies each respondent verified that he or she had fol- of the natural history of arthritis (2) and dia- lowed the instructions. No attempt was made betes mellitus (14) among Pima Indians living to cleanse the bowel with laxatives prior to the on the Gila River Indian Reservation in Arizona cholecystography. (see Figure 1). Initially two radiographs, a 14" x 17" PA A house-to-house census of the Gila Riverscout film of the abdomen and a 10" x 12' Reservation was compiled, using aerial survey oblique film of the right upper quadrant, were photographs to ensure thatallhouses were taken on each respondent. If the gallbladderwas visited. Attempts are made to examine each identified on either of these views,a 10" x 12" reservation Indian aged fifteen years and over upright PA and a 10" x 12" right lateral decubi- every two years. This biennial examination in- tus view of the right upper quadrant were ob- cludes a 75-gram oral glucose tolerance test; a tained. After this, most respondents were given history and physical examination pertaining pri- a fatty meal consisty of 30 cc. of Cholestim 3 marily to diabetes and arthritis; radiographs ofgallbladder stimulant and either the uprightor the chest, cervical spine, hands,feet,pelvis, the decubitus film was repeated. thigh, and calf; a12-lead electrocardiogram; Respondents in whom no gallbladder could photographs of both optic fundi;a bio-thesi- be identified on the scout and oblique films were ometer examination (quantitative test of vibra-considered to have a nonvisualizing gallbladder, tory sensation in both great toes and index fin- and an attempt was made to performa repeat gers); a urinalysis; and determinations ofserum cholecystogram on them four to six weeks later. cholesterol, creatinine, uric acid, hematocrit, and Clinical gallbladder disease at surgery or by rheumatoid factor by sheep-c:ellagglutination cholecystography had been documented previ- and bentonite flocculation titers. In addition, the ously in 60 of the 258 members of the sample blood of each subject examined has been typed who have attended the NIAMD Clinic since the for several blood groups. beginning of the gallbladder study. An oral A sample of 600 Pima Indians was selectedcholecystogram has been performed on an addi- with random numbers for a study of gallbladder tional 132 of these respondents. The present re- disease. This sample consisted of 50 males and port analyzes the findings on this subsample of 50 females in each of the six decades between 192 Pima Indians. 15 and 74 years of age. Since the gallbladder Of the remaining 66 respondents, 61 did not study was started on 1 February 1967, 258 of have a radiologic examination-8 because of a the 600 members of the sample have attended contraindication to the procedure, 43 because the Unit for their comprehensive biennial ex- work or school made it impossible for them to amination. The subjects were selected according return at the specified time, and 10 because they to their age at the time of the census (as of 1 refused cholecystography. Five respondents were January 1966), but in the present analyses theyexcluded because of the poor technical quality have been grouped according to their age at the of their radiographs. time of the examination for gallbladder disease. Criteria for Gallbladder Disease Cholecystogram procedure The diagnosis of previously documented gall- Eight Telepaque 2 tablets were ingested on the bladder disease was based on either a hospital evening prior to examination and the respondent record of gallbladder surgery, the patient's his- was instructed to take nothing thereafter by tory of having had gallbladder surgery, or a mouth except water until the cholecystogram hospital record of an abnormal cholecystogram had been completed. Prior to the examination, (stones or a nonvisualizing gallbladder).

2 lopanoic acid manufactured by Winthrop Labora- " Distributed by Gencral Electric X-Ray Department, tories. New York. Milwaukee.

83 L. *SALT LAKE CITY

1431

UNITED PUEBLOS

ALEinuat

M E X I C

Hi FIGURE 1. Indian reservations of southwestern United States. Gila River Reservation is a lower left.

84 The cholecystograms performed at the Clini- TABLE 3. Diagnosis of gallbladder disease in sample cal Field Studies Unit Clinic were analyzed for METHOD OF DOCVMENTATION MALESFEMALES TOTAL the patient's clinical record by a radiologist of the Division of Indian Health. All radiographs Total examined to date 56 136 192 were reanalyzed by one of us (L. J. C.) prior to Normal cholecystogram at 79 this report without knowledge of the age or sex CFSU Clinic 35 44 Number with gallbla&kr disease 21 92 113 of the respondents. A disagreement in interpre- Cholecystectomy 5 41 46 tation was noted in only eight instances; these Abnormal cholecystogram at cases were carefully re-evaluated (by L. j.C.) hospital 1 13 14 without reference to the previous readings and Gallstones 2 2 7 the new interpretation was used. Two nonvisualizations 7 One nonvisualization 4 5 Abnormal cholecystogram at Results CFSU Clinic 15 38 53 Prevalence of gallbladder disease Gallstones 3 14 17 Two nonvisualizations 7 9 16 Table 2 shows that gallbladder disease was One nonvisualization 5 15 20 extraordinarily prevalent in both male and fe- male Pima Indians. In males the disease fre- quency increased with age, with 66.7 per centbladder radiograph. Thus, 15 (30.0 per cent) of of those aged 65 years and over affected; in 50 Pima males without known gallbladder dis- females the prevalence of disease was strikingly ease had abnormal cholecystogram.Of 92 af- high after the age of 24 and reached a remark- fected females, 54 were previously known; thus, able 92.6 per cent in those 65 and over. abnormal radiographs were demonstrated in 38 As Table 3 indicates, 6 of the 21 Pima males (46.3 per cent) of the 82 Pima females without with disease had previously been diagnosed by previously diagnosed disease. cholecystectomy or an abnormal hospital gall- In 5 males and 15 females who were examined at the Clinical Field Studies UnitClinic, the TABU 2. Prevalence of gallbladder diseasein Pima diagnosis of gallbladder disease was based on Indians only a single nonvisualization. However, when cholecystograms were repeated on these 20 cases, WITH GALLBLADDER 17 (85.0 per cent) had either second nonvisuali- DISEASE NO. zations or gallstones on the repeat radiographs. AGE EXAMINED NO. Morbzdity from gallbladder disease Males 15-24 5 0.0 In Table 4 the clinical importance of gall- 25-34 6 0.0 bladder disease among Pima Indians is shown. 35-44 9 11.1 Among males, "silent" gallstones were relatively 45-54 10 4 40.0 55-64 11 6 54.5 frequent: only 6 (28.6 per cent) of the 21 af- 65 and over 15 10 66.7 fected males related "gallbladder symptoms" 4 Total 56 21 37. or had a history of such symptomsnoted in their medical records. Among the females, however, Females gallbladder disease resulted in considerable mor- 15-24 11 9.1 bidity: 39 (42.4 per cent) of the 92 affected 25-34 18 13 72.2 35-44 33 22 66.7 females had had fu previous cholecystectomy and 45-54 28 19 67.9 an additional 25 (27.2 percent) had had gall- 55-64 19 12 63.2 65 and over 27 25 92.6 4 Defined as an affirmative answer to the question "Have you ever had pain inyour stomach or vomited Total 136 92 67.6 aftereating certain kinds offood?"

85 00 TABLE 4. Morbidity from gallbladder disease in Pima Indians

MALES FEMALES TOTAL

NO. NO. NO.

Total examined 56 100.0 136 100.0 192 100.0 With gallbladder disease 21 37.5 92 67.6 113 58.9 Postprandial abdominal pain or nausea 6 10.7 64 47.1 70 36.5 Cholecystectomy 5 A.9 39 28.7 44 22.9 Abn. cholecystogram (hospital) 1 1.8 13 9,6 14 7.3 Abn, cholecystogram (CFSU) 0 0.0 12 8.8 12 6.3 No postprandial distress 15 26.8 28 20.6 43 22.4 With no gallbladder disease 35 62.5 44 32.4 79 41.1 Postprandial abdominal pain or nausea 5 8.9 7 5.1 12 6.3 No postprandial distress 29 51.8 33 24.3 62 32.3 No information 1 1.8 4 2.9 5 2.6 bladder symptoms but still retained their gall-diet, pregnancy, obesity, diabetes, and serum bladders. Thus, 64 (69.6 per cent) of the 92 cholesterol-will be reported when a higher pro- affected females in this sample were having or portion of this sample has been examined. had had clinical manifestations of gallbladder A review of the medical records of 575 mem- pathology at the time of the examination. The bers of the sample for evidence of documented actual magnitude of gallbladder disease as agallbladder disease and possible associated con- clinical problem among Pima women is bestditions has previously been reported (5). In illustrated by the fact that 64 (47.1 per cent) of that study gallbladder disease was found to be the 136 females in this sample had symptomatic related to increased parity but not to diabetes, gallbladder disease. body weight, or serum cholesterol except that in older Pima males ther.; was an association Discussion with diabetes. These data must be interpreted cautiously, since the present study indicates that These data, though as yet incomplete, indicate nearly a third of the Pima males and nearly that the gallbladder was an extremely frequent half of the Pima females without documented site of pathology among Pima Indian females. clinical gallbladder disease actually hadgall- Gallbladder disease was responsible for numer- bladder pathology. ous hospitalizations, which often culminated in In order to advance the study of cholelithiasis a major surgical procedure with its attendant and to further explore the problem of gallstones morbidity and slight mortality. Unfortunately, in Indians, it would be desirable to supplement we have no data on complications that may have compounded the morbidity from gallbladder the epidemiologic studies currently being per- formed among the Pima with selected biochemi- disease in the members of this sample. Gallbladder disease was less common amongcal and physiological investigations. Pima males than among the females. Fewer Fundamental to the concept of gallstone for- males admitted to symptoms and fewer had had mation is the chemistry of the hepatic and gall- surgery. The earlier onset of gallbladder disease bladder bile at the time stones are first being and the more frequent occurrence of biliaryformed. In mice it has been possible to alter bile symptoms in females may perhaps be due to the composition and to produce gallstones by feed- effects of female hormones (10) on gallbladder ing "lithogenic diets" (3). The biochemistry of physiology. human hepatic bile and the significance of the An analysis of other factors possibly cont ')ut-changes found in pathologic bile have recently ing to gallbladder disease in the Pima-5u( as been reviewed (I). Since other groups of Ameri-;

86 "1 can Indians appear to have a high rrevalence of It has been reported that peptic ulcers are rare the disease, it is possible that the bile of Ameri- arr^ng southwestern Indians (9). This observa- can Indians, perhaps as a rault of genetically tion may be relevant to the excess of gallstones determined factors, differs fundamentally in its among Indians, since a recent study found gas- composition from that found in other racial tric secretion to be diminished in the majority of groups. To investigate this possibility, specimens patients with cholelithiasis (4). Further studies of bile from Indians both with and without gall- relating gastric secretion to gallbladder function bk dder disease should be analyzed. These could in Pima Indians appear to be indicated. be obtained from the gallbladder at the time of laparotomy for some unrelated condition and Summary at the time of cholecystectomy. Bile obtained at A preliminary report of a survey based on autopsy would be suitable for study only ifit oral cholecystograms on a population of Pima could be determined that its composition in the Indians from Arizona has been made. The prev- post mortem state was similar to bile examined alence of gallbladder disease among those ex- before death, and particularly if the information amined to date was significantly higher than had could be supplemented with pertinent ante mor- previously been reported, with 2 1 (37.5 per cent) tem data. An analysis of the constituents of gall-of 56 males and 92 (67.6 per cent) of 136 fe- bladder bile and their relation to age and sex in males affected. Clinical symptoms attributable this gallstone-prone population, together with a to gallbladder disease were found among 10.7 comparison of bile from Indians w;th that of per cent of the 56 males and 47.1 per cent of the other racial grups, might add significantly to 136 females. Nearly a third of the males and knowledge concerning the biochemical factors nearly half of the females without previously predisposing to cholelithiasis known disease had abnormal cholecystograms. The Pima Indians have low levels of serum Further biochemical and physiologic studies cholesterol (5), even though their diet seems to to supplement the current epidemiologic investi- be high in saturated fats (8). It is possible that gations are suggested. an unusually efficient mechanism forclearing serum cholesterol may be present, which might Acknowledgments be mediated through exaggerated biliary excre- We wish to thank the Tribal Council of the Gila River tion of cholesterol. Studies of the intake and Indian Community and the Division of Indian Health of absorption of dietary fats and of the synthesis the United States Public Health Service for their encour- agement and assistance.Special thanks are duc to Drs. and excretion of cholesterol might give :1_11es to W. J. Held and Jon E. Gustafson, the radiologists at thc the etiology of gallbladder disease in ;.his ;.!roup. Phoenix lidian Hospital who read the cholecystograms.

REFERENCES 1. BOUCHER, I. A. D., and J. W. FRESTON. The 5. COMM, L.J.,P. H. BENNETT, and T. A. etiology of gallstones (hypothesis). Lancet i:340-344, BURCH. Clinical gallbladder disease in Pima Indians. 1968. Its high prevalence incontrastto Framingham, 2. BumiN, J. J., T. A. 13tAcx, and W. M. O'BRIEN. Massachusetts. New Eng. I. Med. 277:894-898, 1967. Influence of genetic and environmental factors on 6. CUNNINGHAM, J. A., and F. E. HARDENBERGH. occurrence of rheumatoid arthritis and rheumatoid Comparative incidence of :holelithiasis in the Negro factor in American Indians. Bull. Rheum. Dig. 15: and White races. A study of 6185 autopsies. Arch. 349-350, 1964. Intern. Med. 97:68-72, 1956. 3. CALDWELL, F. T., K. LEVITSKY, and B. ROSEN- 7. FRIEDMAN, G. D., W. B. KANNEL, and T. R. BERG. Dietary production and dissolution of choles- DAWBER. The epidemiology of gallbladder disease. terol anstones in the mouse. Amer. I.Physiol. Observations in the Framingham Study. I. Chronic 209:473-478, 1965. Dig. 19:273-292, 1966. 4. CAPPER, W. M., T. J. BUTLER, J. 0. KILBY, and 8. HESSE, F. G. Dietary study of Pima Indians. M. J. GinsoN. Gallstones, gastric secretion, and flalti- Amer. I. Cfin. Nutr. 7:532-537, 1959. lent dyspepsia. Lancet, 1:41 3-415, 1967. 9. HESSE, F. G. Incidence of chokcystitis and

87 92 other diseases among Pima Indians of southern 14. MILLER, M., T. A. BURCH, P. H. BENNETT, Arizona. IRMA 170:1789-1790, 1959. and A. G. STEINBERG. Prevalence of diabetes mellitus 10. IMAMOGLU, K., S. L. WANGENSTEEN, H. D. in American Indians. Results of glucose tolerance Roo; P. A. SALMON, W, 0. GROFEN, and 0. H. tests. in Pima Indians of Arizona. Diabetes 14:439, WANGENSTEEN, Production of gallstones bypro- 1965. longed administration of progesterone and estradiol 15. SIEVERS, M. L. and in rabbits. Surg.Forum 10:246-249, 1959. J.R. MARQUIS. South- 11. KAMEDA, H. Gallstone disease in Japan. A western American Indians' burden. Biliary disease. report of 812 cases. Gastroenterology 46:109-114, MMA 182:570-572, 1962. 1964. 16. VAN DER LINDEN, W. Some biological traits 12. LAM, R. C. Gallbladde diseasesamong Ameri- in female gallstone disease patients. A study of body can Indians. /. Lancet 74:305-309, 1954. build, parity and serum cholesterol level witha 13. LIEBER, M. M. Incidence of gallstones and discussion of some problems of selection in observa- their correlation with other disease. Ann. Stag. tional hospital data. Acta Chit. Scand. 269, Supple- 135:394-405, 1952. ment 1, 1961. HYPERGLYCEMIA IN PIMA INDIANS: A PRELIMINARY APPRAISAL OF ITS SIGNIFICANCE

Max Miller, Peter H. Bennett, and Thomas A. Burch'

There have been several reports in recentlation group, geographically isolated from the years indicating that certain groups of North other. Some 3,300 Indians aged 5 years and over American Indians have a high prevalence of reside in this community; of these 3,041 claim to diabetes mellitus (5, 14). It has also been sug- be half- to full-blooded Pima. gested thatthe frequency and typesof the Between 1 March 1965 and 1967, 1881 of these associated clinical manifestations and complica- Pima Indians received a modified glucose toler- tions differ in the American Indian from thoseance test at the National Institutes ofHealth observed in the Caucasian and the Negro (12). Clinic at the Sacaton PHS Indian Hospital. An This report will examine the frequency of hyper-oral 75-gram glucose equivalent carbohydrate glycemia andrelatedvariablesinaNorth load 2 (6) was given to each subject who at- American Indian tribe, the Pima Indians of tended the examination center regardless of the Arizona. time of the previous meal. Two hours later venous blood was drawn into a tube containing Methods sodium fluorideand another specimen was Thc Pima Indians have resided for at least placed in a plain glass tube. Glucose concentra- several centuries in the region of the Gila River tion was measured on the AutoAnalyzer, using in south-antral Arizona, which is typical of the the modified Hoffman method,8 in plasma ob- hot, dry Sonoran desert of the southwesterntained from the fluoride tube. Serum creatinine United States. The Pima were traditionally an was determined on theadditional serum sample. agricultural group, and they 3r their forebears, Before drinking the carbohydrate load each with the aid of irrigation water from the Gila subject was asked to void, and two hours later a River, have farmeditsvalleyfor about twourine sample was collected, which was tested thousand years (4). The majority of them now for protein and glucose with dipsticks.4 When- reside on the Gila River Indian Reservation, toever protein wa F.found in a concentration of the south and east of Phoenix, Arizona. approximately 30mg/100m1 or more ( >1+ ), The inhabitants of the reservation reside inquantitative determination of the urinary pro- two main regions, one of which receivesits tein concentration in a sample of the urine was medicalcare from the U.S.PublicHealth made together with determination of the urinary Service Indian Hospital in Phoenix and the other from the PHS Indian Hospital in Sacaton, 2 Glucola,Ames Company,Elkhart,Indiana,or latterregion containsseveral Dexcola, Custom Laboratories, Baltimore, Maryland. Arizona. The Technicon AutoAnalyzer Method File N-20, Tech- villages that form an independent natural popu- nicon Instruments Corporation, Chauncey, New York, 1965. I Presented by Dr. Miller. 4 Labstix, Ames Company, Elkhart, Indiana.

89 94 creatinine concentration. Among those aged 15and their outcome among the Pimawomen years and over a subsample was e,;amined moreaged 25 to 44 years on 1 January 1966 (/). thoroughly and retinal photographs, electrocar- Autopsy data were collected on members of diograms, and radiographs of the soft tissues of the Pima Indian population who have died since the left calf (lateral view) and thigh (AP view) our initial survey in 1963, and death certificates were taken!' have been collected systematically since 1964. A prospective study of the outcome of the pregnancies among the Pima women living in Results the same community was initiated in October Table 1 shows the number of Pima subjects 1965. The modified glucose tolerance testwas in the sample with hyperglycemia byage and performed on these womenone to three days sex and the number with previously diagnosed after parturition, and data on thecourse ofdiabetes who have receivedspecific hypogly- pregnancy and the characteristics of the off-cemic therapy in the course of routine medical spring were collected. In addition,a retrospec- care. The prevalence of previously treated dia- tive study was made of all previous pregnancies betes was highest in the males aged 65to 74 (26.2 per cent) and in the 55-to-64-year-old 5 The subsample consisted initially of a one-in-two females (40.8 per cent). sample of those aged 30 years and over but wasex- tended, after examination of the first 1,100 persons, to If "diabetes" is arbitrarily definedas a venous includeallthose examined aged 15 years and over. plasma glucose level of 160mg/10Orn1 ormore Though this is not a truly random sample,we believe that itis unlikely to be significantly biased in relation two hours after a 75-gram glucose equivalent to any findings associated with plasma glucose levels. load, the majority ofcases of the disease were

TABLE 1.Carbohydrate intolerance among Pima (4/8-8/8) Indians of the Gila River Reservationaged 5 years and over, 1965-1967

NO. WITH NO. WITH TWO-HOUR AGE GROUP NO. KNOWN PREVIOUSLY GLUCOSE LEVELS NO. WITH % WITH BY SEX EXAMINED TREATED DIABETES >160 MG "DIABETES'' "DIABETES'.

Males 5-14 238 0 0 0.0 15-24 146 0 4 2.7 25-34 86 0 15 15 17.4 35-44 91 10 20 30 33.0 45-54 70 13 17 30 42.9 55-64 74 16 15 31 41.9 65-74 61 16 12 28 45.9 75 and over 38 4 12 16 42.1 TOTAL 804 59 95 154 19.2

Females 5-14 284 0 7 7 2.5 15-24 230 1 11 12 5.2 25-34 149 9 21 30 20.1 35-44 150 23 50 73 48.7 45-54 93 25 34 59 63.4 55-64 93 38 26 64 68.8 65-74 55 16 15 31 56.4 75 and over 23 3 9 12 52.2 TOTAL 1077 115 173 288 26.7

90 undiagnosed at the time of the modified glucose CUMULATIVE PERCENTAGE DISTRISUTION OF TWO HOUR POST LOAD VENOUS GLUCOSE LEVELS IN PIMA INDIANS tolerance test. The combined prevalence ofpre- viously and newly diagnosed "diabett:- reached a maximum of 45.9 per cent in the males aged 65 to 74 years and 68.8 per cent in the females aged 55 to 64. In all age groups, especially those of 35 years andover, the prevalence of "dia- betes" in the females exceeded therate in the

males, the differences being significant in each *ISLES decade between 35 and 64years. Thus, both male and female Pima Indians hadan extra- ordinary frequency of "diabetes"higher than -worammaw....voinzammimi 100 SOO SOO 800 500 800 700 that reported in any other population studied. PLASMA SLUG Oltittel00 Table 2 and Figure 1 show the distribution of FIGURE I.Age- and sex-specific:umblatiyc per- the two-hour plasma glucose resultson all Pima centagedistributionsoftwo-hoar Indians examined regardless of whether they plasma glucose kvds in Pima Indians had previously diagnosed diabetes, provided they did not have concurrently treated diabetesi.e.,values had a bimodal distribution. Usingan had not taken specific hypoglycemic medication iterative maximum likelihood procedure, these within the period in which the two-hour glucose distributions were dissected mathematically and determination could have been directly influ-each of the observed distributions in those aged enced.° Such a determinationwas made on 1,776 25 to 74 yearsfor example, that shown in of the subjects. The age- and sex-specific cumula- Figure 2 for males aged 55 to 74proved to be tive frequency distributions of the plasma glu- a statistically satisfactory fit for two overlapping cose values indicated that there was a relatively log10 normal (Gaussian) distributioncurves. In steep rise in the cumulative frequency distribu- those below 25 years, however, onlya unimodal tion in the lower range of the glucose valLes in distribution was found, as is shown in Figure 3 for males aged 15 to 24, which all age groups. There was not, however,a uni- was a satisfactory form increase in each successiveage group at fit for a single logio normal distribution (13). all percentile levels. Below the median (fiftiethHence theheterogeneityinferredfromthe percentile level) in both sexes the increaseswere cumulative percentage distribution in the older relatively small, but the upper percentile glucose groups appeared to result from the presence of values were much higher in the olderage groups of both sexes than in theyounger subjects. This 20 MALES 55-74 suggests that, as age increased, the proportion of subjects with hyperglycemia increased mark- edly. From this the inferencewas drawn that the plasma glucose levels in all except theyoungest groups of each sex were not homogeneous but consisted of two or more components. Further analysis indicated that in eachsex and in all 5 decades from 25 to '74 years the frequency distri- bution of the logarithms of the two-hour glucose SO ibo 200 ° For patients taking tolbutamide or phenformin this tASMA GLUCOSI 1.1/100 period was 24 hours;forpatientstaking insulinor FIGURE 2. Frequency distribution of two-hour post- chlorpropamide or other oral agents or combinations of load venous plasma glucose levels (logarithmic scale) in oral agents, 48 hours. Pima Indian males aged 55-74 years

91 66 TABLE 2. Cumulative percentage of two-hour post-load vcnotr, plasma glucose levels (mg/100m1) in Pima Indians AALES FEMALES GLUCOSEPLASMALEVEL NO. 5-14 YEARS CUM. % NO. 15-34 rEmts CUM. % NO.35-54 YEARS CUM. % NO. 55 AND OVER CUM. % NO. 5-14 YEARS CUM. % No. 15-34 YEARS CUM. % NO. 35-54 YEARS CUM. % NO. 55 AND OVER CUM. % 0-50 1 0.4 2 0.9 1 0.7 1 -126-100 -63-80 878636 6 90.854.218.1 2.9 707633 6 80.650.417.7 3.4 432611 2 55.026.5 2.09.3 -3418 6 37.415.5 3.9-- 106 9831 5 84.547.212.7 1.8 138 9429 8 72.135.1 9.92.1 -5912 9 37.6 9.94.2 - 19 42 20.0 0.85.42.3 -159-199 22 100.0 26 35 91.894.0 1118 6 78.174.266.9 401118 74.863.2 37 7 100.097.5 6818 9 97.695290.3 363117 77.069.052.1 381511 69.257.749.2 -398-316-251 - 32 97.496.195.3 18 4 92.780.8 95 91.085.281.9 1 97.9 1112 87.882.6 16,11 90.077.7 -501 - - 3 98.7 6 96.7 11 1 98.1 - - 5 99,2 12 93.4 6 94.6 -796-631 - - 21 100.0 99.6 4 99.3 2 100.0 98.7 - -- - 3 100,0 11 1 99.198.6 34 100.0 97.7 -1260-1000 238 -- - 232 151 - 1 100.0 - 155 284 - - 373 - - 213 2 100.0 130 TotalWith concurrently treated diabetes 238 - 0 - 232 - 0 - 161 10 173 -18 - 284 -- - 379 - 6 - 243 30 171 41 - Among those with diabetes concurrently under treatment, retinopathy was observed in 20.0 per cent of the males and in 19.6 per cent of the females. Proteinuria Other evidence that hyperglycemia among the Pima Indians was associated with vascular com-

plications was investigat 1 by examination for evidence of renal disease. Table 4 shows the relationship of proteinuria ( >1 +) to the two- hour post-load plasma glucose level. The fre-

Pt ASMA G UC OSIIng/190PRI/ quency of proteinuria among both males and FIGURE 3. Frequency distribution of two-hour post- females increased with age and plasma glucose load venous plasma glucose levels (logarithmic scale) in level. Within each age group there wasa gradi- Pima Indian males aged 15-24 years ent of increasing frequency of proteinuria with increasing severity of hyperglycemia. This rela- two components, one characterized by a much tionship was similar in males and females, as higher mean plasma glucose level than the other. was the over-all frequency of proteinuria in both The parameters of the distributionswere such sexes and in every age group of each sex. that bimodality was apparent only because the In 105 of the 128 subjects with proteinuria it proportion of subjects who contributed to thewas possible to make more quantitative esti- higher curve was relatively high. mates of proteinuria by determining albumin In order to determine whether the subjects inand creatinine concentrations in the two-hour the two distributions consisted of those with and urine samples.Sincethedaily excretionof without diabetes mellitus in the clinicalsense, creatinine in the urineisrelatively constant the relationshipof other parameters ofthis (H), the albumin/creatinine (A/C) ratio will disease to glucose tolerance levelswas examined. yield an approximate estimate of the 24-hour urinary excretion of protein. Thus, an A/C ratio Diabetic retinopathy of 1 or more indicates that one or more grams of "Diabetic retinopathy" is allegedly one of the protein are excreted daily. most specific signs of diabetes mellitus. Its fre- Markedly elevated A/C ratios ( >1.0) were quency was assessed in the Pima Indians by the found more frequently in those who had two- evaluation of retinal photographs for thepres- hour post-load glucose levels in excess of 200mg/ ence of microaneurisms, preretinal and vitreous 100m1 and in the concurrently treated diabetics hemorrhages, exudates, and neovascularization.than in subjects with lower values (Table 5). Table 3 shows the frequency of diabetic retinop-Thus, among those with high glucose values, athy in all subjects aged 15 years andover whoproteinuria was both more frequent in any de- had one or more satisfactoryretinal photo- gree and also much more severe. graphs. The nephrotic syndrome associated with se- No definite evidence of diabetic retinopathy vere intercapillary glomerulosclerosis isrecog- was observed in male or female Pima Indians nized as one of the important complications of under 35. In those above thisage, it was found the renal disease of diabetes mellitus. The essen- only in those with the h:gher plasma glucosetial feature of this syndrome is the occurrence levels. In those with two-hour plasma glucose of excessiveproteinuria, usually exceeding 3 levels of 200mg/100m1 andover, retinopathy grams a day. It is noteworthy that of those with was approximately twice as frequent as in thoseplasma glucosevalues of 200mg/100m1 and withlevels between 160 and 199mg/l00m1.over, 11 of the 39 with proteinuria on whom the

93 , TABLE 3.Diabetic retinopathy and two-hour post-load plasma glucose levels in Pima (4/8-8/8) Indians examined 1965-1967

PLASMA GLUCOSE LEVEL (MG/lOOML) AGE GROUP WITH CONCURRENTLY BY SEX 0-100 101-159 160-199 200+ TREATED DIABETES TOTAL

Males 15-34 years No. examined 43 43 0 8 1 95 No. affected 0 0 0 0 0 0 Per cent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 years No. examined 26 35 9 27 9 106 No. affected 0 0 0 4 2 6 Per cent 0.0 0.0 0.0 14.8 22.2 5.7 55 and over No. examined 15 41 11 19 10 96 No. affected 0 0 1 3 2* 6 Per cer t 0.0 0.0 9.1 15.8 20.0 6.3 Total 297 No. examined t-; 4 119 20 54 20 No.affected 0 0 1 7 4 12 Per cent 0.0 0.0 5.0 12.9 20.0 4.0

Females 15-34 years No. examined 43 71 6 12 3 135 No. affected 0 0 0 0 0 0 Per cent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 No. examined 13 67 27 45 23 175 No. affected 0 0 0 4 3 7 Per cent 0.0 0.0 0.0 8.9 13.0 4.0 55 and over No. examined 2 29 6 30 20 87 No. affected 0 0 1 6 6 13 Per cent 0.0 0.0 16.7 20.0 30.0 14.9 Total No. examined 58 167 39 87 46 397 No. affected 0 0 1 10 9 20 Per Lent 0.0 0.0 2.6 11.5 19.6 5.3

a Includes one male who at the time of the retinal photographs had a plasma glucose of 156 mg/100 ml while not on medication. He had, however, lost weight and had carcinoma stomach with metastases,and in1961) had a fasting blood glucose of 165 mg/100 z

4.1C 9 TABLE 4.Proteinuria and two-hour post-load plasmaglucoselevelsin Pima(4/8-8/8)Indians examined 1965-1967 (> 1+ on dipsticks4)

PLASMA GLUCOSE LEVEL (M0/100ML) AGE GROUP WITHCONCURRENTLY BY SEX 0-100 101-159 160-199 200+ TREATED DIABETES TOTAL

Males 15-34 years No. examined 108 90 5 13 0 216 No. affected 6 5 0 1 0 12 Per cent 5.6 5.6 0.0 7.7 0.0 5.6 35-54 years No. examined 39 59 10 36 10 154 No. affected 2 6 2 4 4 18 Per cent 5.1 10.2 20.0 11.1 40.0 11.7 55 and over No. examined 22 70 17 35 17 161 No. affected 2 6 3 10 6 27 Per cent 9.1 8.6 17.6 28.6 35.3 16.8 Total No. examined 169 219 32 84 27 531 No. affected 10 17 5 15 10 57 Pcr cent 5.9 7.8 15.6 17.9 37.0 10.7

Females 15-34 years No. examined Irv/ 176 16 17 6 324 No. affected 4 7 2 5 0 18 Per ccnt 3.7 4.0 12.5 29.4 0.0 5.6 35-54 years No. examined 20 86 36 64 29 235 No. affected 0 4 2 10 5 21 Per cent 0.0 4.7 5.6 15.7 17.2 8.9 55 and over No. examined 8 51 10 44 40 153 No. affected 2 2 0 15 13 32 Per cent 25.0 3.9 0.0 31.8 32.5 20.3 Total No. examined 137 313 62 125 75 712 No. affected 6 13 4 30 18 71 Per ccnt 4.4 4.2 6.5 24.0 24.0 10.0

fuo TABLE 5.Albumin/creatinine ratios in subjects with > 1+ proteinuria

PLASMA GLUCOSE LEVEL (MG/lOOML) WITH CONCURRENTLY 0-100 101-159 160-199 200+ TREATED DIABETES TOTAL

Males No. examined 5 17 4 13 9 48 A/C> 1.0 0 2 2 9 7 20 Per cent 0.0 11.7 50.0 692 77.8 41.7 Females No. examined 5 10 2 26 14 57 A/C> 1.0 2 2 0 14 11 29 Per cent 40.0 20.0 0.0 53.8 78.6 50.9 Total No. examined 10 27 6 39 23 105 A/C> 1.0 2 4 2 23 18 49 Per cent 20.0 14.8 33.3 58.9 78.3 46.7

A/C ratios were determined had values of 3.0 previously had several blood glucose values in and over, as did 11 of the 23 with concurrentlyexcess of 250mg/100m1 and had received insulin treated diabetes. A/C ratios of this magnitude and tolbutamide for many years. Prior to testing were not encountered in subjects with two-hour he had lost at least 26 pounds in weight, and he glucose levels of less than 200mg/100m1 except died 23 days after examination. in one 32-year-old male who was known to have Thus, evidence was found that hyperglycemia had acute glomulonephritisnineyearspre- among the Pima Indians was strongly associated viously. with proteinuria and impairment of renal func- tion, especially among the older subjects of both Serum creatinine levels sexes. Inordertodeterminethefrequ,:ncyof Vascular calcification subjects with significantreductionsinrenal function, serum creatinine levels were examined. Evidence of disease of the peripheral vascula- Elevated levels were defined for this :....pose as ture was evaluated in soft-tissue radiographs of 15mg/100m1 and over in males, and 1.3mg/the thigh (femoral artery) and the calf (tibial 100m1 and over in females. Such levels were arteries), since vascular calcification has been found mainly among subjects with two-hourshown to be more frequent among diabetics plasma glucose levels of 200mg/100m1 and over than among nondiabetics (3) and to develop and among those with concurrently treated dia-concomitantly with increasing duration of dia- betes (Table 6). They occurred predominantlybetes among the younger subjects with diabetes among the older subjects in the same groups in (1 5).Tables 7 and 8 show the frequency of which marked proteinuria was found. calcification in these vessels. In both sites calci- The records of four male subjects with plasma fication was found more frequently in the males glucose levels of 101 to 200mg/100m1 who had than in the females, was age-related, and was elevated serum creatinine levels were examinedencountered more frequently in the tibial arte- to determine possible reasons for these. Two had ries than in the femoral artery. The calcification histories of hypertension and congestive heartobserved was mainly medial in type; the intimal failure, and another had received renal dialysis type was rarely seen and never occurred alone. following trauma and shock. The fourth, with Calcification in the femoral artery and in the a glucose value of 160-199mg/100m1, was a tibial arteries was not found in any Pima Indians

known diabetic t.f many years' duration. He had below the age of 35 years. In the older groups WI TABLE 6.Elevated serum creatinine and two-hourpost-loadplasmaglucoselevelsinPima(4/8-8/8) Indians examined 1965-1967 (Males > 13 mg/100 ml : Females1.3 mg/100 ml)

PLASMA GLUCOSE LEVEL (MG/100ML) AGE GROUP WITH CONCURRENTLY BY SF.X 0-100 101-159 160-199 200+ TREATED DIABETES TOTAL

Males 15-34 years No. examined 84 61 2 8 0 155 No. affected 0 0 0 0 0 0 Per cent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 years No. examined 27 49 9 30 10 125 No. affected 0 2* 0 0 2 4 Per cent 0.0 4.1 0.0 0.0 20.0 3.2 35 r nd over No. examined 19 64 17 26 16 142

No. affected 0 1* 1' 2 1 5 Per cent 0.0 1.6 5.9 7.7 63 3.5 Total No. examined 130 174 28 64 26 422 No. affected 0 3 1 2 3 9 Per cent 0.0 1.7 3.6 3.1 11.5 2.1

Females 15-34 years No. examined 62 84 6 11 2 165

No. affected 0 0 0 1 0 1 Per cent 0.0 0.0 0.0 9.1 0.0 0.6 35-54 years No. examined 14 67 26 52 26 185 No. affected 0 0 0 0 2 2 Per cent 0.0 0.0 0,0 0.0 7.7 1.1 55 and over No. examined 6 48 6 41 34 135 No. affected 0 0 0 5 4 9 Per cent 0.0 0.0 0.0 12.2 11.8 6.7 Total No, examined 82 199 38 104 62 485 No. affected 0 0 0 6 6 12 Per cent 0.0 0.0 0.0 5,8 9.7 2.5 4 See text.

97. 102 TABLE 7.Medial calcification of the femoral artery and two-hour post-load plasma glucose levelsin Pima (4/8-8/8) Indians examined 1965-1967

PLASMA GLUCOSE LEVEL (MG/IOOML) AGE GROUP WITH CONCURRENTLY BY SEX 0-100 101-159 160-199 200+ TREATED DIABETES TOTAL

Males 15-34 years No. examined 59 49 3 7 0 118 No. affected 0 0 0 0 0 0 Per cent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 years No. examined 23 38 5 28 9 103 4 No. affected 0 0 1 2 1 Per cent 0.0 0.0 20.0 7.1 11.1 3.9 55 and over No. examined 17 59 14 26 15 131 No. affected 5 17 5 7 8 42 Per cent 29.4 28.8 35.7 26.9 53.3 32.1 Total No. examined 99 146 22 61 24 352 No. affected 5 17 6 9 9 46 Per cent 5.1 11.6 27.3 14.8 37.5 13.1

Females 15-34 years No. examined 39 68 10 11 3 131 No. affected 0 0 0 0 0 0 Per cent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 years No. examined 12 54 24 49 22 161 No. affected 0 0 0 2 0 2 Per cent 0.0 0.0 0.0 4.1 0.0 1.2 55 and over No. examined 5 42 7 36 34 124 No. affected 0 2 0 7 6 15 Per cent 0.0 4.8 0.0 19.4 17.6 12.1 Total No. examined 56 164 41 96 59 416 No. affected 0 2 0 9 6 17 Per cent 0.0 1.2 0.0 9.4 10.2 4.1

1C3 TABLE 8.Medialcakification in thetibialarteriesand two-hour post-load plasma glucose levels in Pima (4/8-8/8) Indians examined 1965-1967

PLASMA GLUCOSE LEVEL (MG/100ML) AGE GROUP WITH CONCURRENTLY BY SEX 0-100 101-159 160-199 200+ TREATED DIABETES TOTAL

Males 15-34 years No. examined 58 47 3 7 0 115 No. affected 0 0 0 0 0 0 Per cent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 years No. examined 21 38 5 28 9 101 No. affected 0 2 1 6 2 11 Per cent 0.0 5.3 20.0 21.4 22.2 10.9 55 and over No. examined 16 56 14 26 14 126 No. affected 4 24 5 11 8 52 Per cent 25.0 42.9 35.7 42.3 57.1 41.3 Total No. examined 95 141 22 61 23 342 No. affected 4 26 6 17 10 63 Per cent 4.2 18.4 27.3 27.9 43.5 18.4

Females 15-34 years No. examined 45 68 10 11 3 137 No. affected 0 0 0 0 0 0 Per cent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 years No. examined 12 58 21 49 23 163 No. affected 0 0 3 2 0 2 Per cent 0.0 0.0 0.0 4.1 0.0 1.2 55 and over No. examined 5 42 7 34 32 120 No. affected 0 3 2 10 7 22 Per cent 0.0 7.1 28.6 29.4 21.9 18.3 Total No. examined 62 168 38 94 58 420 No. affected 0 3 2 12 7 74 Per cent 0.0 1.8 5.3 12.8 12.1 5.7

fb4 both sites wete affected more frequently, but not 1,253 pregnancies were considered "diabetic," exclusively, in subjects with the higher two-hour as against a frequency in the Cleveland, Ohio, plasma glucose values and with concurrently metropolitan area of 0.28 per cent (8). The fre- treated diabetes. In the majority of subjects withquency of "diabetic" pregnancy was therefore femoral artery involvement, the tibial arteriessome 10 to 15 times greater among the Pima were also affected. Indians than among the women of Cleveland. The frequency of offspring weighing nine Hyperglycemia and pregnancy pounds or more at birth was significantly higher Diabetes mellitus is well recognized as having in the Pima "diabetic" group than in the "nor- characteristic effects on pregnancy and its out- moglycemic" mothers. The frequency of con- come. In order to determine whether similargenital anomalies and the perinatal mortality effects were found in the Pima Indians a review rate were also significantly higher among the was made of the past pregnancies of females"diabetic" group. The perinatal mortality rate aged 25 to 44 years on 1 January 1966 who bad in the latter was 25.5 per cent, which is in the received a moditied glucose tolerance test and same range as that reported in other populations had borne at least one child. On the basis of(10). this recent test and clinical history, the females A prospective study of the current pregnancies were divided into three groups: those with two- of Pima women was started in 1965. The modi- hour post-load venous plasma glucose levels of fied glucose tolerance test was performed on the less than 140mg/100m1, those with either clini- cal diabetes requiring hypoglycemic treatment mother generally two to three days postpartum, or plasma glucose values of 160mg/100m1 orbut irt a few instances it was done on the first over, and those who had intermediate plasma day. Table 10 shows the relationship of the glucose levels. The last group were considered infant birth weight to the two-hour post-load to be in a "gray zone" and were not analyzed. plasma venous glucose level among such women. Pregnancies among the second group were di-While the number with high postpartum glu- vided into those that occurred before the clinical cose values was small, there was a highly sig- diagnosis of diabetes ("prediabetic") and those nificant relationship between birth weight and that occurred after the diagnosis ("diabetic"). maternal postpartum glucose level, an association The results appear in Table 9. Among thecor,sistent with that noted in the retrospective Pima females studied, 47 (3.8 per cent) of the study.

TABLE 9.Outcome of pregnancy among 215 Pima females (retrospective study)

OUTCOME 'NORMOGLYCEMIe' '.PREDIABETIC" "DIABETIC" TOTAL

Birth weight No. of offspring 571 229 38 838 >9 pounds at birth 37 26 15 78 Per cent 6.5 11.4 42.9 9.3 Zongenital anomalies No. of offspring 785 375 47 1207 With congenital anomalies 31 13 9 53 Per cent 3.9 3.5 19.1 4.4 Perinatal mortalily No. of births 815 391 47 1253 No. of deaths 10 12 12 34 Per cent 1.2 3.1 25.5 2.7

" Includes stillbirths and neonatal deaths occurring in first 28 days of life.

100 JAZ TABLE 10. Infant birth weight and maternalpost diagnosis was sometimes basedupon less-than- partum two-hour post-load plasma glucoselevelsin ideal criteria. The rate Pima (4/8-8/8) Indians examined Oct. 1965Dec. 1967 was approximately twice as high in the females and at least 40 per cent

OFFSPRING higher in the males with "diabetes." The find- WEIGHING ings indicate that mortality MATERNAL rates among Pima >4000 GRAMS PLASMA subjects with hyperglycemiawere substantially AT BIRTH GLUC6SE NO. OF higher than among those with "normal" plasma (MG/ HOMO PREGNANCIES NO. glucose levels.

0 63 17 0 0.0 Post-mortem study 64 80 49 2 4.1 81-100 90 5 5.6 Of the Pima Indians who died sinceour 101-126 76 5 6.6 initial study in 1963, autopsy datawere available 127-159 33 9 27.3 on 34 subjects with "diabetes" and on 19 with- 160-199 out. Evidence of nodular intercapillary glomu- 2}33.3 lerosclerosis was reported in six of the "dia- 200+ 3 1 betics" and in none of the others. Seven ofthe Total 274 24 8.8 34 diabetics had myocardial infarcts, compared to 2 of the 19 "nondiabetics." Tuberculosiswas Mortality feund in 7 "diabetics" and in only 1"nondia- betic." Pyelonephritis, howover, Death certificates have been collectedon the was reported as frequently among the"nondiabetics"asthe Pima Indians of the Gila River IndianReserva- "diabetics." Amputations had been performed tion since 1964, and autopsy datawere obtained on persons who have died since 1963. About half on 2, and mucormycosis was present in1 of those with "diabetes." of those who died receivedpost mortem exami- nations. Over a four-year period therewere 53 Discussion male and 49 female deathsamong persons aged The data plc ented show that hyperglycemia 15 years andover who had available glucose is extremely frequent among the Pima Indians. estimations or whose diabeticstatus was known, As complete glucose tolerance tests excluding those dueto accidents or violence were not per- (Table 11). formed routinely and as onlya single venous glucose determination made two hours aftera For comparative purposes the subjectswere 75-gram glucose equi lt load was available classified as "diabetic" on the basis ofa clinical on all subjects, the ust... history of diabetes and/or high glucose values. ;teria for the diagno- sis of diabetes mellitus Age-standardized death rates for subjects with cob not be applied. In order that comparisons might be made with "diabetes" in bothsexes we e found to be higher other studies, however, the frequency of subjects than those of the "nondiabetics," althoughthe who had received specific treatment for diabetes was determined, as was the frequency of subjects TABLE 11.Four-yearmortality, excluding accidental and violent death (1964-1967) in PimaIndians aged who had two-hourvenous plasma glucose levels 15 years and over of 160mg/10Gml andover. It was felt that this level, which corresponds toa whole-blood sugar MALES FEMALES of approximately 140mg/100m1,was suitable for

Totalno. deaths 53 49 comparison with several studies in whichsome- No. with diabetes 27 34 what ,irrai;ar testing methodswere used. It was Age-standardized rates/ found that known diabetes and hyperglycemia, 100 persons as so defined, occurred more frequently among Diabetics 9.8 the Pima Indians than inany other population Nondiabetics 7.8 4.4 thus far studied. Unusually high frequenciesof

101 106 diabetes have been found among the Cheiokeealso been performed on the same subjects, and Indians of North Carolina, but the prevalencethe findings will be reported separately. Post observed in the Pima Indians was even greater mortem studies, however, have already indicated (14). It was also very much greatcr than had that myocardial infarction occurred twice as fre- been found among Alaskan Eskimos (9) andquently among the "diabetics" in the autopsy was 10 to 15 times greater than the rate believed series. to be representative of the United States as a Over a four-year period the age-standardized whole (7). mortality rate amorig Pima Indian subjects on To determine whether hyperglycemia in Pimawhom glucose determination had been made, Indians was associated with the chronic vasculareither in the course of routine medical care or complications that are characteristic of diabetesfor research purposes, was shown to be greater in other populations, analyses were made toamong those of each sex with the higher glucose discover the frequency of diabetic retinopathy, levels than among those with lower levels. This renal disease, and vascular calcification among finding was consistent with the increased mor- Pima subjects aged 15 years and over as a func-tality from diabetes mellitus reported in Jos lin tion of their plasma glucose level or previouslyClinic patients, which is mainly related to the known diabetes. Retinopathy, as shown on fun- chronicvascular complications(2). Theugh dus photographs, and impaired renal function,similar processes appear to be responsible for as indicated by elevated serum creatinine levelsthe excessive Pima mortality, the number of or heavy proteinuria (A/C ratio >3.0), weredeaths is so far too small for deducing the rela- fount.. exclusively among those with two-hour tive importance of the various types of vascular glucose levels in excess of 200mg/100m1 or withcomplications. Serial studies of renal function concurrently treateddiabetes,except inrare and electrocardiograms as well as systematic individuals in whom the findings were clearly autopsy, whenever possible, will make possible explained on another basis. Proteinuria of allsuch a definition in the future. degrees was more frequent among those with The frequency of "diabetic" pregnancy among high glucose levelsor treated diabetes and,the Pima Indians was in accordance with that when present, was moderately severe (A/C ratio which might be predicted from the over-all >1.0) four times as frequently as in those with prevalence of hyperglycemia. The demonstration glucose levels below 200mg/100m1. of significantlyincreasedperinatal mortality, Since retinopathy and nephropathy are consid-high infant birth weight, and high frequency ered highly specific findings in diabetes mellitus,of congenital anomalies resulting from these their frequent occurrence in Pima Indians with pregnancies further implies that hyperglycemia elevated two-hour plasma glucose levels implies in Pima Indians has the same pathologic signifi- that the hyperglycemia in this population was a cance as diabetes mellitus among Caucasians and manifestation of the diabetic syndrome as recog-Negroes (8, 10). nized elsewhere. This conclusion was strength- As the classic symptoms of diabetes mellitus ened by the reports of cases of nodular inter-are related mainly to the individual levels of capillary glomerulosclerosis at autopsy amonghyperglycemia and are ameliorated by the spe- such subjects. cific hypoglycemic medications, the substatrial On the other hand, disease of the larger vessels, proportion of subjects who had received such as shown by the presence of vascular calcification treatment may reflect the past frequency of the in the lower limbs, occurred among older sub-characteristic symptoms ofdiabett'smellitus jects over the Whole range of glucose levels but among the Pirna Indians. While only 40 per cent was again more frequent among those of theof the subjects with glucose values of 160mg/ same age and sex with glucose levels in the upper 100m1 and over had received such therapy, they range or with concurrently treateddiabetes. included 60 per cent of the females and 50 per Standard twelve-leadelectrocardiogramshave cent of the males aged 55 to 64 years with such

102

167 levels. This observationsuggests that substantialdiffering tribes with high and low prevalences numbers of the youngerpersons with hyper-of diabetes at present could contributeto our glycemia will become symptomatic in the future.knowledge of the etiology, especially if identical Beforethesystematic study of the Pima methods of evaluation could be used in several Indian communitywas begun, diabetes mellitusof the groups. Such investigations could be was locally recognized as an important healthenhanced by parallel study ofgroups of the problem but the true frequency of hypergly-same race in Central and South America. As cemia and of the vascular manifestationswere these groups are subjected to change andas their unknown. Althoughsome other southwestern life expectation is increased, it is possiblethat Indian tribes are recognizedas having a highothers may go the sameway: in them diabetes frequency of the disease, othersare said to have mellitus may emergeas a major chronic disease a low prevalence. The significance of this varia- problem, as it is today among the Pima Indians. tion, if it were confirmed and if thereasons for the differences could be identified, would bean Acknowledgments important contribution to our understanding of the etiology of diabetes mellitus. We gratefully acknowledge the help and assistance of many persons who have contributed to this studyin A definition of the geography of the diseasein particular, Dr. E. I. Ballintine for his evaluations of the the southwestern American Indianseems to be retinal photographs, Dr, F. A. Rose for the soft-tissue needed as a matter ofsome urgency, since these radiographic readings, and Miss P. E. Sperling for the emerging groups are likely to be subjectedto majorityofthelabo-atorydeterminations. We also radical and rapid change. Somemay lose their thank the Division of Indian Health, the Tribal Council, and members of the Gila River Indian community, identity; others may adoptways widely differ- whose collaboration and cooperation made this study ent from those of their forebears. The study of possible.

REFERENCES 1. Com Ess, L. J., P. H. BENNETT, T. A. BuRcH, and 9. MOURATOFF, G. J., N. V. CARROLL, and E.M. M. MILLER. Congenital anomalies and diabetes in SCOTT. Diabetes mellitus in Eskim3s. IAMA 199: the Pima Indians of Arizona. To be published. 961-966, 1967. 2. ENTMACHER, P.S., H. F. RooT, and H. H. 10. PEDERSEN, J. Perinatal mortality in babies of MARKS. Longevity of diabetic patients in recent years. diabetic mothers. In B. S. Liebel and G. A.Wren- Diabetes 13:373-377, 1964. shall(eds.), On the Nature and Treatmento 3. FERRIER, T. M. Radio logically demonstrable Diabetes. Amsterdam, Excerpta Medica Foundation, arterial calcification in diabetes mellitus. Aust.Ann. 1965, pp. 722-726. Med. 13:222-228, 1964. 11. PETERS, J. P., and D. D. VANSLYKE, Quantita- 4. HAtmv, E. W. The Hohokam, first masters of tive Clinical Chemistry; Interpretations. Baltimore, the American desert. Nat. Geographic 131:670-701, Williams & Wilkins, 1946, p. 908. 1967. 12. RIMOIN, D. L., Genetics of diabetes mellitus. 5. JoHNsoN, J. E., and C. W. Mc Nun. Diabetes Diabetes 16:346-351, 1967. mellitus in an American Indian populationisolate. Texas Rep. Biol. Med. 22:110-125, 1964. 13. RUSHFORTH, N. B., P. H. BENNETT, A. G. STEINBERG, T. A. BURCH, and M. MILLER. Studies 6, KENT, G. T., and J. R. LEONARDs. Massscreen- ing for diabetes in a metropolitanarea using finger of plasma glucose measurements in the Pima Indians blood glucose after a carbohydrate load.Diabetes of Arizona. I. Application ofa composite of two 14:295-299, 1965. Gaussian distributions asa model for plasma glu- 7. MCDONALD, G. W., and G. F. FISHER. Diabetes cose distributions. In preparation. prevalence in the i..Tnited States. Public Health Rep. 14. STEIN, J. H., K. M. WEST, 7. M. ROBEY, D. F. 82:334-338, 1967, TIRADOR, and G. W. MCDONALD. The highpreva- 8. MILLER, M. Diabetic pregnancy and foetal lence of abnormal glucose tolerance in the Chero- survival in a large metropolitanarea. In B. S. Liebe! kee Indians of North Carolina. Arch. Intern. Med. and G. A. Wrenshall (eds.), On the Nature and 116:842-845, 1965. Treatment of Diabetes. Amsterdam, Excerpta Medica 15. WHITE, P. Naturalcourse and prognosis of Foundation, 1965, pp. 714-717. juvenile diabetes. Diabetes 5:445-450, 1956. MALARIA IN THE AMERICAN INDIAN

George G. Gig lion

Malaria prevails, or has prevailed up to re-being made in respect to the different ethnic cently, over very wide areas of the Westerngroups that might be present. Given the magni- Hemisphere, lying betv een latitudes 50 N and tude of the task, the remote, scattered. and eco- 40' S and from the shores of the Atlantic Ocean nomicallyunimportantIrdiancommunities to those of the Pacific. The American Indianwereto a great extentmissed, overlooked, or ranges from Canyda to Patagonia, and some ignored. thirteen species of American anopheles mos- The present discussion will be restricted to the quitoes have been linked with malaria transmis- Indians of Guyana, who are estimated to num- sion, seven species being vectors of major public ber 32,000, widely scattered over the least accessi- health importance in one country or another. ble parts of the country. At the present day, nine The range of ecological and epidemiological tribes are represented: the Warau of the coastal variation is therefore practically unlimited, and lowlands, mostly in the North-West District, it would be quite impossible to submit a concise, adjoining the delta of the Orinoco; the Arawak, comprehensive account of malaria in the Ameri- of the lowland forests of the near interior; the can Indian, even if adequate information on the Wapishana (Arawak stock), of the South Ru- subject were available. Unfortunately, published pununi Savannah; theCarib,formerlyvery data are conspicuously absent from the literature, numerous and widely distributed, but now re- and none of the principal malaria eradicationduced to very small numbers, mainly on the services have been able to supply information on upper Barama and Barima rivers in the North- this special malariological problem. West District; the Akawai, of the upper Berbice, During the great upsurge of antimalarialDemerara, Cuyuni, and Mazaruni rivers;the activities over the last twenty years, emphasis Patamona, of the southern portion of the Paka- was placed on vector destruction and chemo-raima Plateau;theArekunz,oftheupper therapy, not on epidemiology. Excessive hopesCuyuni and Mazaruni mountains and savan- were founded on the new insecticides with long- nahs;the Makusi,ofthe North Rupuliuni lasting residual effect, and on the powerful anti-Savannah; and the Wai-Wai, of the Essequibo malarial drugs developed during the war and in headwaters. The last five tribes are of Carib the early postwar period. Malaria eradication stock. campaigns were launched widely and waged Some of these tribes extend beyond the politi- with little or no attempt to establish base-line cal frontiers into Venezuela and Brazil. Gen- data on the local epidemiology of the disease. eralizations about malaria are not permissible, For obvious economical, political, and logisti- and my observations refer specifically to Guyana; cal reasons,effortsinthesparsely inhabited it may be noted, however, that few environ- countries of the American tropics were concen- ments pi esent greater stability and u! ;ormity trated on the more populous areas, no distinction than the equatorial rain forest, which covers

104 109 most of the Guianas and Amazonia, wherenortheastern India and in New Guinea. In the Anopheles darlingi, the most efficient malaria Western Hemisphere, however intense malaria carrier in the Western Hemisphere, is the pre-transmission may be, so far as I know it has vailing, and often the only, vector. It is in thisbeen recorded only among the Bush Negroes of vast region that the majority of American In-Surinamdescendants of runaway slaves who dians live still fully exposed to the ravages ofhave lived, for many generations, isolated in uncontrolled and untreated malaria. the Guyana forest, where other races, equally The study of racial differences in .r..spect toexposed, suffer severely and never acquire the the epidemiology of malaria and the clinical and high degree of adult tolerance to malaria infec- immunological reactions induced by the malaria tion that is characteristic of holoendemic ma- parasitespresents many difficulties; for validlaria. This suggests that race may be a factor conclusions studies must be based on observa-connected with holoendemicity. tions made under identical conditions of en- Among the Bush Negroes, Swellengrebel and vironment and exposure. Such situations areVan der Kuyp (12) found a very high P. falci- rare, and this is probably the reason why studies parurn ratein both children andidults;the on this aspect of malariology are few and the mortality at all ag t. was low, howeve-, and they conclusionsdrawn somewhattentativeandwere not much incommoded by the infection. guarded. Thus, in a village on the upper Surinam River, The Negroes of West and Central Africa have P. falciparum accounted for 75 per cent of the evolved, throughout the ages, in continued con-infections in children, and for 73 per cent in frontation with malaria; they have acquiredadults. The actual parasite rate averaged 70.5 practically total immunity to Plasmodium vivaxper cent in children and 37 per cent in adults, and a high degree of tolerance to P. falciparum. yet during the twelve-year period from 1927 to Tolerance to the latter species implies no im- 1939, the population of the village increased by munity to the parasite, but a high degree of30 pet- cent, from 698 to 901, with an average tolerance to the effects of parasitization. Rai-arcual death rate of 21, a birth rate of 39 per chenow (10),in1929, working among the10J0 inhabitants, and infant mortality of113 Hausa of the Cameroons, reached the conclusion per1000 birth !Creole Negroes fromthe that toxin resistance was inherited but parasite Surinam coastland, living in similar surround- resistance was acquired by continued exposureings, showed the same or even higher infection and reached its peak in early adulthood; there- rates andiff,-.red severely. after, the parasite rate and the degree of para- The Amerindians of Surinarn, living under sitization remained low and more or less stable, the same conditions asthe Bush Negroes, Others in Africa have reached similar conclu-showed little evidence of tolerance to the infec- sions. tion. On the Wayombo River, in September Holoendemicmalariais regardedasthe 1939, 172 Indians were examined; P. falciparum highest expression of malaria endemicity: the accounted for 50 per cent of the infections in spleen rate in children 2 to 12 years old is con-children (106 examined) and 61 per cent in stantly above 75 per cent; the spleen rate and the adults (64 examined), and the spleen rates were size of the average enlarged spleen fall progres-82 and 83 per cent respectively. Swellengrebel sively with age, becoming low or negligible in and Van der Kuyp found that the adult population. Mortality may be high in The adult spleen rate and the average size of thc en- young children,buttheadults enjoy good larged adult spleen(2.2 Schaffnerunits), surpassing anything we encountered in Bush Negroes, arc evidence health so far as malariaisconcerned. Ho lo- of the lack of tolerance of these people to malaria para- endemic malaria is typical of Africa but has also sites. We may add that a high spleen rate and a large been recordcd in other rather restricted situa- size tyf the swollen spleen arc peculiar of Indian malaria, as distinct from the push Negro and Creole malaria one tions, as, for instance, among the aboriginals of meets everywherein Surinam. . . Besidesa strong

105

'140 spknic reaction to the presence of malaria parasites in considerable size were found rarely in the older their blood, the Indians give evidence of their lack of children and even less frequently in adults. tolerance by sickness and death. . .The importance of Indians as carriers of malarialinfection does not, Among East Indians, on the contrary, the like the Bush Negroes, consist in a tolerance to the incidence and degree of splenic enlargement was parasite (manifesting itself by a moderate splenic reac- much higher and was hardly affected by age. tion and a low morbidity and mortality), but ir their seminomadic life which enables them to carry rr dignant Clinicalexperienceinsugar-estatehospitals malaria infection from the Bush to the Savannan, showed that the majority of adults went through American Negroes,descendantsofslaves life afflicted by largeoften very largespleens brought from Africa between the sixteenth andand failed to come to terms with the infection; the nineteenth centuries, who have lived forthey suffered frequent attacks of fever, and generations under conditions of more or lessanemia of some degree was practicallyuni- moderate malaria endemicity, still enjoy someversal. Mortality directly due to malaria was of the immunity built up by their ancestors. The highest in children, but the tolerance established North American Negro presents considerable in adults by life-long exposure to infection was resistance to provoked infection with P. vivax both low and unstable; at all ages, malaria and (1), and West Indian Negroes throughout theits sequelae continued to be major causes of Caribbean, though equally susceptible to infec- death. tion, on the average suffer much less and present The 239,000 immigrants who came to Guyana lower spleen rates and smaller average enlarged from India between 1838 and 1917 originated spleens than other raceslivingin the samemainly in Bengal, I3ibar, Orissa, the Central environment (2). Provinces, and Madras, where malaria is mostly, In Guyana, people of sixraces have been though not exclusively, hypoendemic or meso- living for many generations in the same general endemic, and seasonal inits occurrence. This environment and often under the same condi- explains the lack of any inherited resistance to tions of exposure to malaria infection.' The malaria and the severe reaction these immigrants Europeans (mainly of Portuguese descent) and suffered when they were transferred to the the Chinese constitute small minorities and live hyperendemic Guyana coastland. mostly in Georgetown. The rural population of Further inland, along the tidal course of the thecoastlandscompriseslargenumbersof Negroes and East Indians, and ideal conditions Berbice, Demerara, and other rivers, the autoch- exist for comparative racial studies of all kinds. thonous population is formed by Amerindians Up to 1946, when large-scale DDT sprayingof the Aiawak and Akawai tribes and by per- sons of mixed race, usually with Amerindian, operations were introduced, malaria used to be European, and Negro blood in very variable stable and hyperendemic throughout the coast, from the Berbice Estuary to the Pomeroon proportion. There are few permanently settled Negroes, but a considerable fluctuating popula- River (7). Systematic surveys of schoolchildren tion of this race, mainly adult males, is employed were conducted between 1937 and 1946, separate in the mining and timber industries. At the time records being kept for the two main races. The when malaria was hyperendemic in these dis- parasite rate showed no significant racial varia- tricts, the Negroes came not only from the in- tion; splenic reaction in the Negroes was, how- tensely malarial coast but also, and inhigh ever, decidedly and consistently more moderate, proportion, fromthe more mildlymalarial and decreased rapidly with age so that spleens of Leeward and Windward Islands andfrom 1 According tothelastcensus,. thepopulationof malaria-free Barbados. Whatever their origin, Guyana in 1960 totakd 560,406, comprising East Indians, their reaction to malaria infection followed the 267,840; Negroes, 183,980; Mixed. 67,189; Amerindians, same pattern: they contracted the infection soon 25,450; Chinese, 4,074; Europeans, 3.218; and Others, 8,655. after arrival, suffered an attack or two of fever,

106 and after that enjoyed normal health and rarely scattered and can be estimated at 45,000 (1967), returned for treatment. of whom approximately half are Amerindians In contrast, the mixed population born and belonging to the Wapishana, Macusi, Patamona, bred in the area suffered severely and continu-Aka ',Rai, Arakuna, and Carib tribes. On the ously,allages being affected; mortality andRupununi Savannahs and the Pakaraima Pla- morbidity were certainly graver in children, but teau,theinhabitantsare almost exclusively the tolerance acquired by the adults was poor Indians. Among them, malaria surveys have and very unstable. The Amerindians reacted in been few and irregular, and no systematic, long- the same way as the mixed population, but even term observations have been made; the same can more severely;thelargestspleensrecorded, be said of other medical studies. Prior to the reaching the pubis and partly filling the pelvis, introduction of control measures, malaria oc- were seen in adults of this race. curred throughout the year but transmission Blackwater fever was not common in Guyana, tended to be much more seasonal than on the in spite of the very high incidence of P. fda-coast, being related either to the May-August parum; it appeared at long intervals, during rains or to the floods of the larger rivers. Among periods of malaria exacerbation, and tended to the Indians of the lowland forest, spleen rates recur in some localities, with clustering of cases ranged from 40 to 60 per cent. in certain neighborhoods, homes, and families. In some of the less accessible areas, the intro- On the Demerara River between 1923 and 1931, duction of malaria appeared to be recent and 53 cases were recorded, mainly in 1926-1927, possibly related to increasedtrafficwith the with the following racial incidence (4): coastlands.Thus, the savannahs of the Ru- pt., uni, which were difficult to reach before the Incidence of Blackwater Fever advent of the airplane, were reported to be Per 1000 Per 1000 healthy up to 1930; the only Indians who suf- Race inhabitants patients treated fered were those who had visitedthe coast, Negroes 2.67 19 traveling down the Essequibo River or driving East Indians 24.00 191 Amerindians 11.79 84 cattle through the Berbice trail. By 1938 (5), Mixed races 21.83 189 the infection had spread all over the savannahs, Nephrosis and chronic nephritis associated with and more than 30 per cent of the children had P.malariae infection were frequent on the enlarged spleens, the rates increasing with age. Demerara River. The racial incidence per 1000 By 1943 (6), fears were entertained that the patients during eight years was 20.4 in Negroes; Makusi of the North Savannah were dying out. 48.8 in East Indians; 33.6 in Amerindians; and During a malariortelTAsurvey of this area, 424 35.6 in mestizos. Children and young persons Indians of all ages were examined; 68 per cent are more liable to blackwater and nephrosis had either enlarged spleens, parasites, or both, than adults; thus the low incidence of theseand 46 per cent of the adults had large spleens, syndromes in Negroes might be due, to somewhich in 20 per cent of cases extended below extent, to the preponderance of adult males in the transverse umbilical line. In the settlements this ethnic group. Ten years' experience on the situated nearest to the forest, 80 per cent of the Demerara River, in conclusion, showed that the inhabitants showed evidence of malaria. The Arawak and Akawai Indians of the arca suf- lack of old people was very striking; only two fered severely from malaria and its complications were seen, who appeared to be about 60 years and, like the East Indians of the coastlands, of age. This was reported to be a recent develop- acquired only a very relative degree of tolerancement; it was said that before malaria appeared to the infection, suffering fromitseffects at on the savannah, in1930, old and very old all ages. people were numerous. (One rancher who came In the far interior the populationis widely to the area in 1922 had known two Mahwut

107 ancients who remembered quite well the British incidence of the disease has been drastically military expedition to Pirara in 1843.) Duringreduced. The rapid increase that has followed this 1943 survey, many settlements in the Indian in all the tribesisa clearindicationofthe reservation along the northern Kanuku Moun-disastrous effect of malaria on this population tain foothills were found deserted andsome of group over the payears; in 1968, their number the houses burned, asis customary after the is estimated at 32,000. death of an inmate. This high susceptibility and reactivity of the This high susceptibility cf the Guyana Indians Amerindian to malaria is without doubt relat,c1 to malaria is without doz.i.lit one of the major to the recent introduction of the infection in the causes of their decline over the past 150 years. Western Hemisphere. Itisgenerally accepted Smallpox,measles,yellowfever,influenza, that malaria, as well as many other diseases, whooping cough, and tuberculosis have all been came to America with the discoverers, the con- contributing factors, sometimes causing disas-quistadors, and the hundreds of thousands of trous epidemics; none of these diseases, however, slaves brought from West Africa. In the Ameri- was as constant in its onslaught as malaria. casthenew arrivals found, wherever they The early Dutch colonists sought the alliancelanded, native anopheles capable of acquiring of the Indiansparticularly the Caribs, thena and transmitting the disease; given the pro- very numerous and warlike triberelying on longed infectivity of malaria subjects, the lack them forthe procurement ofIndianslaves of curative medicines at the time, and the wide captured in raids on villages and Spanish mis- diffusion of anopheles, itis not surprising that sionsettlementsonthe upper Cuyuni andserious earlyoutbreaks occurred among the Orinoco. Later, during the long years when invaders and that the infection spread rapidly African slaves were brought to the colony, both to the highly receptive Indians. the Dutch and the British planters reliedon In spite of tradition, it appears thateven the Indian help in subduing slave rebellions and in Peruvian Indians were ignorant of the properties the capture of fugitives. From 1778 to the endof cinchona bark. Richard Spruce, the botanist, of slavery, annual official receptionswere held spent fifteen years, from 1849 to 1864, on the in honor of the Indian chiefs. At the end of the Amazon, and in the cinchona forests on the eighteenth century, the Amerindian population slopes of Chimborazo in the Ecuadorean Andes appears to have been numerous and flourishing.(//). It was he whowas responsible for the The figures that have been given, however,are collection and shipment of seeds andyoung unreliable and differ widely; 50,000 would ap- plants from which the cinchona plantations of pear to be a reasonable guess. With the abolition India and Ceylon eventually originated. He of slavery, the Indians ceased to be ofany inter- found that the cascardleros (Indians employed est to the colonists, and were left to themselves in the collection of cinchona bark) not only were and neglected. Their numbers dwindled rapidly, completely ignorant of its medicinal properties and the Caribs, the most rowerful and feared but regarded his explanations with skepticism tribe, practically vanished; onlya few survive at and suspicion, for they were convinced that the thepresent day on the upper Barama and bark was used for the extraction of a valuable Barima rivers. In 1848, afterproper surveys,it brown dye. was established that the Guyana Amerindians In equatorial America, altitude, in the Andean aggregated approximately 15,000, of whom only cordillera, has probably been the only barrier to 7,000 were still in the aboriginalor semi-aborigi- the spread of malaria to even the most remote nal state (3). and isolated Indian communities. The Wai-Wai Systematicmalaria-controloperationswere Indians are an entirely primitive tribe that has inaugurated in the interiorin1947; though its main settlements in Brazil on the Mapueira eradication has notas yet been achieved, the River, which flows, through the Trombetas, to

108 the Amazon, some 400 miles to the south. A few localities where conditionsforabundant A. used to visit the headwaters of the Essequibo in darlingi production exist. Guyana. Their nearest neighbors are the Wapi- The more permanent type of house consti- shana of the South Rupununi Savannah, some tutes the Indian's operational base, but, with his 70 miles to the northwest, through high rainentire family, he spends a large part of his forest. Cassava graters and feather ornaments existenceelsewhereworking onhisfarm, have been traditional items of trade betweenusually situated at a very considerable distance; these two tribes, but contacts are maintained by on hunting or fishing expeditions; attending only a few individuals. For the last 15 years, a sprees; or just visiting his neighbors. In the more mission has been established at Kanashen, on civilized areas, he may seek temporary employ- the upper Essequibo, and it has attracted con- mentas a guide, a porter, or a boathand, bleed- siderable numbers of Wai-Wai from across theing balata (wild rubber), collecting Brazil nuts, border. In 1966 a group of 80 paid their firstand so onand when so engaged he is usually visit to the mission, after a journey of severalaccompanied by his family. When away from weeks; of these, 14 were found infected with their permanent home, the Indians camp under malaria, P. falciparurn, P. vivax, and P. rnalariaetemporary shelters, ranging from a few large all being identified. palm leaves stuck into the ground to an open So farI have dealt with the incidence of palm-thatched shed. malaria and its effects on the American Indian. This great and continuous mobility of the The Indian's way of life, however, also has Indian is without doubt a factor favoring the important bearings on malaria epidemiology and wide and rapid diffusion of malaria infection. even more so on the practical development ofIn 1947 a sharp outbreak occurred among the antimalaria operations. Patamona Indians on the Potaro-Ireng water- Indian villages are small, with widely spaced shed, on the Pakaraima Plateau, a mountainous, houses. The larger buildings usually accommo-mainly forested region at an elevation of 1,000 date several families, each having its fireplace to 2,000 feet above sea level (7). Infected persons and allotted space. Completely isolated houses were discovered scattered widely over some 500 ar.: frequent. The large villages to be found insquare miles; all of them, however, had worked the more civilized areas are due to missionaryat, or had visited recently, a diamond placer, influence. The houses are made of materials that and this was the only locality where transmis- are easily procurable on site: round wood poles, sion was actually occurring; a creek had been bush ropes, palm leaves for roof thatching, bark dammed and many gravelpits sunk in the and split palm trunks for walls and floors, andvalley bottom, and in these A. darlingi found so forth. By community effort even the larger ideal conditions for continued production. The houses can be erected in the course of a fewmigration of 80 Wai-Wai, of whom 14 were days, and a serviceable shelter may be put up malaria parasite carriers, has already been men- in a matter of hours. This makes it easy for the tioned. Indians to move from one site to another for These characteristics of the Indian way of life security,forbetter hunting orfishing, and, introduce serious difficulties in practical malaria probably, for the purpose of escaping disease. control and may frustrate even the most efficient As has been mentioned, houses in which aeradication program. In the sparsely inhabited death has taken place are frequently abandoned interior, A. darlingi, in adaptation to the en- and burned down. This instability of settlements vironment, is by no means as anthropophylic has been recorded by most observers, and it is and endophylic as it is on the coast; it is widely reasonable to expect that such a practice may distributed as a silvatic mosquito, and malaria help in the avoidance of malaria, which in the transmissiontakes place not in villages and forest is eminently a place disease, prevailing in houses but at camping sites, on the farms, and

109 114 3*.

P. vIvax. RESIDUAL FOCI 1111:* AND LINES Of SPREAD

++1-P. faklpxium. RESIDUAL FOCUS oa + Of SILVATIC IRANSMISSION

MALARIA ERADICATED 4. 90% OF POPULATION

B It A ZIL

*. 10.

FIGURE I. Map of Guyana. showing residual Plasmodium vivax foci that survived 1961-1965 chloroquinized-salt campaign; lines of spread of infection by wandering Amer- indians; small focus of P. falciparum in south, persisting among wild-rubber bleeders; and point of intrusion of primitive Wai-Wai Indians from Brazil. Elsewhere medicated salt and/or DDT house spraying have eradicated malaria or greatly reduced its incidence(see text). In south, the campaign failed, following the introduction of a chloroquine-tolerant P. ft:hi- parurn strain from Brazil in 1962. DDT house spraying on both sides of the border has interrupted transmission on the permanently inhabited Rupununi Savannahs, but a small focus of P. falciparum, transmitted by silvatic A. darlingi, persists among Amerindian balata- bleeders who camp, with. their families, in the otherwise uninhabited Rewa-Kwitaro forest, from May to November. In all, 3,500 persons are involved, with an annual average of only 24 cases. In the extreme south, the point of intrusion of primitive Wai-Wai Indians is indicated.

110 115 in the forest. The main villages can be sprayed soryinproclaimedareas and regulatingits with insecticides without excessive difficulty, but shipping, transportation, and sale in the interior. the existence of remote, isolated houses is very Epidemiologicalevaluauonpreceded,accom- frequently unknown, and the systematic spray- panied, and followed the campaign. In the areas ing of temporary shelters is not only impossible in which chloroquinized salt was used, con- but futile, since they offer next to nothing in the tinued DDT operations, from 1947 to 1960, had way of sptayable surfaces. Under such condi- greatly reduced the frequency of the disease; the tions'.1-,'cticida1 methods can reduce, but not reservoir of infection within the population was eradic nalaria, and their application is diffi- therefore relatively small. cultand expensive.Systematicsprayingof In the North-West District and in the Paka- hammocks with DDT solution or emulsion is a raima sectors, aggregating an area of 35,000 useful practice, forthe Indian always traveissquare miles and a 1964 population of 39,550, with his hammock and the insecticide residuemalaria disappeared within six months (the resists occasional washing. parasite rate in 1960 had been 8.3per cent); Mass chemotherapy, through systematic, per:- thereafter, only four sporadic, cryptic cases of odic distribution of antimalarial drugs, is obvi-P. vivax infection were recorded up to April ously impracticable, owing to the terrain, the 1965, when the campaign was closed down. widely scattered population, and its elusiveness. Early in 1966, ten months after the cessation Even when a case of malaria has actually been of all antimalaria measures in the area, a small discovered, treating it is a difficult matter; radi- focus of P.vivaxinfection wasdiscovered cal treatment of P. falciparum infection needsamong the Caribs of the upper Barama River at least three days, and for P. vivax 14 days area very inaccessible localitythat couldreceive required. Pinning down an Indian for regularchloroquinized salt only irregularly and prob- daily dosage is no easy matter, and he usually ably in insufficient quantity, as no shops or other fades into the forest in mid-treatment. normal means of salt distribution exist. These In Guyana, malaria, and its carrier A. dar- Indians, traveling down the Barama, spread the lingi, were eradicated from the coastlands and infection to the Waini and Moruca rivers and, the more accessible and populated inland areas crossing overland, caused an outbreakatthe (93 per cent of the population) exclusively by manganese mines on the upper Barima and DDT house spraying (7). In the far interior the Kaituma rivers(see arrows attop of map) results were by no means satisfactory, and nia- From there the infection spread rapidly to thy laria control only was established. This partial lower course of these streams and to the Aruka. failure of DDT was due to the wide scattering In June and July the outbreak reachedits of the population over an impervious terrain, its peak, with 372 widely scattered cases. During instability and mobility, and, above all,the 1966 a total of 643 cases were recorded on a silvatic adaptation of A. darlingi (8). total of 20,811 slides examined, out of a popula- From January 1961to December 1965 antion of 16,850; P. vivax was the only parasite attempt was made to eradicate these residual involved. The ontbreak has been controlled by foci of malariaintheinterior by means ofDDT house spraying and the reissue of chloro- chloroquinized salt (9); in order to assess fully quinized salt, these techniques being usedin the value of this new technique, DDT house combination for the first time. Only 15 cases spra}qng was suspended. This campaigr covered were recorded in 1967 (9,116 slides examined) an area of 42,000 square miles and a population and only 3 from December 1967 to J, 1968. of 48,500, mainly Amerindians. The campaign An identicalincident occurred among the was conducted under strict administrative and 4 vv.0ndians of the Wenamu and upper scientific control; specific legislation was enacted Ct\ .1 along the Venezuelan border, the first making the use of chloroquinized salt compul- cases ixing discovered in April 1966, one year 4 afterthe cessation of antimalaria operations. in the ferest during the rainyseason, from April This again is a very inaccessible area, with ato November. Among 700 bleeders and their small population of Akawai Indians whocan dependents (estimated at 2,800), 24cases were obtain salt and other supplies more easily from discovered in 1966 and 22 in 1967. Venezuela than from the Guyana side; P. vivax For the 1968 season it is proposedto issue to was responsible for all cases discovered. Indians all bleeders, along with the other rationsthey crossing overland from the Wenamu tothe receive, 20 pounds of salt medicated withchloro- Kamerang, an affluent of the Mazaruni, spread quine (0.43per cent chloroquine base) and ..he infection to the upper Mazaruni andit3 pyrimethamine (0.043per cent), and, in addi. tributaries, a sparsely inhabitedarca with antion, to give to every indi :dual whocan be estimatedpopulation of 3,000Indians and "caught" before leaving for the foresta two-day diamond miners. During 1966, 280 caseswere treatrient with a combination of Fanasil (Sul- discovered in this sector. Many infectedpersons furthomidine) and pyrimethamine. The result reached the coastlands, but were intercepted by of this new approach will not be known tillthe systematic examination of plane passengers from end of the year. the interior. DDT house spraying and chloro- These various developments in thecourse of qv inizedsalt to (issued free Indians) have the Guyana campaignillustrateclearlythe brought the situation under control, though peculiar difficulties encountered in malariacon- progresshasbeen slow, owing to the very trol and eradication operationsamong the Amer- difficult terrain. indians. As I have .said, in the early planning of Itis important to note that both these re- the many campaigns that have been undertaken crudescences were due exclusively to P. vivax; of late years throughout tropical America, little the 1961-1965 medicated-salt campaign,even in importance was attached to the specific problem remote and poorly supplied areas, appears to of malaria in the Amerindian, priority being have achieved the eradication of both P. falci. given to the larger, more accessible communities, parum and P. malariae, which before the intro- which were economicallymore important and duction of chloroquinized salt accounted for 57 politically more voca.. Today, with eradication and 15 per cent, respectively, of the infections. or, more frequently, control established over Among the 10,000 Makusi and Wapishana wide areas, the problem of malariaamong the Indians who inhabit the savannahs of the Ru- Indians assumes a new dimension, foritis pununi, close to the Brazilian border, chloro-evident that eradicationwill remain a dream quinizedsaltfailed,afterafavorablestart until means have been discoveredtoextend during 1961, owing to the introduction ofa control to eventhe most remoteofIndian chloroqu:nc-tolerant P. falciparum strain from settlements.Ihave shown how rapidly and the neighboring Rio Branco Tcf-ritory (9). This widely malaria can be broadcast from extremely new strain spread rapidly over the whole area, limited foci, by wandering or migrating Indians. even where chloroquinized-salt coverage was Of the preventive techniques known atpres- adequate. DDT housespraying,interrupted ent, medicated saltappearstobethe most since 1960, was reintroduced in 1962, andas promising, since it can findits own way far soonassystematicspraying operations werebeyond the reach of organized public health established by the Braziliansacross the border, agencies. We do not yet possess, however, the beginning in January 1965, malaria transmission ideal drug for this purpose. Those inuse today, on the savannahs was interrupted. Si !vatic A. among other shortcomings, fail to act on certain darlingi still persists in the surrounding forest, parasite strains and may thus bring about the however, and occasionalcases of P. falciparum emergence and favor the diffusionof drug- malaria continue to occur, but exclusivelyamong tolerant or drug-resistant infections, introducing Indians engaged in bleeding balata, whocamp therapeutic problems whichmay be serious.

112 417 I have shown how, in Guyana, chloroquinized fo i. the fish and game on which they subsist and ialt was completely succesul against all forms which tend to abound in certain seasons and not cf native malaria parasite, only failing where in others; the traditional method of preservation and when a chloroquine-tolerant strain was in- employed is smoking. The entirely primitive troduced from Brazil. Our campaign, it thould Wai-Wai Indians,forinstance, who reach be emphasized, was inaugurated at a stage when Kanashen Mission fromthedepthofthe the reservoir of infection had beim greatly de- Brazilian virgin forest, take to the use of salt pleted by DDT spraying. Under such conditions, with greatreadiness as soon asitis made theprobabilities of the presence of recistant available. strains were much redur- "ith the drugs at Obviously, spreading the use of salt to the present available, we benn- , at medicated saltmore remote tribes will take years, and the should not be used under conditions of high same can be said for eradicatingmalaria where endemicity. primitive Indians exist; at the present stage of Indians still in the aboriginal stage of civiliza- mala.lological knowledge and antimalaria tech- tion are not acquainted with commercial salt nique, it appears likely that treatment and pre- and manufacture salt substitutes by washing certain soils or fr3m the ash of certain palms. vention may be extended more eas;ly to the Natural deposits of salt are not likely to occur primitive and semiprimitiveIndians through throughout the vast equatorial rain forest. On theestablishedchailnels ofintertri:3albarter the other hand, civilized and semicivilized In- trade, in a bag of salt, than through a spray gun dians take to the use of commercial salt very or a bottle of tablets. For this purpose,however, readily, as in it they find not only a means for more effective drugs for salt medicationhave seasoning their food but a valuable preservative to be found.

REFERENCES I. BOYD, M. F., and W. K. STRATMAN THOMAS. 6. Gtomou, G. Malaria Survey of the North Studies on benign terian malaria. 4. On the re- Rupununi Savannah. Georgetown, 1939, pp. 1-14. fractoriness of .7.-,1cgioes to inoculation with P. vivax. (Legislat. Council Paper No. 3/1944) Amer. Hyg. 18:485-489, 1933. 7. CIGLIOLL G. Malaria,Filariasis and Yellow 2. CLARK, H. C. A comparison of the spleen and Fever in BritishGuiana:Control by Residual parasite rates as measures of malaria incidence in D.D.T.Methods. Georgetown,Medical Depart- the races of the mainland of Central America. In ment, 1948. 226 pp. Medical Department, United Fruit Company, Seven- 8. thomou, G. Fifteen years experience in malaria teenth Annual Report, 1928, pp. 77-83. eradication in British Guiana. Maintenance policies 3. COMMISSIONER OF THE INTERIOR. (1948) The and residual problems. Riv. Parassit. xx(4) :280-288, Amerindianproblem.InDevelopmentReport. 1959. Georgetown, Government of British Guiana, 1948, 9. GIGLIOLL G., F. J. RUTTEN, and S. RAMJATTAN. pp. 10-22. (Mimeo.) Interruptionof malaria transmission by chloro- 4. Giamota, G. Malarial Nephritis. Epidemiologi- quinised salt in Guyana. Bull. Wld. Hlth. Org. 36: cal Notes on Malaria, Blackwater Fever, Albuminu- 283-301, 1967. ria and Nephritis in the Interior of British Guiana, 10. RAtcriENow, E. Zur frar - der Malaria-resisten, Based on Seven Years Continual Observation. Lon- bei Negren. Behefte Z. Arch. f. Schiff; u. Tropen don, Churchill, 1930. 164 pp. Hyg. 3358-64, 1929. 5. Giouou, G. Notes on health conditions in the 11. SPRUCE. R. Notes of a Botanist on the Ama- South Rupununi Savannah. Appendix to Report of zon and in the Andes, II, ed. by A. R.Wallace. the British Guiana Commission to the President's London, Macmillan, 1908, pp. 271-272. Commission on Political Refugees. Georgetown, 12. SWELLENGREBEL, N. H., and E. VAN DER KUYP. 1939, pp. 56-59. (Legislative Council Paper No. 12/ Health of White Settlers in Surinam. Amsterdam, 1939) Colonial Institute, 1940, pp. 92-97.

113 '118 FOOD AND NUTRITION OF THE MAYA BEFORE THE CONQUEST AND AT THE PRESENT TIME'

Moisés Béhor

".. . and grinding then the yellow and the white cars In this paper I shall try to review our unfortu of corn, the goddess hmucané prepared nine beverages, nately small store of information on the clietir) and from this food came strength and stoutness and with practices and nutritional status of the Ma)ar it they created the muscle and vigor of man.. .." populations before the Spanish conquest of tai: "Their flesh was made from yellow and from white corn; from ground and kneaded corn were moulded the arms region. I shall examine the same conditions ir and legs of man. Only damp, ground corn went into the their direct descendants, the Guatemalan In flesh of our fathers. .." dians of today, and attempt to determine tht A greatcivilizationdevelopedinMiddle main changes that have come to pass. It is m) Americo on thc basis of corn. The above quota- hope that, in so doing, some general principle! tions from the Topol Vuh, the sacred book of applicable not only to this particular group the ancient Quiche Maya, emphasize the im- American Indians but to others as yell wil portance ascribed to this staple grain by theemerge. This seems to me more useft.:, than Mayan populations. Nevertheless, our present purely descriptive paper on the nutritional prob knowledge of nutritica has demonstrated that lems now afflicting the American Indian groups corn,ifnot properly supplemented(16),is Dietary practices and nutritional status of the nutritionally very poor as a staple. As in most of pre-Columbian Mayan Indians the cereal grains,itsprotein concentration is low; in addition, corn proteins have a very low Corn, prepared in many different ways, was biological value. They are deficient in at leasteaten at every meal, every day, as the most irn two of the essential amino acids:lysine and portant .:ood. In addition, the Mayas cultivated tryptophane (5). Furthermore, the deficiency ofanu consumed beans, vegetables,fruits,roots tryptophane in corn, plus a low concentration tubers, cacao, and spices (15). Animal products. and/or availability of niacin, favor the develop- mainly meat and to a much lesser extent eggs. ment of pellagra in those whose dietis based were apparently not a .-egular itemintheirs on it (10). dietary; they were consumed sporadically and Tt must therefore be asked how an important on special occasions, and were obtained mainlyt civilization like the Mayan could develop with by hunting. Very significant in this regard is such a great dependence on corn as its staplethe existence, in a Mayan dialect of the Guate-i food; and why the present-day descendants ofmalan highlands, of a spcial word, tcibex, fori the Mayas have serious nutritional problems, "to eat meat," as distinct from the general word; also because of their dependence on corn. lobex meaning "to eat" (14). Only the groups living on the coasts consumed fish. Milk was noti 1 Research supported by Contract PH43-65-640, Na tional Institute of Child Health and Human Develop- used at all. ment (NICHD), Bethesda, Maryland, U.S.A. Our present knowledge of nutrition indicates; that, with such a diet, it is difficult to satisfy the because the biological value of corn protein is requirements for at least three important nutri-about 50 per cent(4). This means that he mts: protein, calcium, and niacin. The defi-would have to consume about 450 g of corn a ziency of calcium and niacin was corrected byday, or about 800 g of tortillas, to satisfy his the way in which corn was processed. For theprotein needs. Not only isthis physically im- ?reparation of tortillas and other corn products,possible but, in addition, the child would be the grain was first soaked overnightin hale receiving many more calories than he needs. water a-od then cooked in the same water (the Of course, the Mayas did consume other foods purpose was to remove the husk). The resultbesides corn. From what we know, the other was a product very rich in calcium, for which important items in the regular daily diet weie the Mayas had no other adequatedietary beans, fruits and vegetables, and rootsand sources. This method also yielded a higher con- tubers. Beans are a more concentrated source of centration of tryptophane inrelationtothe protein, but the remaining food items, although other amino acids, and more available niacin they contribute other needed nutrients, reduce (13), thus preventing pellagra. We can only the over-all concentration of protein in the diet. speculate on how this practice was adopted, butThe situation would again be one in which it is now evident that it was indispensable foradults can satisfy their nutritional needs but their survival. small children cannot. We are still left with the problem of how they I have mentioned that the Mayas did not con- satisfied their nutritional requirements for pro-sume milk, and thatother animal products tein. As I have said, the main drawback of corn (meat aPcl eggs) were only eaten irregularly. proteins is that they are deficient in some essen- It is also very unlikely that they fed meat to tial amino acids, particularly lysine and trypto-infants and small children, for they had not phane. However, an amount of 39 grams of corn developedthzrequit..:dprocessingfacilities. protein a day, which satisfies the average protein There was only one solution, and they applied requirementinan adult male weighing 65itprolooged lactation. The information now kilograms(17),doescontaintheneeded available indicates, in fact, that Mayan mothers amounts of all essential amino acids (12). This breast-fed their children for at least threeyears amount of protein is provided by about 370 gand frequently even longer (15, p. 44). of corn, which is not an unreasonable quantity Our present knowledge indicates that breast for a man to consume per dayin iact, Guate- milk can satisfy all the nutritional requirements malan Indians now consume around 500. In of a baby up to approximately six months of age. spite of the low biological value and low con-It is probable that from about that age Mayan centration of proteins in corn,itistherefore mothers started to give their children what the possible for aduits to satisfy their protein needs rest of the family ate but continued to feed them with tliis grain as the main staple, if they con-breast milk as an indispensable protein supple- sutne enough of it. ment. By the time they were completely weaned, But the situationi,completely different for after the age of three, they were able to eat small children, who have relatively higher pro- enough of the family food to satisfy their nutri- tein requirements and in whom the need fortional requirements without the useof any essential amino acids is also greater. A two-year- special product, such as milk inthe case of old child, for instance, would need about 18 gWestern civilization, Itisalso probable that of protein a day to satisfy his nitrogen require-during this transitional period, from about six ments if the protein has an ideal composition,months to three or four years of age, they did with a biological value of 100 per cent; in other not grow to their full genetic potential. This words, if it is fully utilized (17). With corn ascould be the main reason for the small size of the source, the child would need 3G g of protein, the adults, whichiswell dormmented. This

115 reduction in body size, a consequence ofen- Dietary practices and nutritional condition of vironmental conditions and particularly of nutri- the Guatemalan Indians of today ent availability, can also, I believe, be considered The dietary practices of the Guatemalan In- a mechanism of adaptation to those environ- dians of our time have not changed very much mental conditions. In fact, with reduction in from those of their pre-Columbian body size resulting in a similar reduction in ancestors. The changes that have occurred, however, have nutrient requirements,titt. Mayas were better gradually brought about a deterioration in their able to meet these requirements with the avail- diet and have upset the balance under which able foods. they lived. Unfortunately, we have no informationon Corn, processed in the same way, is still the the health conditions of children in thosepopu- lations.Data on infant andearly childhood main staple. It provides up to 80 per cent of the calories and 70 per cent of the proteins in the mortality could be particularly illustrati- The information available indicates that, in general,diet of the adult population (9). Beans,vege- tables, and fruits complement the basic diet (8). the children were healthy. De Landa (15)says: These products, however, now have a market They grew wonderfully handsome and fat during the value for which the more affluent non-Inditn first two years. Later, their skin waxed dark withcon- tinuous bathing by mothers and many suns; theywere groups in the population are better able to com- nonetheless bonny and mischievous throughout child- pete. This is even more true of animal products. hood. never ceasing to w,,nder about carrying bows and Wild animals have practically disappeared, and arrows and frolicking among themselves and thus they hunting isno longer an important means of grew until they began to adopt the ways of the youths and to acquire a more exalted self.concept, and to leave obtaining precious foods. Domestic animals are behind the things of childhood. now the only important source of animal prod- We cannot say thesame about the majority ofucts, and the market for them is very competi- piTschool children of Mayan descent today; theytive. The majority of the Indian families raise are usually apathetic. and very ofter sick. some animals in their homes, mainly poulti y Anotheris'aponont dedu&on thatcan be and pigs. But these are morea source of badly made from the wtion lace we,ri;rw needed cash than of food. They and their prod- concerns birth spacing.th' pro:aably ucts are consumed only on very rare occasions; cl-kost of the time, !!'),Pv are sold. The meat most had at least three years betwcin 0;1,1 the next. To what extent prolonged l. tati:;n frequently ,2onsuined is beef, which has to be contributedtothis situation we still do Lot trorn i:he market and therefore is one know. However, it was definitelyvery con- lor hidi rash is needed. venient, because only inthis way could the Not eini);La-ce indianl,;1'wtheirold mothers breast-feed their childrenas required. sources of food in the wild forests, but also the Accurate information on the nutritional condi-amount of land available to them for cultivation tionof pre-Columbian Mayan adultsisalso has been drastically reduced. Before the Con nonexistent, but all known facts indicate that,quest, they had all the land they needed to although small, they were in generalstrong and produce their corn. Although the method of healthy. We can thus conclude that, interms of cultivation they used cannot be consideredvery nutrition, they did not have serious problems efficient today, its relative inadequacywas com- and were well adapted to their environmental pensated for by the availability of sufficient land circumstances and degree of cultural and tech- for a rotation system. They did not need to nologicaldevelopment. Theydid,however, produce cash crops, which now compete with undergo acute crises, and there is informationon food crops for the best lands. the famines they suffered when natural disasters Under present-day circumstances, the Indians (droughts or locust invasions) interfered withare frequently forced to buy even their corn, their corn crops. which they had always produced before. Owing

116 :o their poverty, this greaterdependence on the at the present time is barely adequate for adults market to satisfy food needs has caused their and frequently inadequate for children. As a diet to deteriorate. Another change in the adultresult, environmental stresses,particularlythe lietisthe introduction of coffee, which hasheavy burden of frequentinfectionsin early replaced the more nutritious corn and chocolatechildhood, have caused poor nutritional condi- beverages they used to drink, Distilled alcoholic tions for the general population, with frequent beverages have also become part of their con- and F. cvere cases of nutritional deficiencies found sumption habits, with deleterious effects fromparticularly in early childhood. Briefly, the nu- the nutritional and even more from the eco- tritional peoblems now affecting the Guatemalan nomic standpoint. Indians art;.,ksfollows: Changes in dietary practices concerning small The most important by far is protein-calorie children are even more important, and these malnutrition (19). As has been indicated,it is have been mainly the effect of cultural influ- the result of the synergistic effect of an insuffi- ences. In the first place, theperiod of weaning cient and inadequate dietaryintake and fre- has been shortened. In pure Guatemalan In- quent infectious episodes, particularly diarrheal dians, prolonged lactationisstillessential toprocesses and the common diseases ofchildhood children's survival. Complete weaning in these (3). This condition contributes very significantly populations now takes place between the ages of to the high mortality rates in infancy andearly about two and two and a half years; this time, childhood (2). It causes a relatively high preva- already shorter than it used to be before thelence of severe cases of protein-calorie malnu- Conquest, is being progressively reduced as thetrition (kwashiorkor and marasmus) (/), which process of integration into the ladino orWestern are usually fatalif not properlytreatedin a culture advances (11). This situationisvery hospital. Even more important from the public dangerous andhas caused acute nutritional health standpoint arethe chronic, subclinical problems, since the populations adopting earlier forms of malnutrition, which usually go unrec- weaning are not as a rule prepared culturallyognized. These affect a large majority of the and economically to feed their children properly population, both children and adults (18). The during the critical post-weaning period. Their consequences of this condition are not yet clear, general ignorance in matters of hygiene, and the but there is information suggesting that it may lack of the most elementary physical facilities in interfere significantly with mental development their homes, renders the use of milk very dan- in children (6) and with work performance in gerous. Most of them, in any case, cannotafford adults (20). If these observations are confirmed to feed milk to their children. and studies are now under way to test the The ,itu Itionisfurther aggravated by the hypothesisit is easy to understand the tremen- introduction, by foreign civilizations, of refined cous importance of theproblem in terms of the products with a much lower nutritive value than general well-being of these populations and of the foods they replace. This is particularly true tt-eir participation in the social and economic and deleterious for the child during the post- development of their societies and countries. weaning period. A v, example of this problem is Among the other nutritional problems may be the pure starch preparations made from corn or mentioned vitamin A deficiency, primarily the manioc that are otten used nowadays in gruels result of diets in which the sources of t.1,15 vita- for small children, instead of the more nutritieus min are vegetable prodects with iow whole-corn gruels of old. activity consumed in insufficient amounts (8). The Guatemalan Indians of today thus seem This deficiency rewIts in problems as serious as to be in a less favorable balance with their en-keratomalacia, which ends ;n te talblindness vironment than their ancestors were, and their during early childhood. Riboflavin deficiency is dietary practices have deteriorated. Food intake among the most prevalent of the vitamindefi-

117

w ciencies (8). It is closely interrelated with pro- the mother's breast for as Icni; as Fo,sible. They teindeficiency,sincetheadequatedietary do not have at their disposal the 'benefits" ol sources of the two are frequently the same. civilization, such as the practice of bottle feeding Mucosal and cutaneous lesions characteristic ofand very inadequate weaning foods. We must this deficiency are frequently seen. Its possible consider the situation of these groups of prirni. general effects are not clearly understood. tive Indians as they become incorporated into Nutritional anemias are also highly prevalent; our societies, with the hope that we will find the deficiencies of iron and folic acid are themeans to prevent or minimize the nutritional factors most frequently responsible, but againdeterioration to which they may be exposed in protein malnutrition contributes to their devel- the process of integration. opment (21). Anemias may also be important in restricting work capacity. Endemic goiter Summary used to be highly prevalent in Guatemala, but An analysis of the dietary patfrr,:.f3nd the fortunately it has now been controlled bymeans nutritional conditi.,-1 of of salt iodization (7). Mayrn populatiotIs. before the ihe The nutritional problems herein summarized ,t indicatestLy. a:tL .or,Acoist :11e: do -1t3t affic t.the hidian pouPil,,onr. Guate- r,ain stapk kr both pnpul-auongroups, the mala viittifn7 or i.ieterentiaily. Theyare asso- pre-Columbian Mayas were probablybetter uiated wit:0 !w,A, cur. ,ioial inomic levels adapted to ecological conditions. The availability and poor sanitary conditions. Thereare non- of game, greater availability of land for cultiva- Indian populations1,1 .,Lilar conditions, but tion, the absence of competitive markets, pro- practically allthe Indian populations fall into longed lactationof infants, and the lack of this category. Their difficultyis one of proper industrialized products were among the factors integration into and adaptation tonew ecologi- that brought about a better diet for the Indians cal and cultural conditions and socialstructure. at that time. The sociocultural and economic In my opinion, the situationdescribedin relation to the Mayan Indians, and the changes changes that have taken place since the Con- tbat have come to pass, are applicableto a lesser quest have worsened the diet of the present -..1.can:r extent to other Indian groups now Indians and resulted in serious nutritional prob- with Western civilization in the lems, among which protein deficiencyparticu- kmcricali cotintries. On the other hand,larly in small childrenis the most important. many intl;an groups ;Ail: 13ving in isolation from This experience should be considered both in Western Cure and from thepresent Latin tr, mg to solve the nutritional problems of this American societics have less serioLs m.tritional ahd other American Indian gro..ips now living problems; some of them seem to be liNiug under with pui.,Intions of Western culture and inpre- excellent conditions in thisrespeu. e groups venting similar changes that could occur as the are still very dependent on hunting and Usliing still primitive and isolated Indiangroups are to satisfy their food needs, and still follow theincorporated into modern Latin American cul- "primitive" practice of raising their childrenat ture.

REFERENCES 1. Au Tiirr, M., and M. BEHAR. Sindrome poli- in four rural communities in Git9tcipall. Bull. Wld. carencial in/anti! (kwashiorkor) and its prevention Hlth. Org. 19:1093-1102, 195e. in Central America. Rome, Italy, Food and Agri- 3. BEHAR, M., and N. S. SCRIMSHAW. Epidemi- culture Organization of the United Nations, 1954. ology of protein ii.alnutrition.InIago Caldston (FAO Nutritional Studies, 13) (ed)., Human Nuto,i Historic Scientific. 2. BEHAR, M., W. ASCOLI, and N. S. SCRIMSHAW. New York, International Universities Press, 1960, An investigation into the causes of death in children pp.257-273.(InstituteofSocialandHistorical

118 123 Medicine, The New York Academy of Medicine, nutrients-Amino acids. Fed. Proc. 22:1424-1430, Monograph III) 1963. 4. BLOCK, R. J., and H. H. MITCHELL. The corre- 13. KODICEK, E., R. BRAUDE, S. K. KON, and K. G. lationof the amino-acid composition of proteins MITcHELL. The availability to pigs of nicotinic acid with their nutritive value. Nutrition Abstr. Rev. intortilla baked from maize treatedwith lime- 16:249-278, 1946-1947. water. Brit. J. Nutrition 13:363-384, 1959. 5. BRESSANI, R., N. S. SCRIMSHAW, M. BEHAR. d 14. LA FARGE, 0. The Religion of a Cuehumatan F. VITERI. Supplementation of cereal prot": .'rd;an Town. Chicago, University of Chicago Press, amino acids. 11. Effect of amino acid sth:ple:1 1947, p. 37. tion of corn-masa at intermediate -levels o rotein 15. LANDA, FRAY DIEGO DE. Relación de lascosas intake on the nitrogen retention of young children. de Yucatan (compuesto por el alio 1560). 8a. ed. 1. Nutrition 66:501-513, 1958. Mexico, Editorial Porrua, 1959, pp. 36, 37. (Biblio- 6. CRAVIOTO, J. Application of newer knowledge teca Porrua, 13) of nutrition 1-.1physical and mental growth and 16. ORGANIZACION DE LAS NACIONES UNIDAS PARA (I. )1. t Am. 1. Pub. Health 53 :1803-1809, LA AGRICULTURA Y LA ALIMENTACION. El maiz en 11,1;3. lu alimentaeión. Estudio sobre su valor nutritivo. 7. DE LEON, J. R., and 0. G. RETANA. La erradi- Roma, FAO, 1954. (Estudios sobre NutriciOn, 9) caciOn del bocio endémico en Guatemala; informe 17. Protein requirements. Report of a Joint FAO/ final. Nutrición Bromatol. Toxicol. 5:43-54, 1966. WHO Expert Group. Published jointly by FAO and 8. FLORES, M., B. GARCIA, Z. FLORES, and M. Y. WHO. Rome, Food and Agriculture Organization LARA. Annual patterns of family and children's oftheUnited Nations,1965. (FAO Nutrition diet in three Guatemalan Indian communities. Brit. Meeting Report Series, 37; WHO Tech.Report 1. Nutrition 18:281-293, 1964. Series, 301) 9. FLORES, M., and E. REH. Estudios de htibitos 18. SCRIMSHAW, N. S., and M. BinAR. Malnutri- dieteticos en poblaciones de Guatemala. I. Magda- tion in underdeveloped countries. New England J. lena Milpas Altas. Bol. Ofic. San. Pan. Sup. 2, 90- Med. 272:137-144, 193-198, 1965. 128, 1955. 19. SCRIMSHAW, N. S., and M. BEHAR. Protein 10. GowsmITH, G. A., G IEBE NS, W. G. UNGLAUB, malnutrition in young children. Science 133:2039- and 0. NEAL MILLER. Studies of niacin requirement 2047, 1961. in man. III. Comparative effects of diets containing 20. ToRt3N, B., F. VITERI, F. SARANTIA, and M. lime-treated and untreated corn inthe production FLORES. Body composition in Guatemalan adults of of experimental pellagra. Am. 1. Clin. Nutrition 4: different nutriture. Fed. Proc.27:679, Abt. 2605, 151-160, 1956. 1968. 11. GoNzAuz, M. L. SOLIEN DE.Breast-feeding, 21. VITERI, F., J. ALvARAno, D. G. LUTHRINGER, weaning and acculturation.J.Pediat. 62:577-581, and R. P. Wool), H. Hematological changes in pro- 1963. tein-caloriemalnutrition.Vitamins & Hormones. 12. HEGSTED, D. M. Variation in requirements of 26, 1968. In press.

"9124 IODINE DEFICIENCY WITHOUT GOITER INISOLATED YANOMAMA INDIANS: PRELIMINARY NOTE

René Rivière, Dominique Comer, Monte& Colonomos,Jean Desenne, and Marcel Roche'

Roche et al. (10) showed that, ina region of endemic goiter, subjects withoutas well as those with goiter had an elevated thyroid radioactive iodine uptake. A similar findingwas reported in New Guinea by Cboufoer, Van Rhijn, Kas- senaar, and Querido (1). In an isolated primi- tive Venezuelan Indian community, Roche (9) demonstrated a generally elevated thyroidup- take without goiter; and thiswas again de- scribed in the eastern regionofthe Congo Republic by Delange, Thilly, and Ermans (3). In the present preliminarynote, further quan- titativeaspectsofthyroidmetabolismina Yanomama group from theupper Orinoco region are presented.

Material and methods The area chosen for study lies roughly be- 72 ?0 tween the Ocamo and Mavaca rivers on the FIGURE 1. Area selected for study of thyroid metabolism Orinoco (Figure I),in the extreme south of in Yanomarna Indians (see page 77 for map of Vene- Venezuela. Itis reached by plane to La Esrne- zuda) raldaan old mission site visited by Humboldt (7) in 1800and upstream from there by fast eithernot palpableorbarelypalpable. A river boat in about four hours. Two expeditionsCatholic and a Protestant mission were estab- were carried out. lished at the site, but the Indians still lived in During a preliminary one in February 1962,wide-open thatch huts, ate their own food, and thyroid iodine uptakewas determined on 41 went completely naked. Yanomarna Indians living in Bisaasiteri, at the Another expedition was carried out in Janu- confluence of the Orinoco and the Mavaca. All ary, February, and March 1968 to Bisaasiteri the subjects studiedwere ad- quately nourished, and to the neighboring Ocamo River (Tyowci- in good health, and with thyroids thatwere teri). The living conditions of the Indians had changed, although they remained very primi- I Presented by Dr. Roche, tive. They no longer lived in the characteristii:

120 123 huts, but rather in closed dwellings; they occa- Results sionally received food and medicines from the missions. A third group, from Macorimateri on Figure 2 shows the thyroid iodine uptake in the Manaviche river, was also contacted this Bisaasiteri in 1962 (left) and the uptake in the time; this group, unlike the first two, was still combined populations of Bisaasiteri, Tyoweiteri, culturally isolated in all respects. and Macorimateri in 1968. Table 1 shows data on On the second expedition, 56 adult subjects- iodine metabolism. The iodine content of river 44 male and 12 femalewere studied. Theywater was 3.2 pg/liter (average of six samples). differed from those studied on the first expedi- tion. They were inapparently good clinical nutritional state, of short stature (males 1.57 m Discussion and 52 kg, females 1.44 m and 45 kg, on the As Figure 2 shows, the thyroid iodine uptake average) and without any clinical evidence of is higher in all the subjects studied in 1962. In thyroid dysfunction. In 45 of them the thyroid 1968, although many uptakes are still high, this was not palpable; in 10it was palpable but time five individuals have uptakes below 40 per estimated to be of normal size. In one subject cent. It is presumed that the lowereduptakes are (a male) there was a small diffuse goiter, esti- due to the introduction of foreign food and mated to weigh around 50 g. medications; none of the uptakes in the more Twenty-four-hour thyroid uptake was deter- isolated group were low, and the average in that mined after the oral or intravenous administra- group was higher than in the other two(Table tion of 5 to 30c of "'I, in the manner recom- 1). mended by the Intei national Atomic Energy The urinary iodine/creatinine ratios (Table Agency (12). A figure on the uptake of the first 1) are low compared to normal values (5). The group has been published (6). Kineticstudies total 24-hour iodine excretion can only be indi- are being continued after the oraladministration rectly evaluated. The values obtained are ap- of 100 fic of laq and will be reported later, as proximations, since urinary excretion of iodine will the effect of TSH and of methimazole and isnot uniform throughout theday and since other measurements. 24-hour urinary creatinine is not an accurately Radioactivity was measured on the site of the fixed value. As a matter of fact, creatinine excre- study; the various iodinated fractions of plasma tion is related to muscle mass and in theseshort, were separated on ion exchange resins and mea- thin Indians is likely to be less than is taken sured separately. Measurements of urinary '271 here as normal (4.) In a group of Makiritare In. and creatinine were performed on single urinary samples because of the practical impossibility of obtaining reliable 24-hour samples. 1968 The blood, urine, and river drinking water samples were refrigerated around 40 C when indicated and sent by plane to the central labora- 0- tory in Caracas. Stable iodine was measured by activation analysis in the Nuclear Chemistry Department of the Venezuelan Institutefor 5 Scientific Research and at the Service Hospitalier Frederic Jo liotin France (2). The iodine/

creatinine ratio was measured to give an idea of 102030 70 100

the daily urinary iodine excretion. The average Thyroid Uptake i',44 24-hour creatinine excretion was assumed to be FIGURE 2. Frequency distribution of iodine131 thyroid 1.6 g for mcn and 1.0 g for women (5, 8). uptake of Yanamama Indians

121 126 TABLE 1.Iodine metabolism in isolated Venezuelan Indians (average, standard deviation,range, and number of subjects) - --- 24-nouR PLASMA MAXIMUM 1 25I THYROID 1311 127I0DINE (AO / THYROXINE PLASMA LIPTAK E CREATIN1NE (g) I 271 URINARY 127I THYROXIN SITE ( OF DOSE) RATIO (jig/ l 00 cc) (Aig/DAY) (70/LITER)

64.3 29.1 4.42 42.6 0.445 ±-7.4 ±-7.8 Bisaasiteri ±.0.107 36.0-81.3 10.0-104.0 15.0-166.0 0.194-1.100 (26) (35) (22) (35) (18) 61.1 0.679 Tyoweiteri 48.6-70.0 0.207-1.450 (10) (10) 74.1 19.1 5.07 30.5 ±-6.4 Macorimateri 32.9-93.0 30.0-70.0 2.6- 7.4 16.0-48.0 (12) (10) (12) (10) dians, at the Ventuari River, Roche, Perinetti, pears evident that goiter may fail to occur even and Barbeito (Li) found a 24-hour iodine excre- though, as is likely in this case, iodine deficiency tion of 21.2 p.g/day with a mode between 10 and hasbeen present through many generations. 20 pg. Iodine in the drinking water was low in What other factor or factors are needed remains the Orinoco, although not as low as in the in question. Ventuari, where, by a different method, 0.2 to 0.6 lig of iodine was found. Summary Plasma thyroid 127I was normal in both of A study of iodine metabolism was carried out the groups studied, which is in accord with thein 48 Indians from the Yanomama tribe living observatioilthatthesesubjects areclinically near the confluence of the Orinoco and Mavaca euthyroid. rivers in Venezuela. Elevated thyroid iodine up- The high thyroid iodine uptake and the low take, low urinary iodine/creatinirie ratio, normal urinary 19 excretion are comparable to those plasma thyronine, and apparently normal clinical found in a goitrous Andean region of Venezuela thyroid status were demonstrated, in the absence (Riviere, Camuzzini, Comar, and Roche, to beof goiter. Itis suggested that iodine lack is a published) and indicate that these goiter-free necessary condition for the formation of endemic Indians are indeed iodine-deficient. It again ap- goiter.

REFERENCES

1. CHOUFOER, J.C., M. VAN RHIJN, A. A. M. 3. DELANGE, F., C. THILLY, and A. M. ERMANS. KASSENAAR, and A. QUERIDO. Endemic goiterin Iodine deficiency,aprimitive conditioninthe western New Guinea: iodine metabolism in goitnous development of endemic goiter. I. Clin. Endocrinol. and nongoitrous subjects. I. Clin. Endocrinol.and and Metab. 28:114-116, 1968. Metab.23:1203-1217,1963. 4. DE VENANZI, F., M. ROCHE, and A. GERARDL 2. COMAR, D., R. RIVIERE, and C. KELLERSHOHN. Captación de yodo radiactivo(1131 )por sujetos L'analyse paractivationetsesapplications aux eutiroideos de nuestro medic), y algunas considera- sciences biologiques. In Troi.iietne Congres Interna- clones sobre aplicaciOn al diagnOstico de las enfer- tional de Biologiede Saclay.Paris, Presses Univer- medades tiroideas. ActaMed. Venez. 3:114-118, sitaires de France,1964, pp. 307-336. 1955. 5. DWORIlsi, H. J.,J.P. CHANDLER, and W. H. the absence of goiter in isolated Venezuelan Indians. BEIERWALTES. Estimation of fecal and urinaryexcre- J. Clin. Endocrinol. ind Metal,. 19:1440-1145, 1959. tion of iodine, J.Clin. Endocrinol. and Metab. 25: 149-151, 1965. 10. ROCHE, M., F. DE VENANZI, J. VERA, E. COLI., 6. GAEDE, K. Algunos eswdios recientes sobre el M. SPINETTI-BERTI, J. MENDEZ MARTINEZ, A. GER- bocio endemico. Acta Cientifica Venez. Supp. 1:58- ARD!, and J. FORERO. Endemic goiter in Venezuela, 74, 1963. studied with P31. I.cli,i. Endocrinol. and Metab. 7. HUMBOLDT, A. DE. Voyageaux regions equi- 17:99-110, 1957. noxiales du Nouveau Continent.Paris, Librairie 11. ROCHE, Y., H. PERINETTI, and A. BARBEITO. Grecque-Latine-Allemande, 1816. Urinary excre,.ion of stable iodine ina small group 8. JOLIN, T., and F. ESCOBAR DEL REY. Evaluation of isolated Venezuelan Indians. f. Clin. Endocrinol. of iodine/creatiaineratios of casual samples as 21:1009-1012, 1961. indices of daily urinary iodine output during field 12. The Calibration and Standardization of Thy- studies. J. Clin. Endocrinol. and Metab. 25:540-54', roid Radioiodine +Uptake Measurements. Vienna, 1965. International Atomic Energy Agency, November 9. ROCHE, M. Elevated thyroidal P31 uptake in 28-30, 1960. STUDY OF ENDEMIC GOITER IN THE AMERICAN INDIAN'

José Bane lotto and Edmundo Covarrabias2

The existence of endemic goiter in America mented examples are those of the United Statcs can be traced back to pre-Columbian times (5, (22), MLxico(40),Guatemala(39),Colombia 4i), although the opposite view can be sustained (6),and Argentina(33).Iodine prophylaxis (14,15).Endemicgoiterareashavebeen programs are being increased in number and demonstrated during the last decades in practi- effectiveness in America and will soon influence cally every couutry in the Americas. From the goiterepidemiology,atleast among Indians review published by Kelly and Snedden in 1960 more closely integrated with the surrounding (21),it can be estimated that atleast 20 per soc iety. cent of the populationisaffected by goiter. The idea that endemic goiter reflects only the From epidemiologicalsurveys comprisingin adaptation of man to iodine deficiency can no some cases huge samples, it can be inferred that longer be sustained. Asn other parts of the all races are affected. Unfortunately, there is no wczld,itis contradictedir America by such precision about racial distribution in most ofexamples as the finding of groups of Indians the studies; present-day American Indians are without goiter who were iodine-deficient(37) mixed in different degrees and have been in-andthepersisting development of endemic cluded in variable proportions. Those Indians goiter in children born in areas where effective who live in prinlitive conditions and are pre- iodine prophylaxis was under way(13).Natural sumably less mixed have been the subjects of goitrogens occurring in food have been suspected only a few specific goiter surveys(35,46). of playing a role in the pathogenesis of some Iodine deficiency in the populations of en- endemic goiter (a23. 32, 34).An example in demic goiter areas in America, as judged by our continent is the pilion or nut of theArauca- urinary excretion, has been found in prac,ically nia araucana,a pine tree abundant in south- every case where it has been looked for (2.17, central Chile(42). 18, 19, 24, 26, 36, 4!).This is not surpris.ng, Contrasting with thesituationin sporadic since except for the nitrate deposits in northcm goiter, genetic mechanisms have not been well Chile(2d)iodine is not abundant ib the surfaci documented inthepathogenesis of endemic of this hemisphere as itisin the rest of the goiter. Enzymatic detects have not been demon- world. Wherever thediethas been supple- :grated initsetiology. A concentration of en- mented with iodine, the incidence of endemic demic goiter cases within familieshas been goiterhasbeengreatlyreduced;well-docu- reported in America(16,25)and elsewhere (7, 27). Apositive association of nodular goiter I Work supported by Comisi6n de Ayuda a la Investi- with the distribution of nontasters of phenyl- gaci(inCientifica.Facuityof Medicine, Universityof thiocarbarnide (PTC) I as been reported in a Chile, Contvact 64/4, and Comisión Nacional de Investi- gad6n Cientifica y Tecnol6gica, Project 41. general mixed population in Brazil (28). How- 2Presented by Dr. Barzelatto. ever,thesearchforthisassccidtion among

124 4;9 American Indiansis rendered difficult by the Endemic goiter in Pedregoso small proportion of nontasters usually found In accordance with the foregoing line ot among them, as was the case in a study of the thought, we have been involved for the last five Pewenche(10). years in a study of endemic goiter on a Pewenche Another aspect to be considered is the prob- Indian reservation in Chile. Some of our find- lem of endemic goiter variability. The anatomic ings will provide a good basis for discussion of and physiological characteristics of goiter may these ideas. vary in a single individual during his rietime; Pedregoso is a geographically isolated Andean other levels of variation are to be defined among valley at an altitude of 1310 meters, with a the members of a given community and among cold climate. According to a 1965 census, 592 different populations. These three types of vari- Pewenches livethere, by relativelyprimitive ability make it a hard task to compare results agriculture and some cattle raising. Pinon isa of different authors, especizlly because there is standarditemintheirdiet.The available still no general agreement on definitions. Some genetic data show that these IndiansL....eless methodological limitations should also be borne mixed than other groups of the Araucanian in mind, such subjective differences when not complex to which they belong(8, 10, 29, 30, 38) all individuals are examined by the same per- and constitute a highly inbred populationin son. Even if these are overcome, difficulties in which a quarter of the marriages are con- classification persist, such as estimating the size sanguineous(10). of a goiter in the presence of retrosternal exten- Endemic goiter in Pedregoso was found to sion and the error induced by missing goiters have a prevalence of 66.8 per cent in a sample that have become totally retrostemal. comprising 81 per cent of the total population The impact of endemic goiter in public health (the remaining19percent,mainly young people, were not examined in our house-to- isanother matter of debate. Severe endemic house survey because for one reason or another goiterisassociated with endemiccretinism, they were away from home). The distribution deaf-mutism and neur)logical defects, but the according to sex and age is shown in Table I, mechanisms involved are not clearly understood. which also gives the frequencies of clinically There are also no adequate studies of mortality nodular and large goiters (including those esti- and fertility differentials in comparable popula- mated to weigh 60 grams or more and all those tions with and without endemic goiter, or inwith retrosternal extensions). As in most other individuals within an endemic goiter area. Instudies, women were found more frequently both cases fruitful information may be obtained goitrous at all ages than men. The fact that the from looking into relations with endemic goiter frequency does not diminish after puberty and variability. the finding of nodular goiters in the first decades Comparative studies among different groups of lifeare indications of the severity of the of American Indians could provide an excellent endemic. On the other hand, huge goiters were opportunity to study the pathogenesis of en-e.;ceptional and only one cretin and no deaf- demic goiter. They presumably have very differ- mutes were found. enz genetic endowment, and also are influenced The first problem to be considered is whether by different environments. The interplay of all the increases in size and in nodularity with age these factors in the genesis of endemic goiter isare independenteffects.Nodularityinereases probably unique in eachpopulation; hence with age in both sexes, appearing earlier among studies in depth of endemic goiter in certainwomen (Table 2). Large goiters are more fre- communities may be scientifically rewarding,quently nodular than small ones with advance besides providing the basisforrational pro- in age, but some individuals have small nodviar phylaxis programs. goiters in the first decades. Itis obvious from

125

, TABLE I. Goiter in Pedregoso

MEN WOMEN

EXAMINED GOITROUS EXAMINED GOITROUS

AGE % (YEARS) GOITROUS N NODULAR LARGE N GOITROUS NODULAR LARGE

0-9 63 25.4 16 0.0 6.2 71 39.4 28 3.6 0.0 10-19 55 72.7 40 2.5 2.5 55 72.7 40 15.0 15_0 20-29 37 62.2 23 26.1 21.7 44 95.5 42 23.8 45.2 30-39 28 67.9 19 47.4 42.1 33 100 0 33 54.6 30.3 40-49 16 75.0 12 66.7 50.0 17 94.1 16 81.2 68.8 18 50 and over 29 76.9 22 68.2 63.6 96.4 27 77.8 81.5 Total 228 57.9 132 29.5 2(.3 248 75.0 186 37.1 36.6 these data that if th2 influence of other etiolog:- which may have been ignored when diagnosing cal factors upon endemic goiter variability is to small goiters, particularly if nodular. be studied, sex, age, size, and nodularity shaild The data should therefore be analyzed both be considered separately. A number 2e facts by sex and by age group. in each et' these cat..:- reinforce this conclusion: (I) The greater sever- gories, five subgroups have to he studied: with- ity of goiter among women, as judged here by out goiter and with small diffuse, large diffuse, greater frequency, larger size, and morenodu- small nodular, and large nodular goiter-. Since larity, and also by the earlier appearance of these the size of the population aqd the distribution of characteristics,isduetopoorlyunderstood goiter in Pedregoso do not generally allow such mechanisms related to sex. (2) Men frequently analysis, more inclusive groups are necessary. In remain away from Pedregoso for long periods the first decade of lifeitis only important to workinginnearby townsorfarms, while compare subjects with and without goiter,ignor- women and children tend to stay onthe reser- ing their sex. The second decade is a transitional vation. (3) Since most of the goiters becomegroup, in which largeand nodular goiters are nodular with age, nodularity among younger still infrequent and cannot be considered apart. irdividuals may have a different meaning from Among the ar'ailts there are relatively adequate that seen in most older subjects. (4) Estimat2s numbers of all groups except that of women of size among older individuals are liable to without goiter. Finally, among older individuals greav-.1 error, because cretrosternal extension, the meaning of thzse group: becomes less pre-

TAItl_,E 2. Nodularity and goiter size in Pedregoso

MEN WOMEN

SMALL .JOITERS LARGE GOITERS SMALL GOITERS LARGE GOITERS AGE (YEARS) % NODULAR % NODULAR % NODULAR N NODULAR

0-9 15 0.0 1 0.0 28 3.6 0 0,0

10-19 39 2.6 1 0.0 34 14.7 6 16.7 20-29 18 22.2 5 40.0 23 13.0 19 36.8 30-39 11 36.4 8 62.5 23 39.1 10 90.0 40-49 6 50:0 83.3 5 80.0 11 81.8 50 and over 8 25.0 14 92.8 5 40.0 22 81.8 Total 97 14.4 35 71.4 118 21.2 68 64.7

126 13t cise, except for small diffuse goiters,which arc those whose parentswere born in different less subject to errors in classification. reservations. Other comparisons showno signifi- Let us first consider the effect ofpregnancy cant differences. To sum up, itcan be said that upon goiter variabilicy in Pedregoso. In each inbred -people in Pedregoso developgoiters at a decade, women with either largeor nodularlater age and that ther goiters are somewhat goiters have hada greater average number of less prone to develop nodules. pregnancies (Table 3)which wouldsuggest PTC-tasting abilitywas determined, with the that gestation increases goitersize and nodu- Harris and kalmus technique, in 255persons in larity if this interpretationwere not challenged Pedregoso. Since only 4.3per cent were non- by the fact that thesame tendencies are ob- tasters, this characteristic could not be properly served amongmen. Men and women under 30related togoiter. Nevertheless,a comparison years of age were not included in Table 3, made among tasters showed that bothmen and although they showed thesesame iendcncies. women with nodular goiters in the third and The differences are significant only formenfourth decades of life had significantlylower with large as against those with smallgoiters inmean threshold values than those with diffuse the older group ("t"(3,)=2.337;p<0.05). Itgoiters (9). should be added thatmen without goiter aver- Height was also correlated with goiter (Table age a slightly higher number of descendants5), and it was shown that taller subjectsof both then goitrous men in thesetwo age groups. sexes had larger goiters, in the third and fourth Inbreeding alsoseems toaffectgoiterin ..iecade. An analysis of variancecombining all Pedregoso --(Table 4). The prevalence ofgoiter these groups showed that this relationis signifi- among children under 10 yeafs whose parents cant at the .05 level (F{,,loe)--7-4.04). Nongoi- are consanguineous is significrmtly lower than trous men of the same age are in general taller among children of parents who were born inthan goitrousones, but this differenceisnot Pedregoso but in whom consanguinitywas notsignificant even when the comparisonis made detected (x( 1)2(Yates) = 4.275; p<0.05) or chil-only with males with small goiters("t",,," = dren of parents bornon daerent reservations 1.857). (x, )2(yti.$) = 4.275;p<0.05). The opposite Nutritional surveys made in Pedregosore- trend seen amongpersons over 20 years of agevealed two relevant facts: first, theexistence of is not significant. Inbred adults showa consist- iodine deficiencyas judged by an average daily ent and significant tendency to have fewer nodu- urinary excretion of 33.2 micrograms(2); and lar goiters (x(r)a(Tate.)=6.264; p<0.025)than second, the fact that allyear round the people consumed pifion (1), which has beendemon- TABLE 3. Fertility and goiter in Pedregoso(total off- strated to be goitrogenic inrats (42). Goiter is spring, abortions excluded) apparentlyinfluencedbythese twofactors. Subjects with largeor nodular goiters have 30-39 YEARS 40 YEARS OR OVER lower mean urinary iodide excretions thanthose GOITER N X se se with small or diffuse goiters (Table6). These Women differences are better demonstrated bythe per- centage of subjects in each group excreting 50 Diffuse 11 4.73 0.406 7 7.43 0.922 Nodular 15 5.73 0.547 31 8.29 0,605 micrograms or more. With this approach,sta- Small 18 4.89 0.427 9 7.33 ).414 tistical significance at the .05 level isreached Large 8 6.25 0.620 29 8.38 0.531 when large goitersare compared with small Men (X(1)2(Yates)=4.162); nongoitrous subjects do Diffuse 9 4.88 0.389 11 7.55 0.790 not differ significantly from those with diffuse Nodular 6 6.00 0.775 22 9.59 0.973 and small goiters. On the other hand,there is a Small 5.00 0.408 13 7.00 1.104 direct relation betweenmean piiion ingestion Large 6 5.83 0.792 20 10.20 0.841 and thyroid size (Table 7),nongoitrous subjects

127 132 TABLE 4. Inbrce&ng and goiter in Pedregoso (men and women)

EXAMINED GOITROUS

AGE GROUP INERELDING % GOITROUS % NODULAR % LARGE

0-9 a 35 14.3 5 0.0 0 72 38.9 28 3.6 3.6 27 40.8 11 0.0 0 10-19 18 61.1 11 18.2 9.1 56 80.4 45 4.4 11.1 35 65.7 23 13.0 4.3 20-29 a 8 87.5 7 0.0 14.3 53 84.9 45 24.4 40.0 19 63.2 12 33.3 33.3 30-39 a 5 100.0 5 40.0 20.0 42 88.1 37 54.0 35.1 14 71.4 10 50.0 40.0 40 and over a 4 100.0 4 50.0 50.0 43 86.0 r 70.3 73.0 33 87.9 29 82.8 65.5

°Parents born in Pedregoso; consanguinity detected. b Parents born in Pedregoso, consanguinity not detected. e Parents born on different reservations. showing the lowest consumption. Sincethe comparable and slightly-abovf.-normalfigures values observed differ widely, it is preferable tofor net iodine intake by the thyroid and periph- study in this case the proportion of subjectseral consumption of thyroid hormones; (2) a eating greater amounts, say 100 or more grams considerable "iodine kak," since the total out- daily; again the same trend is seen, the difference put of iodint by the thyroid is five to ten times being significant when subjects with large thy-thc amountincorporatedintothyroidhor- roids are compared with the rest (NX(1)2( 'Yates) = mones; and (3) the abnormal presence the 4.061; p <0.025). Diffuse and iiodular goiters do blood of increased amounts of endogenously not differ by this approach. labeled triiodothyronine (T3), iodinated poly- Kinetic studies of iodine metabolismper- peptides(NBEI), andiodotyrosines.These formed in Pedregoso (3, 4) are summarized in facts, plusincreased radioiodine uptake and Figure 1, the most striking findings being (1) PBP25 levels, express an accelerated turnove- of

TABLE 5. Height (centimeters) anc. ,.ademic goiter in Pedregoso

GOITROUS

NONGOITROUS DIFFUSE NODULAR SMALL LARGE AGE (YEARS) se se se S, N 3c" se

Men 20-29 14 162.1 1.85 17 159.81.22 6 160.02.81 18 159.41,41 5161.51.00 30-39 9 164.22.41 9 160.81.18 8 163.1 1.47 11 160.7 1.02 6164.2 1.67 40 an I over 10159.5 1.10 11 159.81.3720159.01.48 13 159.01.04 18159.4 1.15 Women 20-29 2 150.02.50 32 150.20.90 10150.51.86 23 148.41.07 19 152.51.01 30-39 - 15 150.81.54 17 149.0 1.12 22 149.31.0210151.01.07 40 and over 2 150.02.50 9 151.91.7630149.80.86 9 150.31.8830150.30.85

128 TABLE 6.Urinaryiodide and goiter in Pcdregoso TABLE 7.Piiion ingestion and goiterinPeciregoso (micrograms per day) (grams per day)

50 Aeg oR 100 G OR S2 MORE (%) N X 52 MORE (%) Nongoitrous 12 36.6 388.99 25.0 Nongoitrous 22 16.0 1604.67 4.5 Goitrous Goitrous Diffuse 30 38.1 545.13 26.7 Diffuse 43 61,6 3315.72 18.6 Nodular 37 31.2 324.20 13.5 Nodular 50 75.7 13665.77 16.0 Small 32 38.5 560.13 31.2 Small 41 57.5 2921.71 9.8 Large 35 30.4 288.24 8.6 Large 52 78.4 13462.44 23.1

radioiodine by the gland and considerable ineffi-shows that nodular goiters havesignificantly ciency in handling iodine interms of the per- higher values for thisparameter, as is also true centage used for the synthesis of thyroid har-for NBEI125. Ina previous publication (4) we mones. Some of these findings have been found have shown that nodular goiters have thewidest in other instances of endemic goiter in America range of values of thyroidal iodine pool,com- (31,41)and elsewhere (12). prising both the smallest and the largestfigures How should radioiodine studies be correlated observed. This findingmay explain why PBP2a to goiter variation? The PB1125 reflects the sp.ted values have the greatest varianceamong nodular of the thyroid in incorporating radioiodine intogoiter, small pools determining thehigh values circulating iodinated compounds, and Table8 observed. In support of this view, thethyroidal

Thyroid iodine poola 11 mg.

r 97,6ugioyj + TYROSINE MIT- DIT MIT-DIT

NBEI

13 - T4

102 ,ug/doy

T3 14 1- +TYROSINE

Thyroid cltput a 512- 833 - 974 iug/doy

FIGURE I. Iodine metabolism in Pedregoso

129 134 TAB1E 8. P8112;1 and NBEI12 in Pedregoso(%/dose/ p<0.01), while there is no significant difference liter) between subjects with small and large goiters. Nongoitrous persons have labeled iodotyrosines pm 125 NBEI 125 in a smaller proportion than goitrous ones, 52 52 N .7c 71-Z particularly in comparison to those with nodular goiters. Subjects with labeled iodotyrosines have 100.170.0135 70.0120.04176 Nongoitrous significantly higher PBP25 values("t"(40)= Gpitrous 2.31; p<0.05); this is not true for those with 300.180.0204 190.0120.04823 '9Diffuse labeled triiodothyronine. The demonstration of Nodular 340.500.1874290.0280.038736 Small 300.310.1147230.0220.017575 iodotyrosines can be directly ascribed to small Large 340.390.1529250.0220.035058 thyroid iodine pools, as is suggested by the fact F for nodularity 13.709 a 8.005 b that six subjects in whom they were present had F for size 2.323 0.097 an average pool of 1.97mg±0.327, while five others in whom they were not demonstrated had a p < 0.001. 1.757 ("t"{4,= b p < 0.010. an average value of 6.22 mg± 2.37; p<0.05). iodine pool among 21subjects in Pedregoso, calculated by the Riggs isotopic.,..quilibrium Discussion method (4), shows a significant inverse rela- Multiple causes are involved in the etiology tionship with the PBP25 values observed inand pathogenesis of endemic goiter, many of these subjects (r=0.472; p<0.025). A similar whichstillrequire properidentification and correlation has been observed in euthyroid pa- evaluation. Iodine deficiency is usually the pre- tientsin Scotland (45). Furthermore, small dominant etiological factor; other environmental iodine pools could also explain high NBEP25 influences are probably also present and, where values, since this parameter shows a significant thereisadequateiodine,would sometimes positive correlation with P131125 (r =0.50; p< suffice to explain endemic goiter. Natural goitro- 0.001). gens are the best.knownexample. Genetic fac- Endogenously labeled triiodothyronine is chro- tors have not been shown toinfluence the preva- matographically present in the same proportion lence of endemic goiter, although the possibility among all goitrous subjectsand absent in theof their playing some role cannot be discarded. nongoitrous (Table 9), although the difference The participation of all these factors should is not significant (x(n2,1-1,tes)=2.549). Onthe be studied in relation not only to the occurrence other hand, endogenously labeled iodotyrosines but also to the variability of goiter. Such studies were present in asignificantly higher percentage are seldom made, presumablybecause of the of the Indians with nodular goiters thanof difficulties in definition and methodology that those with diffuse goiters (x1( )2( yates)=5.71; have been mentioned. It is upon variation that genetic factors have been demonstrated, as ex- TABLE 9. Endogenously labeled triiodothyronine and emplifiedbytheassociation of PTC-tasting iodotyrosinesinPedrcgoso ability with nodular goiter. Weak environmental ,r 1ODOTYROSIN ES '3 etiological factors may also be better studied

NO. NO. with this approach, when a predominant cause like iodine deficiency is present. Nongoitrous 8 0.0 14 21.4 In Pedregoso we have looked into goiter varia- Goitrous tion with three different elements in mind: Diffuse 11 36.4 28 10.7 Nodular 25 36.0 34 41.2 presence, size, and nodularityof goiter. These Small 14 35.7 26 19.2 obviously interact in some measure, but each is Large 22 36.4 36 33.3 characterized in our data by a particular con-

130 stellation of statistically significant factors. Nodu-seem necessary to apply to all these dataa multi- larity is related to the two clearly defined genetic variate analysis through which theproper value aspects studied so farinbreeding and PTC-could be attached bothto main factors and to tasting abilityand to some radioiodine parame- the many kinds of interactionspresent.This ters. Inbred people develop fewer nodular goi- might not be possible, however, fortwo main ters; Indians with nodular goiters have lowerreas xis: the small size (34 our sample, especially thresholdsfortasting PTC; and radioiodine since the subsamples in whichparameters wep: studies shuv high 1313I125 and NBEI125 values recorded did not always comprise the and a greater proportion of endogenously la- same indi- viduals, and the fact that the categories beled iodotyrosines in subjectswith nodular of goiter contain different types of goiters.All these rahoiodine findingscan be error in a nonrandom way and have different meanings. For example, linked to the predominance of smallthyroidal relationships with age iodine pools among those with nodulargoiters. are difficult to study be- Large goiters, on the othcr hand,are associated cause small diffuse goiters at an early agemay with the two best-defined environmental factors develop into large and/or nodulargoiters. Our studied by us: the subjectseat less iodine and interpretation of the Pedregoso datarequires us more piiion. Sizeisalso positively associated to extend these studies to other similar Pewenche with two complex factorsheight and fertility. communities, but even then comparisons with Presence, unlike the other two elements, hasentirely different situations using similartypes been cot nected significantly only with inbreed- of analysis would benecessary to determine the ing and during the first decade. it is noticeablebearing of such findingson the general problem here that eally appearance of goiter, like nodu- f,f endemic goiter. Is endemic goiter theadapta- larity,is negatively influenced by inbreeding. tion of man to iodine deficiency? Ourdata lead Sex and age, as such,are involved with all three us to suggest that subjects with certain types of elements, but we feel that their relevance should goiter are better adapted than others. Thefact be sought mainly through interactiontypes ofthat Indians with large goitersare taller and effects. have more children might bc in line with such In view of the complexity and number ofan interpretation, even if we are unable to in- factors to be considered acting together, it would terpret the mechanisms involved.

REFERENCES

I. ARTEAGA, A., J.BARZELATTO, E. COVARRUBIAS, L. T. MOYA, and A. 0. CLEMENTE. In Radioisotopes A. VALIENTE, E. ROSALES, C. MICHEL!, andC. in Tropical Medicine, Vienna, IAEA, 1962,p. 151. TORRES. Nutr. Bromatol Toxicol. 4:125, 1965. 8. COVARRUBIAS, E. BiolOgica (Chile), FaL. XXX- 2. BARZELATTO,J.,C. BECQUERS, C. STEVENSON, VII:62, 1965. E. COVARRUBIAS, A. GIANETTI, E. BOBADILLA,A. 9. COVARRUBIAS, E., and J.BARZELATTO. Third PARDO, H. DONOR), and A. ATRIA. ActaEndocr. Meeting PAHO Study Group on Endemic Goiter, 54:577, 1967. Puebla, Mexico, 1968. To be published. 3. BARZELATTO, J., C. BECQUERS, C. STEVENSON, 10. COVARRUBIAS, E., J. BARZELATTO, C. STEVENSON, A. GIANETTI, E. BOBADILLA, and A. PARDO.Minerva E. BoBADILLA, A. PARDO, and C. BECQUERS. Nature Nucleare 9:259, 1965. 205:1036, 1965. 4. BECQUERS, C.,J.BARZELATTO, C. STEVENSON, 11. EKpEctu, 0. L., A. DIMITRIADOU, and R.T. A. GIANETTI, A. PARDO, E. BOBADILLA, and M. DE FRASER. In C. Cassano and M. Andreoli (eds.), VISSCHER. Acta En doer. 54:591, 1967. C'urrent Topics in Thyroid Research. New York, 5. Bolumvi, S. F., and N. S. SCRIMSHAW. 13o1. Academic Press, 1965, p. 866. Ofic. Sanit. Panamer, Suplemento 3:157, 1959. 12. ERMANs, A. M., J.E. DUMONT, and P. A. 6. CALLEJAS, L.,J. GOMEz, R. ALMANZAR, and BASTENIE. 1. Clin. Endocr. 23:550, 1963. A. UCROS-CUELLAR. Rev. Soc. Colomb. Endocr. 4: 13. GAYTAN, E. Personal communication. 55, 1966. 14. GREENWALD, I. Texas Rep. Biol. Med. 17467, 7. CAMPOS, P. C., B. S. BALTASAR, N.GRABATO, 1959.

131

136 * 15. GRE ENWALD, I. Texas Rep. Biol. Med. 15:874, 32. PELTO L A, P. In C. Cassano and M. Andreoli 1957. (eds.), Current Topics in Thyroid Research. New 16. GREENWALD, I. In G. B. Astwood (ed.), Clini- York, Academic Press, 1965, p. 872. cal Endocrinology, I. New York, Grune & Stratton, 33. PERINETTI, H. et al. In Advances in Thyroid 1960, p.123. Research. London, Pergamon Press, 1961, p. 283. 17. FIERRo, R., L. DE G ROOT, M. PAREDES, and 34. PoDovn, J., and P. LANGER. Naturally Occur- W. PERAFIEL. Rev. Ecuator. Wed. Cienc. Biol. 5: ring Goitrogens and Thyroid Function. Bratislava, 15, 1967. Slovak Academy of Sciences, 1964. 18. FoLus, R. H., JR. Bol. Ofic. Sanit. Panamer. 35. ROCH E. M. J.Clin. Endocr. Metab. 19:1440, 60:28, 1966. 1959. 19. Fouls, R. H. Am. J. Clin.Nutr. 14:253, 1964. .3('. ROCHE, M., F. DE VENANZI, J. VE RA, E. COLL, 20. iodine Facts, I (217). London, Iodine Educa- M. SPINETTI.BE RTI, J. MENDEZFARTINES, A. GERARD, tional Bureau, 1940-1946. and J. FORE RO. J. Clin. Endocr. 17:99, 1957. 21. KELLY, F. C., and W. W. SN EDDE N. Bull. Wld. 37. ROCHE, M., H. PERI NETTI, and A. BARBE ITO. Hlth. Org. 44:27, 1960. 1. Clin. Endocr. 21:1009, 1961. 22. KIMBALL, 0. P. Arch. Int. Med. 107:290, 1961. 38. ROTHHAMMER, F., R. BLANCO, E. COVARRU- 23. KRuslus, F. E., and P. PELTOI.A. Acta Endocr. BIAS, and M. DIXON. Ztschr. f Morph. v Anthrop. 53:342, 1966. In press. 24. LOBO, L. C. G., J. FRIDMAN, D. ROSENTHAL, 39. SCRIMSHAW, N. S. Public Health Rep. 75:731, R. ULYSSEA , and S. FRANCO. J. Endocr. 22: 1960. 1182, 1962. 40. STACPOOLE, H. H. Bull. Wld. HIth. Org. 9: 25. LOBO, L. C. G., A. QUELCESALGADO, D. ROSEN 283, 1953. THAL, A. FREIRE-MAIA, and M. A. RE BELLO. VI 41. STANBURY, J. B., G. L. BROWNELL, D. S. R IGGS, Pan American Congress of Endocrinology. Excerpta H. PERINETTI.J.I TOIZ, and E. B. DEL CASTILLO. Medica, Internat. Congress Series 99, 1965. Endemic Goiter: The Adaptation of Man to Iodine 26. MA ISTERRE NA,J.A., E. TOVA R, A. CANCI Deficiency. Cambridge, Massachusetts, Harvard Uni- and 0. SERRANO. J. Clin. Endocr. 24:166, 1964. versity Press, 1954. 27.. MALAMOS, B., K. MIRAS,P.KOSTAMIS,J. MANTZOS, A. C. KRALIOS, G. RworouLos, N. ZE RE 42. TELLEZ, M., A. GIAN ETTI,E. COVARRUBIAS, FOS, and D. A. KOUTRAS. In C. Cassano and M. and J. BA 11 ZELATTO. Proc. Third Meeting PAHO Andreoli (eds.), Current Topics in Thyroid Re- Study Group on Endemic Goiter, Puebla, Mexico, search, New York, Academic Press, 1965, p. 851. 1968. To be publishcd. 28. MORTON, N. E. Cold Spring Harbor Symp. 43. UCROS CUELLAR, A. Unidia 7, Suplemento 1:1, Quant. Biol. 29:69, 1964. 1960. 29. NAGEL, R., d R. ETCHEVERR Y. Nature 197: 44. WAHNER, H. W., E. GAYTAN, and P. CORREA. 187, 1963. I. Clin. Endocr. 26:279, 1966. 30. NAGEL, R., R. ETCHEVERRY, C. GUZMA N, A. 45. WAY N E. E. J., D. A. KourRns, and W. D. HILLE, J. BARZELATTO, and E. COVARRUBIAS. Rev. ALEXANDER. Clinical Aspects of Iodine Metabolism. Med. Chile 90:616, 1962. Philadelphia, F. A. Davis, 1964, p. 48. 31. PAR RA4IM ENEZ, N., P. RODhIGUEZGARC IA, M. 46. WEINSTEIN, E. D., J. V. NEEL, and F. M. ROCHE, and K. GAEDE. J. Clin. Endocr. 22:754, 1962. SALZA N O. Amer. J. Hum. Genet. 19:532, 1967.

132 1:37 DISCUSSION

J. D. Niswander

The speakers today have each consideredone diabetes mellitus. The rank order for the hos- special diease problem of the Indian. I shouldpitals changes very little if, instead of the total like to Jiscuss three of these in relationto one number of discharges, the total beneficiarypopu- another. I will discuss diabetes mellitus, gall-lation of each hospital is usedas the denomi- bladder disease, and nutrition in thecontext of nator. I believe there is enough uniformity be- the over-all variability in frequency of diabetes tween Public Health Service Indian Hospitals among various Indian tribes in the United States. to make these data reasonably adequate for rela Specific studies have reportedan unusuallytive frequency comparisons, althoughone cer- trequency of diabetes among several North tainly would not want to attributea great deal Amei ican triheswith the studies beingcon- of precision to them. ducted among the Pima, reported by Dr. Miller, The map shows the location of the various being the foremost example. Thereare, how-Public Health Service Hospitals throughout the ever, very few data on the question of juzt how United States and their grouping intoseven prevalent diabetesisthroughout thevarious administrative areas. The eighth area (Portland) American Indian groups or whether it ismore delivers servicP throughcontract with local fa- or less uniquely confined to particular tribes.cilities and not directly through Public Health Certairly such a hypothesis as that put forwar:' Service Hospitals. by Dr. Neelthat diabetesmay represent the The mean frequency for the 46 Indian hos- effects of changing dietarypatterns on genomes pitals is 2.2 per cent. Thiscompares with figures previously adapted tt-,a "feast or famine" situa- of 1 to 1.5 per cent for similar data for several tionwould. uggestthat a high frequency of surveys of general non-Indian hospitals through- diabetes might be expectedamong populations out the United States representing over 26,000,- who have been both recently and rapidlytrans- 000 hospital admissions. formed from this primitive patternto a more The table groups the hospitals by the relative sedentary and overfed condition. frequency of admission for diabetes and indi- The only source of information thatI am cates the major tribes served by each. As can be aware of on this subject, although the data are seen, the range expressed by the mean frequency less than :deal, is the records of the U.S. Public of 2,2 per cent is very wide, with roughlya Health Service, Division of Indian Health Hos-twentyfold difference between the lowest and pitals. Accordingly, I have obtained tabulationsthe highest. The table also shows the relative of discharge diagnoses fiorn these hospitals forfrequency of admission for gallbladder disease, the four-yea priod 1963-1967. Thisrepresents which very closely pa.-allels that for diabetes. 1.1- approximately 250,000 discharges. I have tabu-fact, the rank correlation is slightly higher than latrd the proportion of discharges from each .5 and extremely significant. There is, however, hospital in which the primary cr.agnosiswas a considerably narrower range of variability than

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i4111110111 WM* lassosaisa Unisys Adontniesod two Os Oliakmos Wins Heil* ANA Whelk for diabetes. It can be seen that thereis anvarious Alaskan Indiansand also the Atha- equally striking association between diabetes and bascan-speaking Navajo and the closely related average birth weight. Here again the regression Apaches. I think it is noteworthy that the Alas- of birth weight on frequency of diabetes, bykan natives and the Navajo are probably the hospital, is highly significant. least acculturated of all the Indians remaining Specificallyin regard to diabetes there are in the United States. several pints of interest. First of all, in almost The possibility that their low frequency of two thirds of the Indian hospitals diabetes seems diabetes is due to the inherent genetic make-up to represent a significant! y greater proportion of of these groups, indepent:ent of environmental the patient population tl an in non-Indian hos- factors, cannot be excluded. Likewise, it could pitals. Most of the major linguistic groups arebe argued that birth weight and diabetes are represented in the high and intermediate dia-related by virtue of both being tne result of a betes categories with the exception of the Aleut- particular genetic constitution. Alternatively, the Eskimo group and possibly the Athabascans. birth-weight gradient might be considered a sec- The highest diabetes group is composed of sev-ondary effect of maternal diabetes. It seems to eral different tribes, but most conspicuous are ine, however, that the relationship between gall- the so-called five civilized tribes from the south- bladder disease, birth weight, and diabetes, when eastern portion of the country. All had relatively viewed together with the admittedly limited highly developed agriculture prior to European information on cultural variation, suggests at contact. Also notable among this group are sev- least one other possibility: the operation of com- eral of the southwestern agricultural tribes in- mon environmental factors, presumably dietary. cluding the Pima. It might be suspected thatTo speculate a bit more than is perhaps war- these early agriculturists would be the ones mostranted by the data, I would suggest that virtually susceptible to acculturation and hence wouldall Indians may be of a genetic constitution have undergone the most significant changes in especially susceptible to diabetes and its expres- dietary patterns. sion is the result of dietary and nutritional fac- The low-chabetes group is notable for includ-tors associated with prtsent-day reservation life. ing all of Alaskathe Aleuts, Eskimos, and In this regard, it is unfortunate that there are

134 '

Frequency of diabetes mellitus and gallbladder disease andmean birth weight in Public Health Service, Division of Indian Health Hospitals

DIABETES GALLBLADDER ' N 1 EAN MAJOR (To OF ALL (% OF ALL BIRTH HOSPITALS TRIBES ADMISSIONS) ADMISSIONS WEIGH,' I

Window Rock Area (6 hospitalsNew Mexico, Navajo Arizona)

Beicourt, North Dakota Chippewa

Mescalero, New Mexico; White River, Arizona Apache 0-1 1.9 6.(8 Keams Canyon, Arizona Navajo, Hopi Alaska (7 hospitals) Aleut, Eskimos "Alaskan Indians"

Montana (3 hospitals) Blackfoot, Crow Cheyenne, Gros Ventres Assiniboine, Cree Cass Lakc, Minnesota; Rcd Lake, Minnesota Chippewa North Dakota, South Dakota, Nebraska "Sioux" 2-3 2.3 6.88 (6 hospitals) Santa Fe, New Mexico; Zuni, New Mexico "Pueblo," Zuni San Carlos, Arizona Apache Clinton, Oklahoma; Lawton, Oklahoma Cheyenne, Arapaho, Kiowa, Comanche, Caddo Sells, Arizona Papago

Cherokee, North Carolina; Tahlequah, Cherokee Oklahoma Claremore, Oklahoma; Pawnee Oklahoma; Cherokee, Creek, Philadelphia, Mississippi Ponca, Choctaw, Seminole Talihina, Oklahoma Choctaw, Chickasaw, 4-10 2.6 7.05 Cherokee, Creek Wagner, South Dakota Sioux Sacaton, Arizona Pima, Papag,i Shurz, Nevada; Owyhee, Nevada Paiute, Shoshone Winterhaven, California; Parker, Artzona Cocopa, Mohave, Chemehuevi Based on 257,320 discharge diagnoses, 1963-1967. t Based on 27,288 births.

135 ..,140 no very adequate data avvilable on relativeof increased mortality and morbidity figures. obesity between tribes or on the increasein Difficult related questions, not yet answered,con- obesity that may have occurred in recentyears. cern the proportion of genes potentially capable There is, however, one study that reportsap- of prccoking disease that have passed to the proximately a six-kilogram increase in weight Indians from the other races with which they among the Navajo, with no increase in stature,are in the process of mixing. overatwenty-yearperiod.The reservation Ail the specific health problems mentioned in Navajo, however, are still considered relatively thissession are characterized bya complex lean in body build, without the great tendency causality involving multiple genetic, environ- toward obesity that is seen insome of the othermental, anti cultural factors. The genetics ofa tribes. It would be of considerable interestto disease like diabetes is a complicated subject, in follow Navajos whu havenow moved to the which no precision as to the part played bygenes major urban centers andcompare them both for has thus far been reached; moreover, diabetes obesity and for serum glucose levels with their may represent a genetically heterogeneous group presumably less acculturated counterparts whoof disorders.' A similarsituation existsfor have remained on reservations. endemic goiter. In the case of malaria, in which My purpose in this discussion has been pri- increased resistence is foundamong individuals marily to point up the need not only for detailedheterozygous for hemoglobin S, other stillun- studies of special diseases in specificgroups but known genetic factors, probably polygenic,may alsofor data gi ing comprehensivecoverage be involved.2 over many Indian populations. Certainly, studies What I have said in relation to genetic etiology of human adaptation and attempts to understand is paralleled in the field of environmentalcauses. the interactions betweengenotype and changing Thus, the variability of allthese diseasesis cultural patterns demand considerable knowl-caused by an interaction of factors that is prob- edge of the variation which may exist within ably unique in each population, andno geneti- and between various human ppulations. cism or environmentalism will be of real value If my speculations prove to becorrect, diabetes to an over-all comprehension of the problems.2 may become an increasingly important medical Among the specific questiors raised to-day. problem as the acculturation of the American should like to point out the associations between Indian increases. Further, such considerations gene distributions and such phenomena as the would seem important in terms of general plan- relationshipsuggestedfor Pewenche ning for dealing with the health problems of Indians between nodul.ar goiter and lcwer phenylthio- emerging primitive groups. carbamide thresholds in the group of "tasters" Acknowledgment of this substance. Similar results may appear from the studies of diabetes and gallbladder I am grateful to Mr. Abraham Ranofsky.Program disease among the Pima Indians, in relation to Analysis awl Statistics Branch. Division of Indian Health, blood groups. Whether or not such associations for furnishing thespecial tabulations on diabetes and gallbladder disease. are real is to be found in the future, but they may constitute r.arting points for the analysis of polygenic systems in man. Another interesting Edmundo Covorrublos question here concerns how the epidemiology of The presentationswe have heard this after- noon make it clear that the American Indians ' Rimoin. D. L. Genetics of diabetes mdlitus. Diabetes received from the conquisatadors 16:346-351. 1967. a rather heavy 2 Allison, A. C. Genetic factors in resistance to malaria. burden of evils: biologicalagents of disease and Ann. N.Y. Acad. Sci. 91:i10-729, 1961. the pressure ofa different culture, stresses to 8 Thoday. J. M. Geneticism and environmentalism. In I. E. Mead (ed.), Biological Aspects of Social Problems. which they had to become adaptedat the toll Edinburgh and London, Oliver and Boyd. 1965.

136 a disease would be influenced by the distribution ecosystem for many centuries have evolved and of genes, which vary notably from tribe to tribe, adapted their own practice of infant feeding, as has been shown for blood groups in some ofand thatz,f we rush in with our own outside the papers submitted today. ideas without taking these into careful consid- The mental diseases present a set of problems eration we may do very great harm. even more formidable in view of the difficulties Second, there are lessons tobe learned, of diagnosis due to cultural differences. Onebelieve, from different groups of people and effort in this direction was recently made in different traditional systems. I think thatwe in Chile: a new structural theory of psychoses led the so-called Western world havea considerable to transcultural definitions that made it possible body of dogma in our own infant feeding that to distinguish corriderabk differences in the fre- in fact might very well be re-examined in the quencies of the proposed types of psychoses be- light of practice elsewhere. For example,as tween Mapuche Indian patients and the controls.4 Dr. Behar pointed out, in most of pre-conquis- Finally, I feel tempted to generalize to other tador South America and also in Polynesia systems the words of Book5 concerning theanimal milk was not traditionally drunk. Yet etiology of mentalillness:".. my general people there reared childien and fed themper- medical and biological background together with fectly adequately without it. Thereare messages my fragmentary knowledge of psychiatricre- for us from these differe-,tgroups. search work has led to the tentative conclusion Another important aspect of Dr. Behar'spaper that any unitary explanation ofthe etiology is the fact that disruption ofa culture can very even of what is now considered a special type frequently have itsmost sensitive and devas- of mental illness, or a clinical entity, will not tating effect on the age group that ismost pre- contrib,ite to the advancement of psychiatric cariously poised physiologically. That, ofcourse, research." is the young child--the weanling, who is ina To sum up, biomedical research among the process, if you like, of biocultural transition. American Indians is apt to disch new facts When we consider the Indian communities in that may be of value both for srecific publicthe Americas, we should, I think, bear in mind health policies and for a general understanding what has happened elsewhere in the world. For of the diseases studied. example, the Hadza hunters of northern Tan- zania were surveyed some six years ago and General Discussion were found to have a pattern of infant feeding well adapted to their communitybreast feed- Mille: I should like to congratulate Dr. Mar on his excellent paper. It seems to me that, late ing, bone marrow, pre-chewed meat, and so as the hour is, it woald be wrong to let this goforth. Five years later another surveyWaS car- by without emphasizing its importance. As those ried out, and the pattern of infant feeding and of us here who are working in the field of child. childhood malnutrition had changed completely. hood malnutrition will agree, the lessons to beIn the first survey there was very little protein- learned from his paperare of great importance. calorie malnutrition. In the second itwas very First, I think that we all haveto realize that common. all groups that have been living ina certain What practical messages are there ta be gained Ivoin Dr. Mar's paper? I think thew are these: 4 Mufioz, L.,J. Marconi, J. Horwitz, and P. Naveil- We. should realize thatas well as the rapid !An. Cross-cultural definitions applied t' the study of spreid of infections due to viruses, suchas functional psychoses in Chilean Mapuches, Brit.J. l'sy- arial. 112:1205-1215, 1966. measles, or to bacteria, such as tuberculosis, the 136iik, J. A. Genetical etiology in mental illness.In spread of inappropriate non-adapted ideascon- Causes of Mental Disorders: A Review of Epidernio- lopical Knowledge. Ncw York, Milbank Memorial Fund, cerning inFant feeding can have a very consid. 1961. erable effect on local child health andon the

137 i44.2 local pattern of childhood malnutrition. How malaria and died from it and never came to can we try to prevent this? I think that very terms with it as the African does. You do find probably, before changes are allowed to occur endemic malaria in a few other groupsin too easily in the dietary habits of Americannortheastern India, for example. But itis so Indian groups, careful thought should be given localized as to suggest a racial factor in the to whc has been responsible for these changes mechanism of holoendemic malaria quite apart in the past. from length of exposure. Many of these inappropriate imported changes Water low: That is what I am getting at are iatrogenic, using the term in its wide sense. whether there is a racial factor even with the Ithink, therefore, that in dealing with the Amerindian. American Indian groups that have been dis- Gig lioli: I think so. cussed here the infant feeding practices taught Roche: What has been said about care with by the health services should be most carefully the food brought into these areas induces me to adapted to the local circumstances and should say a few words about the moral issues involved not merely be cultural imports from elsewhere. in studying Innians. These are people who are Furthermore, any imports of foods into theseat our mercy both morally and physically. The communities through, forinstance, food aid Declaration of Helsinki on research on human programs undertaken by international agencies, beings states that the informed assent of the national services, and voluntary organizations, people on whom one is doing research should be including missions, should be very closely looked obtained. Thi:3 includes research on their internal at for whether their hoped-for immediate bene-milieu, not merely their external milieu. The fits will in fact yield ill effects in the near future. persons responsible for the statement were not, Lastly, "commerciogenic" malnutrition has to Ibelieve, thinking about this type of study be mentioned here. It is probably as dangerous because it is practically impossible, as everybody to introduce a commercial milk company, a com- will agree. to get really informed assent from pany selling milk and promoting feeding bot-the Indians. Thic means that the burden is on tles, to a community of people who are success- the investigator, who must exert the utmost fully breast-feeding as it is to introduce ronsles restraint in his investigation to be sure that his or tuoerculosis. activity is in no way harmful to :he populations Water low. May I ask Dr. Giglio li one ques- he is studying. tion?Isit your belief that malaria did not Cohen: I want to return to the theme of how originally exist in the Amazon forest country? these factors might emerge without a basic Was it one of the things brought in along with genetic alteration.I raise the question whethcr others? we are not dealing with the well-known modula- Gig Both My paper touches on this question. tioneffectsthatderepressors andrepressors Not that I have any competence to discuss it. might have on the transcription and translation I mean, 1.is a moot question whether malaria phenomena. It is now, I think, reasonably well pre-dated the Conquest or was introduced. documented that the inliuence of a variety of think the consensus now is thatit was intro- hormones can lead to the expression of enzyme duced and that this is probably the reason why activity that is not seen at a lower level of certain the Ainerindian reacts so much more actively hormones. to malaria infection than the Negro, who has As a matter of fact, many hormonal patterns been exposed to it lor ages. are now being explained in molecular biological We have no data togo by. Consider the Medi- terms of messenger RNA and all that goes with terranean populations, Nhich were exposed to it. This must mean that there is an interplay by malaria for centuries. Certainly therewas noa variety of factors on the genetic potential of such total endemic malaria. People suffered from an individual to give different degrees of quan-

138 titative expression. I am not sure there would be that a variety of nutritional states will elicita qualitative changes. great pattern of enzymatic changes that are not As I listened to many of these presentations, due to an instantaneous transition of thegenes. diabetes in Indians and so on, I thought it quite We know this. Just a simple high carbohydrate conceivable that this could be a quantitativeintake as against a low, high nitrogen intake variation of one or another enzyme pattern thatas against low, will cause changes of level of would lead to a metabolic anomaly of the kind I high orders of magnitude. Thesecan be feed- have referred to. Considering the influence of backs of various kinds, but many have hormonal diet on a number of these circumstances, I think factors that are known to have their initiating the questionstillremains whether the basic effect on the transcription phase in which the genetic pattern is being significantly altered in gene may be repressed for a particular level of the traditional sense or whether the change is quantitative operation. taking place at the modulating level,as it were. I am only suggesting that beforewe get too Moderator: A discussion of this would takeus involvedingenetic markers as an absolute rather far afield. I am not vire whetheryou are known relationship, which is clearly thecase in using the term modulation in thesense used by many diseases that have been defined in these the bacterial geneticist or whetheryou are re- terms, we give some thought to the possibility ferringto the kind ofnature-nurtureinter- that in many of these adaptive phenomenawe actions for which we have come to havegreat are dealing with something quite different. respect :n man. The idea that in fact the genetic I hope that at some time in the future there readout might be different for ourselves than may be a chance to talk at this level as a follow- for, say, the Indians simply because of theen- up to our discussion today. This is where the vironmental differenceis a challenging idea.new molecular biology may perhaps change My first reaction is that the homeostatic mecha- some of our concepts. nisms of man are such as to buffer this readout Moderaton The final item on the agenda is quite a bit. On the other hand, perhapswe labeled"Commentary andSummarization." should devote some thought to trying to put This will be extremely brief. One of the marks that idea to some kind of critical test, andcer- of a good meeting is the extent to which old tainly one opportunity is in transitional popula- ideasarerescrutinized and questioned, and tions. For instance, if it can be documented that from beginning to tad of this session we have the polymorphisn.c, which area readout forbeen questioning some of our old ideas, begin- proteins, alter ;1.equency more rapidly in a ning with the matter of the origins of the population in cultural transition thancan pos- American Indians and ending with details of sibly be ascribed to genetic selection, this woulddisease susceptibility and resistance. The session be hard evidence that the readout is different. bas been very profitable, and it ismy privilege Thac is the kind of eidence that I think itto thank PAHO for making this gathering would take to satisfy most ofus. possible, both financially and in terms of facili- Cohen: Not to prolong this, itis very clear ties.

139 144 v. V.. 't , ApFendix

TABULATIONS OF PHENOTYPE AND GENE FREQUENCIES FOR 11 DIFFERENT GENETIC SYSTEMS STUDIED IN THE AMERICAN INDIAN'

R. H. Post, J. V. Neel, and W. J. Scholl

With the impetus given to further genetic In preparing the tabulations, a recurrent ques- studies of the American Indian by the forth- tion in the face of an unusual finding has been coming International Biological Program, itis whether the cause might be racial admixture probable that the next decade will witness a (i.e., miscegenation). Furthermore, as we have sharp increase in investigations of this kind.pointed out elsewhere (5), the Indian presents The literature is already voluminous and scot- an unusual challenge to thoseinterestedin tered;it seemed that a compilation of thwe recent developments in genetic taxonomy (see studies which deal with the major genetic poly- I, 2), but attempts to apply these developments morphisms would prove useful in assisting in may be vitiated by substantial racial admixture. vestigators bothin planning and facilitatingAccordingly, we have made an effort here to studies and in comparing their findings with classify each sample in one of three ways: (1) those of others. Accordingly. in Tables 1to 11 probablyessentially no admixture,(2)non- tabulations are presented for the following sys-Indian contribution to the gene pool probably tems: ABO, MN, Rh, Kidd, Duffy, Diego, P, less than 5 per cent, and (3) non-Indian con- Lewis,transferrins,haptoglobins,and Gm. tributiorr to the gene pool probably greater than These are all systems exhibiting marked poly- 5 per cent. morphism. W. have not included systems, such As every student of the problem knows, the astheglucose-6-phosphate dehydrogenase or classification of certain samples into one of these hemoglobin system, that are polymorphicin three categories is difficult, and we are sure not some populations but esselitially nonvariant in only that there are grounds for issue with somc the Indian. Nor have we included some systemsof our decisions, but also that some of these that while probably polymorphic arc still inade- decisionsareincorrect and will have to be quately defined either genetically or phenotypi- modified. In this connection, it should be em- cally, such as sensitivity of taste to phenylthio- phasized that the decision applies only to the carbamide, distribution of hair on phalanges, particular sample that was examined, and not tongue-rolling, and so on. An effort has been necessarily to the entire group or tribe. Each made to survey all publications prior to Novem- classification has been based on the history of ber 1967; these are cited in the Bibliography. the tribe, the opinion of competent observers, the

I Work supportedinpart by US. Atomic Energy results of the genetic studies themselves, and, Commission Contract AT( 11-1)-1952. where given, the author's statement.

ifof 141 145 The results of typings for the ABO systemtribal organization has long since disappeared, have been thought to provide especially critical these names may be confusing and occasionally material for ckcisions regarding admixture. Wemeaningless. Thus thereare two samples of accept the working hypothesis that IB and IA-Mayans so called by the authors,an unsatisfac- are not found among American Indians andtory name since several other samples are tabu- that IA, does not occur in South America. How- lated of Mayan-speaking IndiansChol, Kek- e-ier, we note that in the few South American chi, Mam, Pocomam, Quiche, and others.Five samples where P3 and 01 dooccur, the ratio ofsamples are designated Aymara,two Quechua, the frequency of the former to that of the latter and one Chibchan, each of which isa large tends to be higher th.n in western Europe andlinguistic stock including several distinct cultures Africa, the presumed origin of thesegenes. The and several millions ofpersons. Three linguistic possibility must be recognized that IBwas pres- stocks of simpler culwreare similarly employed ent in South America in pre-Colombinn times asnamesof samplesArawak, Carib, and (see also 6), perhaps introduced by trans-Pacific Guarani. From four to ten samplesare desig- contacts. nated by tribal names that fall under each of Additional genetic grounds for decisionsre- these stocks. garding sampk purity are the presence of the The estimates of gene frequencies inthe following antigens, wherever tested for:Kell, tables that follow were computedon an IBM Lewis(a),I3errian(Be), Henshaw (He), 1130 Model, using programs written in the De- Lutheran (Lu"), Miltenberger (Mia), Sutterpartment of Human Genetics. The ABO pro- (Js"), V of the Rh system, Verweyst (Vw) ofgram has been publishesi (4); the other pro- the MN system, and Wright (Wr). All of these grams are available on iequest. The "goodness are thought to be absent in unmixed Indians, of fit" of the observed phenotype frequencies and we have not included tabulations forany withthoseexpected under Hardy-Weinberg of them. References establishing the virtualor equilibrium and with the calculatedgene fre- complete absence of these traits in samples ofquencies has been tested by chi square. The latter unmixed Indians are listed in Table 12. Weare rot given, but the corresponding probabili- have felt, on the other hand, andcontrary toties when significant are indicated byone of some opinions, that the occurrence of Rh-nega- three letters in the column to the far left of tive individuals (d) need not necessarily indicate each table, as follows: A-- 0.05>P>0.01; B- admixture (see 3), although high frequencies0.01>P>0 001; C 0.001>P. The validity of may be of significance. applying such X2 tests to small sample:t in which In our opinion, many types of criticalcom- the mean coefficient of relationship is probably parisons are impossible when there ismore than not negligible is dubious, but we have followed 5 per cent admixture in the sample. Only the convention in this respect. data on samples with no admixture,or less than Names of countries are abbreviated by three 5 per cent admixture, have been incorpoiatedletters and followed by brief indication of into Tables 1to 11, with samples of the first locality, such as that of a stateor province, class designated by asterisks. The referencesto River (R), Mountain (M), Lake (L),or region samples with more than 5per cent admixture (N for north, and so on). Exceptionsare made are given in Table 13, which lists the samplesfor French Guiana and for Guyana and Suri- in alphabetic order. nam (formerly British and Dutch Guiana, re- The names of tribes andgroups are those 3 spectively), for which no locality is added, since 4 given by authors, to whichan arabic numeral their areas are small. has been added incases of two or more samples In several instances an author has published of the same name. Among the civilized popula-data on two or more "subtribes"as well as the tions of Middle and South America, wherepooled data of the "tribe." The grounds for

142 recognizing subtribes are seldom clear; theya few samples of mixtures of several tribes or sometimes appear to represent different villages groups that appear to have less than 5 per of the same tribe. In all case!), we have retained cent non-Indian ancestry. Such data are not the author's designation but listed his data under tabulated here, but thereferences are listed the main tribalunit, with a cross-referenceapart (Table 14) in alphabetical order of the under subtribe. Thus, the data on the Arecuna tribes, together with notes on geographic local- subtribe of the Pemons will be found underity, the names of the several tribes included in "Pemon-Arecuna" with a cross-reference under the mixture, and our estimate of the amount of "Arecuna." non-Indian ancestry of the pooled sample. Genetic marker data have ken published on

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151 A, 1 153 CODE General Panama PAN Paraguay PAR 1. An asterisk following the name of a tribe or group Peru. PRU indicates essentially no Negro or Caucasian admixture. Salvador SAL 2. When two or more samples are presented from a United States USA tribe or group of the same name (whether from the Venezuela VEN same or different authors), each sample is arbitrarily assigned a serial number. This appears in column 14 Colonies unless the sample is pooled from severalareas or sub- tribes, in which case "P" appears in column 14 prrceded British Honduras BRH by the serial number in column 13. (Subsamplesarc French Guiana FRG Guyana presentedseparately,also,wheneveravailable.)"P" GUY stands for "pooled." Surinam SUR

Abbreviationsofnations andcolonies and Other abbreviations other abbreviations North East Nations South; Saint; Argentina ARG Sierra Bolivia BOL West Brazil BRZ Mountain(s) M(TS) Canada CAN Another; Others ANO Chile CHL River Colombia COL Lake Costa Rica COS Delta Ecuador ECD Island (s) I(S) Guatemala GUA Pooled data, Honduras HON from several Nicaragua samples TABLE 1A. THE Alio A2. e. Ale 0 GENE pAroueNcres. As OLTEAmmeo wtt., AN71.40 ANT1A1. AND ANIP4 SENA. TRIBE OR GROUP NATION REGION REF SAMPLE SIZE 0 Al PHENOTYPES A2 8 Alt A29 Al GENE FRovENcits A2 et 0 P APACHEAPACHE*ALACALUPAGUARUNA*ACAWA10* 2 1 9 USACHLPROGUY NEWMAGELLANARIZR MEXMARANON 0900421641.68130 179151 4490 105151 4190 74 30 0 0 00 0 0.2340.0350.000 0.000 0.0000.000 0.7660.9651.000 ATHAPASCANATACAMENOS"MECUMARAWAK* 1 USACHLSEESUR ANTOPAGSTALASKA PERON 062164060 108 788088 73748759 48 430 001 042 002 0 0.0190.0000.266 0.0000.0000.006 0.029000190.006 0.9990.9940.7011 AYMARA*mla/KOMA 314 ANO* POLPAUPRUBRZ 3PUNO AREASS CATRINA P 169168028233 903122 9358 491121 9358 070 10 201 0 0 0.0000.026 0.0000.00000000 040000.0000.004 1.0000.9960.974 CAINGANGCARECAR*BRIM/.BARI* BRZCOSVEN RUJARPAS GSALITREW SULZULIA 231177132 126 62SO 126 6250 0 10 0 00 0 0.00006007 0.0000.003 0.000040000.002 0.9871.00010000 CAMPA*CAMARACOTOcAlconourtcAlmstoucLAcAlcommet i32 SEEGUA SUMPANGOSOLOLAI. ALTI1LMPEMON 268170171 137150341 9 112142334 9 2044 00 3042 0 0 0.0070.0000.0130.006 0.0000.0000.001 0.0110.0000.0130.003 0.9811.0000.9730.989 CHEROKEE*CHAHULAACHAMA*CHACORO*CAYAPA* 2 PEABOLLCDPRU CHIAPASR MADIDIYATACAYAPASumeAme 024169167168 244 63301489 240 63301489 00 0 04 00 00 0.0000.000 0.000 0.0000.008 1.0000.9921.000 CUM*CUIVA*COLORADO*CHOCO* 2 2 PANWENLCDUSA APURESDARIENN DOMINGOBLASCARLINA 15 178134167209 82368078 8236SO75 0 01 002 00 00 0.0000.000 0.0000.006 0.0000.013 1.0000.980100001.000 GUARANI*GuAmtso.GOAJIRODIEGUENO 32 PRZWENVENUSA APUREWCAL ZULU,. 134128202 109119388 '18 1071.14388 56 13 00 120 0 0 0.0050.0130.0090.000 0.0000.000 0.0050.00$0.0090.000 009900.9790.9821.000 HUASTEC*HOPIGUAYO*GUAYM1* AND 1TEMA MEXUSASEEPAN VERANEWN WS COASTCATARINmuCRUZWARAO 024042233178 123240 5234 115240 5234 030 09 0 0 00 0.0120.000 0.021040000.000 0.0000.000 0.9661.0001.000 KARAPALOJIVARO*1SCONAHUA*!NAPA*ICA 2ANO*2 2 BRZPROECDSEECOL MAARAP/COSR UCAYALI SGROSSO REV YUPA 167168131 233114 14 233110 14 01 0 01 00 00 0.0000.0000.013 0.00000000 0.0000.004 1.0001.0000.98216000 LACANDON*KEKCHI*KEKCH1 1 2 / MEXORHGUA CHIAPAS'mowCOPAN 170176171201 119162 337$ 117154 3273 2014 010 020 001 0 0.0150.0080.0160.000 0.000040000.003 0.0000.009 0.9920.9721.000 MAKIRITARE*MACUSHI*MAC0/7A*LACANDON* 12 WENGUYSEEMEX S BOLIVARENIAPASYUPA 130171 119 8661 119 elA61 0 0 0 0 0 0.00000000 0.000 0.000 1.0000.9851.000 mAR1COPA*MAPUCHEMANHAP 2 t 2 USACHLGUAGUA ARIZLONOUIMAYHUEHUETGOSAN JUAN 042163170171 141116 24 130112 22 92o3 0o1 01o 00o 00o 0.0320.0430.0130.000 0.0040.0000.000 0.0040.000 0.9190.9570.9821.000 MOJAVE* USA ARIZ 042 117 3$ 109 38 07 0 10 0 0 0.0300.000 0.0000.000 0.0040.000 0.96916000 I OTOMI*NAVAJONASKAP/ 42 MEXUSACAN QuenecHIDALGOARIZAR/2 042024063033 152237106 81 101173 7661 506145 5 03 01 0 00 0.0310.1160.1810.241 0.0000.007 0.000040000.003 0.9690.8540.7590.815 PARIAI*PANAAEPPEKIN* CAMARACOTO*ARECUNA* 3P VENYENSEEveN LAw mouvAlit SABANAYUPA 124130124 109267 7033 109267 7033 0 0 0 0 00 0.0000.000 0.000 0.000 1.0001.000 P/A0*PIMA*PIAROA* 1 TAUREPANs USAYEAVENPAU RARIZLA UNMAN,' APUAESARANA 042134124168 489111 90 395111 90 93 0 0 0 01 010 0.0000.0000.101 0.000 0.0000.001 0.8981.0001.000 RAMAQUICHEQUECHUA 121 ECDNICGUAECD RAMASCENTRALNORTHERN CENTRAL CAT 175170167 203372732 36 354188217 37 13 51 001 4902 00 0 0.0130.0330.01$0.011 0.0000.0010.002 0.0000.00504022 0.9870.9620.9750.964 C SHIP1B0*SNAOARUABECOYASANEMA* 1 PAUSEEECD ACUYARENO UCAYALIYANOMAMAYUPA 166107 142 AA 141 48 00 00 01 0 0 0.000 0.000 0.0040.0000.000 0.9961.000 'VICUNA*TERRAPAS*TAUREPANSUMO*S/R/ONO* COSSEENICROL ER COASTMACHUPOTERRARAPEMON 168177175169 122101 4027 122103 4027 0 0 0 0 0 0.000 0.000 0.000 1.000 Yumemo.TOTONAC 21 cotMEXPRU CHIAPASCOCUYVERAR AMAZON CRUZNTS 170131024 111100150 45 110142 9841 15 0 0123 0 0 0.0050.0340.017 0.0000.000 0.0110.0000.0050.010 0.9950.9900.9540.973 bro. UTEuyr7207271*TZELTAL*TZOTZIL* 21 2 1 USAusAMEX UTAHCHIAPAS 161163170171 104118 9180 102135 6979 2301 0 02 0 0 0.0100.0110.0000.006 0.000 0.0000.011 6.9900.9890.9030.994 WARAO*WAP15HANAMAMA* GUAYOm YENVENGUYSEE ORINOCO DYANOMAMA 120134 119127 7281 127119 7281 0 0 0 00 00 0.0000.000 0.000 0.00004000 1.0001.000 VANOMAMA.YAGUA*XAVAATE* WIMIKINA* 10P 3P vemPAUeluVEN ROR/NOCO AMAZONsMA sottvAN GROSSO 0 014168091120 572539 9 572539 9 0 0 00 00 0 0.000 0.000 0.0000.00C 1.000 YUMA.VARURO* WA1CA*SANEMA40 USAVENYEN ARIZAPURES BOLIVAR 042123134129 141182102156 167102141156 14 0 0 010 00 0 0.0390.000 0.000 0.0030.000 0.9881.000 YUPA. PARIQI9MACOITAIISMAPARUmIRAPAA 2 39 VENVEN E ZULIAZUL/A2ULIA 126126130 138176 7424ys 138176 247476 00 00 0 0 0 0.000 0.000 0.000 1.000

;0 " 04r, 4.0? '-e.00 4O 044 TAGLE 118.b NARD,WEINSEROTME EQUILIORIUN A. D AND 0 NOTEDRENE FREOUENCIESIN THE COLUMN AS HEADEDDETERMINED P IS GTIN ANTP"A EACN CASEAND ANTI'MOCCASIONEDLARSELY SERA* SY THE OCCURRENCE TOE SIGNIFICANT DEPARTURE FROM OF ONE ON NONETR/RE TYPE OR GROUPAS PERSONS IN A SAMPLE WHERE EXPECTATION FOR THIS EVENT IS VERY LON. 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Sti 1444 IV 04 4 11.0040 TANA sin. Comumottems IntIN CALCULATINSow eat IINDUANCINS cmoomosome AS OCINAMINNO moutmens WITHANI ASPOUR siscaten ANTI.SENA MA INATI.CePAWS SA. ANTI.40LITTLE C. ANTI*0. AMD Aneviet. ANTIC + + +... + PHENOTYPES . .+ + ... + + . e GENE FREQUENCIES e -148.1431/4. TR/RE OR GROUP RATION REGION REP SAmPti SIZE AkTI'EANT104411.c *- *+ + ** * ... * * * * * *+ 4. + EDC tDC ED t0 1d C aeC de E it4e 11"7,7 T CMAPULAPCAINGANGAPACHE* 1 mex382USA R GCHIAPASARIZ SUL 024231042 166178 57 A0 o0 0 196447 22 $ 1 13 00 0 0 11 72 40 64 060680.0510.009 0.2830.5920.439 0.345u0:070.390 0.0690.1010.161 0.000 0.202040000000 0.000 0.0440.000 404.12,10*8VERILLONDIEGUEMOCHIPPEWA 2 VENUSAFRGUSA MMINN ZULIACAL 193114202139 1!8161 3658 o0 0o 0 44212816 1312 01 10 0/ 0 00 10 052 3919:di 5 21ii 05 0.0000.1650.05806000 0.5930.7650.6620.313 0.3090.0690.2450.357 0.0900.0000.200 0.0000.129 080000.000 0.00006000 0.0400.0000.0140.5e. muAsnee.MOP1GUAYORIRAPA* 6000 11TEWA HEX682SEEUSA MAVERA14* GROSSO CRUZWARAOYUPA 201024042 106 7350 0 0 0 221220 23 9I 21 0 0 1 01 00 430 3223 7 35 69 0.000 084030.130 0.3160.233 00430.436 0.1410.000 0.00006000 0.0040.000 0400Napo MAPNACO/TM*KALAPALO 144002 40 DOASEE YUPA 262 235 2 0 1 63 12 3 1 7 0 17 94 II 33 0.0390.030 0.58604424 0.281 0.0460.000 0.000 0400 0.000 0.042 NAVAL",ROCETENES**ANICOPA* 2 USAROL WAND64812AMAZONIAARIZ SW 042113 877633 0 0 0 242312 14 II3 40 00 0 00 30 1 1522 292010 0.0090.0310.000 0.4430.30400530 0.3210.3370.37200274 0.2250.000 0.0000.0000.034 0.0000.0000.093 0.0000.063 0.0000.10404070.041 PALLKOUR8OVAPPI.07031*CAYAMA* MEXFRGPRA HIDALGO 114116024 100 997579 00 0 1 0 46376023 05 o01 o0 1 o0 00 1512 6 24431024 1714 3 0.1170.074080000.047 0.6900.7740.4570.590 0.1020.192003230.353 0.0000.091 0.4000.1280.606 0.0790.0000000 0.0000400 0.0110.00004000 TOTONACNIONIEWINO.46034448111A* 680 4P 1 Ne4VEN40*USA venAfOENTRAL ZULIAARIZ CRUZ .144024262042 464130121235 020 0o 01 136 306371 167299 7 063 0 1 07 00 132017 9 139 831694 104 35 94 040440.0390.036 0.7810.4240.513 000910.2740.220 0.0000.0460.1310000 0.0000.0340010 04000000950.000 0.0000.06104000 060770.4400.000 VAPARANAtWAWA° WINIC/44+4414706 VENVIM WANAPIAREORNOCOORZNOCO 120134120 147261 0 o0 o00 3237 3 023 0 0 0 0 24 2*37 9 02 0.1530.019000810.037 0.3240.6020.70400552 0.234063230.21900231 0.6000.00004000 0.000 0.00004000 0.00000000.000 0.0000.01704210.012 VUOPAT1ffiRA* otoneenincn.VACOITA*MAP** 1 veAVEMME*USA EARIZ tULIAruLIAruLtA InsIts042194 179125113161 0o 0o 0 75566763 142632 e 0o17 0o 0o 0o 10 9I4 13703919 14 407 000440.0390.0170062 0.7870.7300.7020.559 0.6010.2450.2360.032 0.6000.000040040.129 0.000 0.0000000 0.0000.0210.00,0400 0.0110.1140.000 '11%/#111M0" TANIS SC, WERELISTRO EMPLONTO to MISTME tANTIC. 'MM.PNEIRDNICT IN ANTWO. TotorImam OF ANTI.41oftynoTHE GROUP O DOWbUT OPTHE THE CALCUtATIONS RH COMPLEX. NATION REGION REF was EMPLORMO In. 102. 100 AND IN THE REMAININO. THREE A# DETERMINED bY NI PATID saw, OF THE sm4AmPtes LIMITED TO THE O COMPONENT. ANISSIRA CAIUA*ROR0004 IARZAla mA GROSSO 197 SAHPLE SIZE 103 PHENOTYPESRH+ 0 108RH+ GENE FREQUENCIES 0.000 0+ 1.000 UTFNOOTKAGOAJIRO*CUNA4 2 21 USACANVENPAN UTAHNW ZULtAWSMA COASTRLAS GROSSO IS 161102118072197 275161152135 89 0 161275152135 89 0.0000.000 1.0001.00016000 TRIMTAHLE OR GROUP 404, THE JK(A) Akk JK115) GENE FREQUENCIES, AS DETERMINEDNATION WITH REGION REF SAMPLE PHENOTYPES ANIP.JKIA) AND AN11..',8(t8 GENE FREQUENCIES SERA. CAKCNIOUELARECUNA4ACAWA104 1 GUAsEEGUY L ALTITLNPERON 171130 SIZE 90 A48 22 A-8+ A48424 4471 0.489 JKA 0.511 JK8 NAMLACANOON*KEKCHICHEROKEECAMARACOTO* 1 1 *2 MEMGUAUSASEE CHIAPASCORANN CARLINAPEMON 209171173 154141 6177 241723 4 24572247 337136 0036045930.4680+415 0.6640.60700520.585 PERON*NAVA.JCNASKAPI 4 ARECUMA* 3P VENYENUSACANGUA OUEOECLAARIZSAN SABANA JUAN 124063033124173 267237152112 70 1757644117 2349705462 160124 301713 0.4770.5320.5230.299 0.5230.4680.4770.701 A SUMO*RAMAPIAROA* 1 CAMARACOTO*TAUREMAM. N1CYENVEN*EN RAMAAPURELAV. SABANASAWANA KAY 175134124 109108109 36 3010 47 432126 S 24595773 0.4440.3350.5420.4270.457 0.5560.6650.4560.5730.543 VARURUoWAICA.TZOT2IL*TAUREPAIMI 1 VENYENMEXSEERIC APURESCHIAPASE COASTBOLIVARPERON 223229171175 202141103 79 23341322 25172326 90544355 0.4900.5600.4370+481 0.5100.4400.5630.519 A /ne .10141,11;44, ;44.401.0*, ,4ell'4 I re. TARLE 411. WITH THE USE THEOF JKIA/ANTIJKIBI AND JIM,/ONLY GENE FREQUENCIES. AS DETERMINED %arm ANTI..JKIA1 SERUM. THIS OAS MOT BEEN TABLED). (THERE IS A SINGLE STUDY I114I AGUARUNA*TRIBE OR GROUP NATION PRU * MARANON REGION 166REF SAMPLE SIZE 151 PHENOTYPES 67A+ 64A.. GENE FREQUENCIES0.254 JKA 0.746 JK6 AYMARA*ATHABASCANATACAMENOSALACALUF 31 3 1 PRUUSACHLCML MAGELLANPUN°PUNOALASKAANTOFAGST 028062168164 100 93447880 60514430 3549183614 0.4040.3000.5200.3290.436 0.5960.7000.4800.6710.564 CAKCHIOUEL*CARECAR*BRIBRI*BARI*AYMARA* 4 2 GUACOSVENSOL UJARRASSOLALASALITREW3 AREASZULIA P 177170177132169 503 6250TO 9 315 452654 9 188 172416 a 0.4760.307045220.3891.000 0.0000.5240.6930..4780.611 °AMA*CHACOB0*CAYAPA*CAMPA*CAKCH/OUEL 2 5 SOLLCDPRUGUA RSUMPANGO VATACAYAPASURUBMBAMADMI 169167168268 244137 301489 207 74141167 16372263 3 0.270044370.6110.5030.322 0.7300.4630.3890.4970.67$ GUMMI*GOAJIROCUNA*CHOCO*COLORADO* 2 3 2 PANVENECD NW$DARIEN WBLASDOMINGOZULIA COAST IS 178128167 2401111388 00316 256 61995324 179132 202227 0.1360.5900.4170.4230.419 0.8640.4100.5830.5770.581 KEKCHI*JIVARO*ISCONAMUA*/NAPA*ICA 2 2 INNLCDPRUSEECOL TOLEDOARAPICOSRS NEVADAUCAYALIVUPA 176167168131 119233113 14 154104 73 9 4679 59 0.3780.4180.4020018 0.6220.5820.5980.282 NAMMAKIRITARE*MACUSHI*MACOITA*LACANDON* 2 I 1 GUAVEKGUYMEXSEE HUEHUETGOSCHIAPAS BOLIVARVUPA 170130 119 248633 214790 392912 3 0.6460.3270.50604497 0.3540.6730.4940.603 PARINI*PANARE*NAVA*MAPUCHENAM 3 5 2 SEEVENGUACHL W BOLIVARANDELONOUIMAY PETEN SW YUPA 130263164262 141 338070 56303023 111 4724 3 0.6990.4520.1130.181 0.3010.5480.8870.819 QUECHUAQUICHEcortmemPOCOMAMPIRO* 1 1ANO2 GUALCDECDPRU R URUBMBASCENTRALNORTHERN CENTRAL 170167262168 203572220 4090 179220154 2563 152 2474152/ 0.6560.3610.4300.3880.452 0.3440.6390.5700.6120.548 SHIP180*SHAPARU*SECOVA*QUICHE 2 1 PRUSEEECOGUA CUYABENO*S UCAYALICENTRALYUPA 168167262 142 274860 103 294822 391932 5 0.5700.4760.3710.367 0.4300.5240.6290.633 TUCANOTOTONACTICUNA*TERRABAS.SIRIONO* 2 2 MEXPIMCOSBOLSRL R AMAZONAMAZONASVERATERRA8AR MACHUPO CRUZ 186170168177169 122 204540 913518 8 12103122 0.2250.5290.4960.258 0.7750.4710.5040042 XAVANTE*WAP/SMANATZOTZIL*TZELIAL.TUNES°. 2 3P BRZGUYmelt'exCOL MACHIAPAS CHIAPASCOCUYGROSSO MIS 091134170131 445119111 9198 296104 6984 149 22152714 0.42101,6450.508040070.622 0.5790.3550.4920.4930.378 i* ..TlirvisepoirmvIkTom,..,-,,,, 04'1 .1.4 ,t e'ZIIIIPIO 1.4

996*00900 setipc26600 *II OZ£9 ,Z 9ZI lervavilnz 3 WAA 40WWWWWSININW4 WOO966066Z0000S*0ILVO 999006000000062600ZZ60 06tzzzt 0,996;9ZIIL 016ILI6ICIIL 9t00619ZI WWWWW3 SWOrowW il vilAZvilnzlulu 3I1 w3AW3ANumidW3A WO!df Z *V1IO2,06 Z 41ANWVI 414,100WWA svOSIWA orkA SIT 0,41,44t TABLE S. THE MAI AND FV(8) GENE FREQUENCIES. AS DETERMINED eV TESTING WITH ANT14..FYIAI SERUM. AGUARUNA*ACAWAIO*TRIBE OR GROUP NATION PRUGUY A MARANON REGION 168130REF SAMPLE SIZ:0 151 PHENOTYPES137 76A* 14A.,, GENE FREQUENCIES 0.6960.606 'VA 0.3040.394 FVe ATHABASCANATACAMENOSABECUNA*APACHEALACALUF 2 3 I USACHLSEE ALASKAANTOFAGSTNEWMAGELLAN ME% PEMON 062164090 78808444 776$5838 2612 6 0.8870.6110.4440.631 061130.3870.5560.369 BARI*AYMARA*AWEIKOMA 1S ARO* 80LPIMPROBRZ W3PUNOS ZULIAAREASCATKINA P 132169168028233 503109 709358 4:1 46547$ 241:31 0.8220.6560.7370.467 0.1780.3440.2630.533 CAKCHIOUEL*CAKCHIQUELCABECAP*BRIBRI* I 2 VENGOAGUACOS SMOLALUJARRASSALITRE ALTITLN 177170171177 ISO 6250 9 135 5645 9 15 065 0.6840.6890.4141.000 0.3160.5860.0000.311 CAYAPA*CABAjA*CAMPA*CAMARACOTO*CAKCHIOUEL 2 3 ECDBRZPPUSEEGUA RSSUMPANGO ISABELCAYAPASURUBAMBPEMON 167108168268 244137 4089 229125 2786 151312 1 0.7520.4300.8160.704 0.2480.5700.1840.296 CHIRANTECCHAC000*'NH::::EE* mexUSABOLmot 0A.'4CAR MADIDIN* YALTAARLINA 064209169169 20307814 17623022 16 302 0.6130.5470.6221.000 0.3870.4510.0000.378 COLORADO*CHONTOLCHOLCHOCO*CHIPPEWA 2 ECDHEXMEXPANUSA DARIENSTABASCOCHIAPASMINN DOMINGO 167064159178 101137161 3680 126158 299071 11 793 0.5590.6700.7170.6650.864 0.441063100.2630.3350.136 EmER1LL04*DIEGUENOCUNA*CUIVA*CORA* 2 FRGUSAPANVENMEX CALAPURENAYARITS BIAS IS 11420217$134064 388 27588296 343 52951572 124510 6I 0.3330.6780.6590.6510.898 0.6670.3220.1410.3490.102 GUAY0*OUAYOI*.iUARAN14,QUAHIBO*BOAJIRO 3 2 SEEPAN6R2VEN NSAPUREW WCATRINAZULIA COASTWARAO 178233134125 240109119 33 183102106 23 S71013 7 0.5130.4900.7470.669 0.4870.5500.2530.131 HUICHOLIRAPA*HUASTECICA 1 2 SCOLMEX EE SJALISCOVERA NEVADA CRUZYUPA 064131064 113 6975 107 6267 678 0.6810.7700.673 0.2100.3190.327 KALAPALOJIVARO*IRAPA*ISCONAHUA* 2 2ANO*2 I2 ORZLCDPROPRUSEE MAARAP1COSKR UCAYALI GROSSOVUPA 201167168043 233 731416 192 10 06 7141 86 0.0000.2440.5800.388 1.0000.4200.7560.612 LACANDON*KEKCHI*KOICHI 12 I2 MEXBAHGUA CHIAPASTOLEDOCO/INN 171170176173 119162 6033 112136 5732 24 3I7 0.7760.8260.7570.615 0.174042240.2430.385 MAKIN/TABF*MACUSMI*MACOITA* 2I 1 VIMGUYSEE c ROLtVA*YUPA 148130 119 86 106 81 Il 1 0.8130.696 0.1870.104 PAM I GU11 $AN JUAN III 116 110 6 0.773 MAPUCHEMAN 25 2 CHLGUAGUA LONOUIMAYWHUEHUETGO AND SW 164252170 141 7024 129 5224 1218 0 0.708 :::::0.5070.000 KAMAMANUAMAZATECMAYA* 31 1 NEXMEX VERAPUEBLAOAXACAE WEN CRUZ 064263 141136120 41 123103 40ss 332214 I 1:1;26:0.572 0.156::::: OYANP1*OAYANA*NASKAPTNAVAJO 4 USAFRGFRGCAN :4511:EC 114063033114 237152 2271 2::149 14 16 873 0.7240.860 ::1411;0.6030.3140.276 'ENO**PARINI*::::::" sp YENSEEFRG LAyytItIVAR SABANA 124130114 247 3374 3o50 24 3 :::$2690.699::::1 0.2710.3010.569 PIAROA* 1 TAUREPAN*CAMARACOTO*ARECUNA* VEN LA SABANA 042134124 108109 $170 IS 4o 21211 3 0.8330.6550.733 0.3450.267 POCOMAMPIRO*PIMA* PRUUSA CENTRALRiivi:BANA URUBANB 062166 184 $090 179 :::::6376 1714 0.606:3:15 ::::: RANAQUICHEOUICHCounmuAOUCCHUA 211 2 NICGUAECD RANA$SSOUTHERNNORTHERN CENTRALCAT 262175170167 203372232 3780 183344220 6533 13202$12 4 0.6710.567:::::WS,0.:773 :::::0.314 NO1.... SHIPIBO*SHAPARU*SECOYA*SAMNA* 1 SEE R:Zig:: UCAYALI 168167 142 48 138 41 47 001320.618 :OE SIRIONO*SNIP1150*SUMO* 2 EPRUNICB OL : =gig 169043179 103 2710 669827 450 0.761 0:06:312ii0.000 11144ita/ TAUREPAN*TICUNA*TERRABAS* SEE R AMAZONTERRAIAEP:ZIT 177 122 40 104 37 18 3 0.6160.726 0.274 is3 TOTONACTUCANOTOTONAC 22 3 ECOLBR?NEX :51grr:;SPUEBLAVERA CRUZ 13115606417016$ 100 207945 SO7s43 I 1012 02 iiiii0016: 778590 0.7750.000:3:: WA/CA*T1:;;k:Ttlit 1 SEENNEX*EX EX CHIAPAS YANDNAMA 170171 111 9180 1137694 17 86 0.7040.776 HAVANTE*WARAO*WAPISHANA WINIK/NA*GUAY0* 3P BR?VENGUY ORINOCO D 091120134 539127119 7281 IIS102 7074 1217 27 0.5330.7060.622 00.22::0.307!!!!! YANOMANA*VACUA*YABARANA* 10P VENPRU RIORMANAPIARE AMAZON 014168134 491 14 9 425 11 co 114 57 03 0.3410.00':::::: SANENA* VENYENVEN 134 156 434 !!!!! ::::: VARURO* WATCA* VEN i EFfili :111ZIA 123129 102141 0.525::::: 0.413 YUPA IRAPA* 12 sp 11111:VEN 126193130 138154176 1121Iii 97 426437 0.566065060010:1460309099102 0.4340.608 ZAPOTEC SHAPARO*PARINI*MACOITA* 1 OEXVENYENVEN OAXACAI ZUL1AEI1E 2UL1A2ULIA12!!! 064126195 105117 247478 6382le6253 23122$54 6 Eili0.321 0.4680.500::::::::6: , I J Ala at. f - rovr.,.11famolot, wet1,$10..r.,g ei* TABLF 6. THE DICAI AND DUB) GENF FREOUFNCIES. AS DETERMINED BY TESTING WITH A% ANT1...011A/ SERUM. TRIBE OR GROUP NATION REGION REF SAMPLE SIZE DIA+PHENOTYPES DIA GENE FREQUENCIES DIA DIB ARECUNA*APACHEALACALUF*AGUARUNA.ACAWAI0* 2 3 SEEUSACHLPRuGUY NEWMAGELLANR MARANO% MEXPEMON 090164168130 108151 4490 1237 40 104:39 4453 0.0190.0000.0410.233 0.9811.0000.9590067 AYMARA*AMEIKOMAATHABASCANATACAMENCS 1 ANO* 1I PRUBRZUSACHL PU40SALASKAANTOFAGST CATRINA 028233062164 109 587880 5910 407 48SO7873 0.0900.3230.0000.045 0.9100.677140000.955 ABARI*AYMARA* e24 VEMBOLPRU W3PU40MOCOMOCO AREASZULIA P 132169168259 491126100 9350 !: 4::1:g 96 0.0000.0500.020 0.9500.9160.9801.000 CAKCHIOUELCAINGANGCAMECAR* R/RRI*I GUABRZCOS SDLOLALRUJARRASSALITRE ALTITL4G SUL 231170171177 150257 62 5 040841! 0g 1:i 62 0.0000.0440.152 0.956068481.000 CARAJA*CAMPA*CAMARACOTO.CAKCHIQUEL* 2 BRZPRUVEN SRLA ISABELURUBAmB SABANA 100168124 109 3689 381319 235190 0.2010.2430.091 0.7990.7570.909 CHAMA*CH6C060.CAYAPArCARINA* 2 VEN(SOLBOLECD MONAGASR MADID/YATACAYAPAS 169167118 240170 3014 222150 4 219120 10 8 0.4840.1550.0450.160 0.6400.5160.8450.955 CHOCHOCO*CHIPPEWACHINANTECCHEROKEE* L MEXUSAMEXIX OAXACACHIAPAS:11:7E4N CARL/NA 06420917814e 148 54602078 5316 740 132 4727167e 0.1050.0670.4190.0560.000 0.6950.9330.5810.9461.000 CUIVA*COLORADO*mom.CHONTOL 2 ' 2 VENPANHexECDMEX SAPURENAYARITTABASCO BIASDOMINGO IS 064178134167 317 82963689 242317 0I 293 62733572 0.0390.0000.1280.0140.101 0.9860.899009611.0000.872 GUAHIBO*GOAJIROGOAJIRO*WUEMERILLON* 32I VENVENFRG M ZULI6APURE2046 110116134128114 209119152 7628 112312 84 107144 658624 0.1120.0520.0270.074 0.688049480.9730.926 HUASTECGUAYMI*GUARANI*GUAHIBO*GUM* 2 2 MEXSEEPANPAZ VERA4S WCATRINACRUZ COASTMARAO 064233178 240 7034 14 5 I 239 6520 0.0020.0360.2330.075 0.7670.9250.9640.998 HUICHOLISCONAH0A*ISCONAHUA*IRAPA*ICA I AND 2I 2 PR0PRUSEECOLMEX RSJALISCO UCAYALINEVADAYUPA 168043131064 112 721416 4729 4 65431012 0.1550.1340.2380.227 0.8660.8450.7620.773 LACANDON*KEKCHI*KEKCH/JIVARO* 1 2 I HexBRHGUAECO CHTOLEDOCORMARAPICOS IAPAS 171170176173167 117122233 6133 371011 P7 109115196 5122 0.1840.0350.029041083 048160.9650.9710.917 MANMAKIRITARE*MACUSHZ.MACOITA*LACANDON* 1 12 21 GUAVENGUYMEXSEE SAN$CHIAPAS BOLIVAR JUAN YUPA 171130 116119 86 172634 104 6085 0.0530.1650.1550.086 0.9470.6350.8450.914 "Ver, MAPUCHEPAM 2 2 CHLGUA LONQUIMAYW AND SW 164262 130 70 23 5 125 47 0.0190.181 0.9810.619 MANUAMAZATECMAYA* I1 1 REXMIXMEXGUA VERAPUEBLAOAXACAE PETER CRUZ 064269 128136120 32 13181636 2 110120 SO84 0.0730.0320.0610.163 0.9270.9660.9390.817 OAYANA.NAVAJONASKAPI 4 g:CANFRG ARIZQUEBEC 114063033 237152 989130 113417 226119 647419 0.2040.1920.0980.0230.044 0.7960.8060.9020.9770.956 PERON.PARIRI* CAMARACOTO*ARECUNA* 30 VENV:: LAW BOLIVARSABANA YUPA 124130 109287 7033 192578 209 904514 0.0910.1470.1980.349 0.9090.8020.8530.651 POCOMAMPIRO*PIAROA*PIAROA. ANO21 TAUREPAN* GUAPRU'YENVENVFN CENTRALRAMAZONASAPURELA URUBAMRSABANA 262168134124 111108 809024 13171934 3 6357219274 0.2040.0650.0900.172 0.7960.8870.9350.9100.828 QUICHEQUECHUA 21 21 GUAECO SSOUTHERNNORTHERN CENTRAL 262170167 355220 8046 259250 8 263178 5538 0.0910.1390.1160.113 0.9090.8610.884 SHAPARU*SECOU*SANEMA.RIONEGPINO*RAMA SEEMK CUYARENDORAMA CAY YANOMAMA19UPA 167175 4837 90 3937 0.0990.0000.171 0.9011.0000.829 11114- d. 7 SUmO*SIRIONO,SHIPIBO*SHIPIRO* 21 NICam.PRU ER COASTUCAYALIMACHURO UrA:ALI 043175169168 103141 2770 114291 1 92262850 040550.0190,3670.405 0.9810.6310.595 414 TOTONACTICUNA*TERRABASsTAUREPAN. 2 REXPRUCOSSEE VERARTERRABA AMAZON CRUZPEMON 170168177 122 4140 4.4 90 0.1110.2000.000 0.8890.6000.9451.000 TZELTAL*TUNERO.TUCANOTOTONAC 2 3 MEXCOLBRZ CHIAPASCOCUYAMAZONASPUEBLA MrS 064170131186 111100131 49 1146 I4 100 0.0510.0050.1940.042 0.9490.9950.8060.958 WARAO*WAP1SO4NAWAICA.TZOTZIL*TZOTZIL. 2 1 VENREXMEXWYSEE ORINOCOCHIAPAS DYANOMAMA 114170171 119127 8680 331315 0 127 867365 0.0000.1500.0790.099 1.0000.8500.9210.901 YABARANA.XAVANTE. WINIKINA*GUAYG* 3P VENVIMBR1VEN ORINOCOMANAPIAREMA GROSSO 091120134120 538 728114 164 903 374 7278 5 0.4020.1660.0000.019 0.8340.5980.9811.000 YARURC*YANOHAMA.VAGUA* WAICA*SAME48* 10P VENVIMPRUVEN APURES5R BOLIVARAMAZON /23129134014168 102141156571 5092 24114757! 97 7 0.0250.0000.029040000.118 0.9751.0000.9710.882 YUPA. MAC0f7A*IRAPA*IRADA* 21 3P VENYEN E ZULIAE ZULLAZULIAZU11A 121193195130 138117176 44 46 30 135110 44 01080.0110.0000.141 0.8920.9891.0000.859 ZAPOTFC RIONEGRINO.PAR1RI*MACOIIA*SHADARU* 2 4FXVI%VE4VEN OAXACAFE ZULIAzuLtA 064126194 105125 782474 1531252417 6 901894496193 0.0740.154041330.1860.116 0.9260.8660.6670.8140.884 , %dew* ' ..4...1a.40,3411A4 TARLE 7. LOWERTHE P CASEGENE P.FREQUENCIES.TRIBE OR GROUPAS DETERMINED 87 TESTING WITH AN ANTI*0 SERUM. FOR NATION REGION REF SAMPLE oPP0 READ CAPITAL P AND FOR *Pop AGUARUNA*ACAWAIO* PmGUY R MARANON 168130 SIZE 151 90 PHENOTYPES140 64PI 1126P2 GENE FREQUENCIES0.7300.463 PP 0.2700.537 ATHABASCANATACAMENOSARECUNA*APACHEALACALUF 2 3 I USACHLSEE ALASKAANTOFAGSTNEWMAGELLAN MEXPEMON 062164090164 77614480 61143525 631926 0.5130.1470.343 0.4to70.6530.657 AYMARA*AYMARA*AWE1KOMA 43 1 ANO0 SOLPRUBR2 3MAOPUNOS AREASCATRINA P 028169168253 503122 9558 434 922141 693781 1 0.6300.896',.2010.1850.095 0.1040.7990.8150.905 CAINGANGCASECAR*BRI8k146emu. COSBR!COSVEN RUJARRAS GSALITREW SULZUL1A 231177177132 166 62SO70 61427332 9338 18 0.2520.8730.6000.263 0.7480.1270.4000.7370.370 CAYAPA*CAMPA*CAMARACOTO*CAKCH2OUELCARCHIOUEL 2 13 PRUECDSEEGuAGUA R CAYAPASURUBMPAsuMPANGOL ALTITLNDEMON 167168268171 244137150. 89 134116 6685 110 235234 0.3290.4920.3840.524 0.6710.5080.6160.476 CHIPPEWACHINANTECCHEROKEE*CHAMA*CHACOBO* USAMEXBOLROL MINNOAXACANR MADIDICARLINAYATA 159064209169 161 207$3014 136 652014 0 23121310 0 0.6060.2250.5920.4232.000 0.3940.7750.4080.5770.000 COLORADO*CORA*CHONTOLCHOLCHOCO* 2 MEXECOPAN NAYAR/TSTABASCOCHIAPASDAR/EN DOMINGO 064167064178 101150 963680 7429SS4763 103 462217 7 0.5210.5590.3250.1710.539 0.4790.4410.6750.8290.461 GOAJIROEMERILLON*DIEGUENOCORA*CUIVA* 2 3 VENUSAPARFRGVEN wAPURE ZULIACALS 81.AS IS 128114202178134 388119 5882 7 328 636433 3 56256018 4 0.3140,2440.3430.6070.531 0.3930.6860.7560.6570.469 mu/CHOLHUASTECGUAVW*GUARAN/*GUAHIBO* 2 2 REXMEXPAN8R1VEN JALISCOVERANSAPURE WCATRINA COAST CRUZ 064178233134 240109 727534 197 675624 19434220 4 0.7630.4970.5770.3790.458 0.237065030.4230.6210.542 KALAPALOJIVARO*ISCONAHuA*/RAPA*ICA ANO*2 2 2 BRZECDPRUSEECOL MAARAPICOSRS NEVADA UCAYALIGROSSOYUPA 201167168131 231113 7314 181 301441 435072 0 0.5370.2021.000 0.4630.0000.798 MACOITA*LACANDON*LACANDON.KEKCHI*KEKCHI I 2 2 SEEMEXBIMGUA CHIAPASTOLEDOCOBAN YUPA 171170176173 119162 6133 106137 4724 142325 9 045210.4780.6690.6070.233 0.4790.5220.3310.3930.767 MAPUCHEMAPMAMMAK/RITARE*MACUEHI* 13 2 1 CHLGUAVENGUY LONOU1MAYWSANS ANDBOLIVAR JUAN SW 262164172130 141116119 7086 137 38895779 32272940 4 0.8320.3240.5180.4190.420 0.1680.6760.4820.5810.580 MANUAMAZATECMAYA* I1 I MEXGUA OAXACAE PETENPUEBLA 064261 136120 41 432385 189335 0.3370.1730.460 0.663008270.540 PALIKOUR*OYANA*NAVAJONASKAPTNOWA 1 4 MEXFRGUSACAN OUEBECVERAARIZ CRUZ B63033064114 237152141 7599 142108102 3846 3753954439 0.2980.2680.3670.4620.474 0.7020.7320.6330.5380.526 PERON*PARIRt*PANARE* ARECUNA* 3P VENSEE LAM BOLIVARSABANAYUPA 124130 287 3370 149 5436 8 138 553425 0.2900.3030.3070.130 0.7100.6970.6930.870 PIRO*POCOmAMPIAROA* 1ANO CAMARACOTO*TAUREPAN* GUAPRUVEN APURECENTRALRLA URUBMAASABANA 262168134124 111108109 8090 597584 21274915 0.3260.4860.5920.507 0.5120.4080.4930.674 SANEMA*RAMAQUICHEQUECHUAouemA 2 12 NICGUAECDLCDSEE RAMA$CENTRALNORTHERN CENTRAL KAY YANOMAMA 115262167 372221 3780 338204 3155 342517 6 0.5970.4410.6980.723 0.4030.5020.2770.559 SM/PI11041SMAPARU*SECOVA*SIRIONO* 2 MtPRUSEEECD CUYABENOR UCAYALIMACHUPOYUPA 167175169168 103142 4827 109 642029 393319 7 0.3850.4910.5180.371 0.6150.5090.4820.629 TOTONACTICUNA*TERRABAS*TAUREPAN*SUMO* 2 MEXPRUCOSSEENIC VERARTERRABAE COASTAMAZON CRUZPERON 170168177 122 4045 121 2738 18 2I 0.3670.90904776 0.6330.0910.224 TZELTAL*TUNE90*TOTONAC 3 MEAREXCOLMEX COCUYPUEBLAcHiApAsCHIAPAS MT5 171170131064 101100 758079 54676850 4621133329 0.4710.5970.4280.3220.394 0.5290.4010.5720.6780.606 WAAAO*MAICA*TZOTZIL*MUM*YAGUA*XAVANTE* 12 3P ME%PRU6R2VENSEE RMAORINOCOCHIAPAS AMAZON GROSSO 0YANOMAMA 091170168134 538127 9 468 59 9 7088 0 1.0000.6390.168 0.0000.3610.832 YUPA*YARURO*VANOMAMA* %MICA*SANEMA 10P 3P VENYEN EAPURES BOLIVARZULIA 014130123129134 570176102141156 427 76628179 143100 406077 0.2460.3740.3480.2970.499 0.7540.6260.6520.7030.501 ZAPOTEC SHAPARU*PARINI*MACOITA*!PAPA* MEXVEN OAXACAE ZULIA2ULIA 064126 105138 247478 41234474 9 64513415 0.2090.1700.3400.3190.219 0.8300.6810.7810.7910.660 TAMA sae SOONMA. UNECALCULATIONS PAIONINCItsTwit THENOTT01 SAS NOTCANNOT OCTIENNINSOTIONINcIts me SouTIMELY POOPON ROSTTwo CoMPOID MI6SAMPLCS* AMEN, OtCAult A$ OCTIONINOD tilt Allim DT SWIMTOOTING MOUS WITH wmICK ANTI-LOIAI IS NOCOASAMV AND ANTIeramil TO AGUARUNA4TRIBE OR GROUP NATION PRU R MARANON REGZON 168REF SAMPLE SIZE 151 MA.. A+90.28 PHENOTYPES 0 148.1.123 PHENOTYPE FREQUENCIES0.185 0.000AS 0.115 A+O A.41+ AYMARA*ATHAAASCANAPACHEALACALUF 2 45 3 1 SOLPRUUSACHL 3PUNOALASKANEWMAGELLAN AREAS MEX P 169168062090164 503 93774944 4810 4 200 453 89733934 0.0950.0430.0520.2040.227 0.004 0.0000.000 0.900 0.9570.948067730096 CAXCNIOUEL.CAMPA.CAKCHIOUEL*CARECAR.RRIRRI* 21 PRUGUACOS RSOLALAI.UJARRASSALITRE URUOMAAALTITLN 160170171177 150 8962SO 9 32272016 4 010 122 425734 5 0.3600.1800.4440.3230.320 0.00006007 0.000 0.6400.812 0.5560.6770.680 - ,...... CNACOSOs SOL R TATA 169 3014 11 0 0 1914 0.000 0.000 1.000 4.11.. CORA*COLORADO*CHOCOMCVAMA* 2 2 PANECDsot SDARIENp MA010/ALASDOMINGO IS 178167 388 36SO 173 2711 IS 0 215 2535 0.4460.3370.3670.306 0.2250.000 0.000 04.4370.5540.611 0.694 1/44 KEKCHI*KEKCHIJIVARO*GUAYm/*TSCONANUAC 1 2 2 ORHGUAECOPRUPAN TOLEDOCOAANARAPICOSRN uCAYALIW COAST 176171167168178 119162233240 14 106123 9132 3 0 130127117 2411 0.7650.1980.4550.5120.214 0.000 0.000 0.2350.8020.5450.488 0.766 MAPUCHEMAMNANLACANDON.LACANDON* 2I 2 21 OILGUAMEX LONQUIMAYHUEHUETGOSANCHIAPASCH/APAS JUAN 164170171 141116 246131 341810 59 01 123 52188223 0.1280.2080.2930.1480.303 0.0000.042 0.8720.7500.7070.0520.697 ouecHuAQUICHEPIRO*NAVAJONASKAP1 11 ECDGUAPRUUSACAN NORTHERNSRARIZOUEREC CENTRALURUBMSA 167170168043033 231203237152 90 73422425IT 021 158159212127 73 0416C0.1890.101 4070.164 0.0000.0100.0000.004 0.000 0.6840.7820.8120.895 0.836 OUECHUASUMO*srlitrono*SMIP/O0eRANA 2 1 NICrimPAUNrcECD ERARAmACENTRAL COASTUCAYALINACHUPO KAY 175169168167 103142168 2737 110 372921 4 13 0 113258 532316 0.3590.1480.2040.5680.299 0.1260.0000.00 0.7960.5150.6520.4120.701 TOTONACTICUNA.TERRABAS* 2 MEXPRuCOS VERARTERRAAA AMAZON CRUZ 170168177 111122 4539 2616 62 01 120 843723 0.1780.0160.410 0.000 0.8220.9840.590 YAGUA6KAYANYE*rzortiLeT2OTZ/L'1?ELTALC 21 3P PRU8R2MEXHEN RMACHIAPAS AMAZON GROSSO 168091170171 338 9180 9 641613 0 102 02 172 6773 9 0.0000.1890.1760.1630.234 0.0220.0000.3020.009 0.8011.0000.5080.8370.757 ...4.646ate NiVidAt taidibihk4/6. itiomaii 41* liairt4+4.4,644. 44 . 4.- 04' 1

TAIDLE s. DETERmINEDCANNOT BE ROUTINELYTHEFOR PHENOTyPEMOST sAmPLES.COMPUTED FREQUENCIESTRIBE BECAUSE OR GROUP FORTHE THEABH LEwIsSECRETOR SYSTEm. STATUS As WHICHDETERMINED IS NECESSARY BY TESTING TO SUCHWITH CALCULATIONSANIPoLEIAI. WAS NOT NATION REGION REF SAMPLE PHENOTYPES PHENOTYPE FREQUENCIES GENE FREQuENCIES ATACAmENOSARECuNA*ACAWAIO* 1 CHLSEEGUY ANT0FAGs7PEMON 164130 SIZE 8090 A+ 20 7890A- 0.0250.000 A+ 0.9751.000 A- CAMARACOTO*BARI*AYMARA*AWEIKOMA 1 ANO VENSEEYEWPRUBRZ APuREWPUNO$ ZULEACATRINAPERON 134132028233 118 $27058 69 00 82705849 0.0000.585 1.0000.415 2=GUARANI*TAMPA.ICAGUAMIBO* 2 0 3 SEECOLVENYEN W ZULIASAPURE NEVADACATR/NAyupA 233131134128 113109119 34 22 C01 113108119 12 0.0000.6470.009 0.3530.9911.000 OTOmi*MAKIRITANE*MACUSHI*MACO/TA*1SCONANUA. 11 mExVENGUYPRU S BOLIVARP UCAYALI 024130043 119 818616 80 119 738616 0.0000.000 1.0001.000 PEmON*PARIRI*PANARE* ARECLINA* 3P VENvENSEE WHIDALGO NDLIVARLA SABANAYUPA 130124 267 3370 01 286 7033 0.0000.0030.099 0.9970.9011.000 P/AROA*SECOYA*SANEmA* I TAuREPAN*CAMARACOTO* VENvENLCDSEE APURELA CuYABENOSABANAYANOMAMA 167134124 100109 8148 01 108 4881 0.0000.009 1.0000,9911.000 TuNE80*sMAPARU.7AuREPANwSHIP1B0* 2 coLSEEPRU COCUYR UCAYALI mTsYANOMAMAPEmONYuPA 131043 9970 0 9970 0.0000.000 1.000140000 YARURO*YANOMAMAng: : MAMA*SANEMA4 VENYENvEN APuREsORINOCO 801.1vAR5 BOLIVAR 0 123129134 1411F6102127 0 102141156127 0.0000.000 1.0001.000 YUPA* soAPARu*PARIRI*N co1/100,4A * 3P vENYENVEN'..EN E ZuL1AZULIA2ULIA 126130 136176 247478 02 136176 2478 0.0000.014 1.0000.986 4 TARLE 9. TR/BE OR GROUP SOTHE RARE. TRANSFERRIN GENE FREQUENCIES PHENOTYPES ARE AND GIVEN GENOTYPES. POR C AND FOR ALL NATION REGION REF SAMPLE BECAUSE THE 404C PHENOTYPES ARE SO MANY BUT YET INDIVIDUALLY OTHERSPHENOTYPES COMBINED (EXPRESSED IN THE TABLE AS I:K. AGUARUNA6ACAWAIO* PRUGUY R MARANON 16$022 SIZE 151 64 801 0 82C 0 802C 0 151 84C COCHI 0 CO1 0 DI DCHI0 0 GENE FREQUENCIES000000.000 1.C 100001.000 AYMARA*ATHABASCANAWEIKOMAATACAMENOSALACALUF* I ANO* 31 PRU8RZUSACHL PUNOSALASKAANTOFAG5TMAGELLAN CATARNA 093227092164 5637497645 00 00 00 5637494379 00 00 0 0 0.0000.000 1.0002.000 BRIBRI4BARI*AYMARA* 45 COSVENSOLPRU SALITREWPUNO ZULIA3 AREAS P 237022169168 3871 0 0 0 71535471 17 500 00 0 00 0.12000000060000.000 0.8801.0001.000 CAMPA.CAKCHIQUEL*CAKCHIQUELCAINGANGCABECAR* 12 GUACOSBRZ SMOLAIRUJARRAS GALJUILN SUL 260165236266 150116 1025 00 010 00 142115 1024 001 09 00 00 0.000040270.0040.0200.066 00130.996009800.9341.000 CHINANTECOCHIAPANECACHIAPANECACAYAPAr 2 1 MEXmExMEXECDPnu OAXACACHIAPASRA CAYAPASURUBAmn 165260165167168 226 474099 0 0 00 206 453893 0 20 20 0 0 0.0210.0250.0440.000 0.9750.9561.000 41 126 CUNA*COLORADO*CHOLCHOCO* 2 2 2 PANECOMEX 5SCHIAPASDARIEN ALASDOMINGO IS 166167260 174 36167453 0 00 03 174 36161450 00 00 00 00 0.0000.028 1.000160000.9720079 ; Ch GUAYMIA2SCONAHUA4IRAPA*GUARAN/*GUAH190* 1 PRUSEEPANBR2VEN N$APURE WCATARNA COASTYUPA 166237022 204112 30 00 0 00 180112 90 0 24 0 00 0 0.0590.0000.000 0.9411.000 LAC/000mmKEKCHI*KEKCHIJIVARO* 2 2 1 mExBRHGUAECD cHIAPASTOLEDOCOURARAPICOSR UCAYALI 260166165167043 162221 316516 00 00 01 153220 5816 00 5901 10 0 0.0620.0200.0020.000 0.93800720.9981.000 MAKIRITARE*MACUSHI*MACOITA*LACANDON* 21 VENGuYSEEmEx SCHIAPAS ROLIVARYUPA 022165 116 5459 0 0 10 0 116 544931 0 00 00 00 0.000040850.000 1.00000151.000 NAVAJOHAMMAYAMAP 2 1 43 USABRHGUA ARIZTOLEDOHUEHUETNGSAN JUAN 203166260165 230212116 27 00 01 16 0 195112 27 00 16 04 10 0 0.0420.0000.0170.000 0.9561.0000.983 QUECHUAPIRO*PIAROAPPEMON4PARIRI 1 2P3P PRUVENSEE 4APURE URUBAMBLA SABANAYUPA 022168 867796 00 00 0 213 839652 20 00 300 05 00 0.0170.1950.0000.037 0.9630.8051.0000.963 SANEMA*RAMAQUICHE 1 SHIRISHAKA4 VENNICGUAECO SRAMAN BOL/VARCENTRALANO CAY CEN 022165260167 192 743794 0 0 010 165 2592 0 27 A1 400 00" 0.2160.0110.070 0.7840.98906930 SHIRISMANA4SHIP180*SHIPIBO*SMAPARUSECOYA4 12 PRUSEEPRUECO R UCAYALICUYABENDOYUPASANEMA 043168167 129 4670 00 00 00 129 704774 0 010 00 00 0.00000100.000 1.0001.0000.990 fele TERRABAS*SUMO-4r- - COSNIC- -E COAST- TERRABA 168166165 122101 32 00----00 0 -0-408- 0 122 29 43 0 00--2 00 0 0.0320.000 0.96810100 720711L*TZOTZIL*TZELTAL*TOTORACTICUNA* 2 12 MEXPRU CHIAPASVERAR AMAZON CRUZ 260265 974$8879 00 00 10 67759745 0 04 0 0 0.0060.0250.00006000 0.9940.9751.0002.0001.000 YAGUA*XAVANTE*WARRAU*WAPISHANAMAICA* 3P PRUWIZVENGUYSEE AMAZONORINOCOMA GROSSO 0 YANOMAMA 168247022 521123116 9 0 0 00 521121116 9 02 0 00 00 0.0000.008 1.0000.992 YUPA*YANOMAMA* MACOITA*WAICA*!PAPA* 2 10P 2P VEN ES ZULIA2ULIABOLIVARS BOLIVAR 027017020022014 136450 428991 0 0 0 236428 3884 38 0 450 02 15 0 0.0280.3740.0000.005 0.9720.6261.0000.995 ZOOM2APOTEC SHAPARU*PARIRI44 REXMEXVEN CHIAPASOAXACAE ZULIA 020165260 31802269 0 0 02 78291127 1127 00 20 0 15 0 0.0320.2500.0130.4130.048 0.9680.9870.7500.5*/0.952 111 TABLE 10. PHENOTYPES THEAND HAMTOGLOBIRTHE °OTHER' PHENOTYPESPHENOTYPES ANDHAVE GEMF BEEN FRFOUEMENS* OMITTED. IN THE COMPUTATION OF THE LATTER. THE AHAPTOGLOGINEMIA TRIBE OR GROUP NATION REGION 018REF SAMPLE size 87 1.=.1 38 2.-1 42 2.-1MPNENOTYPES 0 5 0 2 OTHER 0 GENE FREQUENCIES HP1 HP2 P ATACAMENOSAPACHE.ALACALUF*AGUARUNA*ACAWA10* 2 31 CHLUSAPRUGUY ANIOFAGSTMAGELLANNEWR MARANON MEX 164261148 151 799843 32541350 42471478 0 171545 5 01 00 0.6710.587064760.4570.494 0.3290.41300260.5450.306 AYMARA*AWE1KOMAATHABASCAN 1 ANC* 132 PRU8R2USA POMOSALASKA CATRINAALASKA 093237032245 122284104202 56 2453SI55113 244089:4498 01 136 411763 4 0183 03 0.3460.7140.4640.5650.429 0.6540.2460.3360.4350.571 CAINGANGCAPECAR*BRIM*AYMARA*AYVARA* 43 ERZCOSCO3ece.PRU RUJARRASSALITREPUNO G3 SULAREAS P 236146237169168 326 253871 103 5636 41 110 141326 12 0 2124 78 01 0 0.75e0.1970.7000.4400.700 0.2420.5600.8030.300 :-. CAKCHIOUEL*CAKCIONEL I 2 GUA SOLOLAI ALT1TLN 260165 150 20 56 4 73 4 0 18 1 13 0 0.46704629 0.5950471 CAMPA*CHIAPANECACEARACAVAPA44 2 21 MExMEX8R2ECDPRU R URUSAMRCHIAPASNOTR CAVAPAS REPTD 240145105147148 226 47249335 219833 48 1620139737 012 231124 5 0 03 0.4570.6740.4770.554 0.4440.5430.3240.523 COLORADOCHOLCN1NANTECCHOCO* 2 2 * 2 (CDHEXPANMEX OAXACA$CHIAPASDARIEN DOMINGO 147145144 34167453 28III 95 2829 48 0 2419 03 804 0 0.8090.4840.4570.368 0.1110.3120.5430.432 GUARANI*GUAH160*GUANARIRO*CUNA* 2 2 ON2VENPAM AMAZDNASSAPURE CATRINABLAS 13 237134017166 117174 3419 551519 9 45911583 4 04 334015 90 012 0 0.5000.6740.8950.173 0.5000.3240440S0.62704,385 JIVARO*HA!DAGUMMI*ISCONAHUA*IRAPA* 2 1 PouSEECANPAN ROUEEN UCAYALIN W CCOAST IYUPA 167043150 222413204 16 113 92SO I 100221 13 00 2879 2 20 0 044450.4690.5410.415 0.5310.459 LACANDON.LACANDON,KEKCHI*KWH/ 1 2 12 MEX8RHGUAECD CHIAPASTOLEDOCONNARAP1COS 165260166165 162 593145 492474 6624 94 00 2213 00 1320 0 0.9220.9290.5870.641 0.0710.4130.33110055 MAKIRITARE.MAKIRITARE*MACUSNI*MACOM* 12 YENGUYSEE AMAZONASS BOLIVARYUPA 265017019018 129 5985 31271220 55183054 200 372714 18 40 00 0.6100.3420.465 0.3900.6580.535 NAVAJOPANARE.MAYANANMAR 1 254 VENUSA15104OURGUA itARIZTOLEDOSAN BOLIVAR JUAN 023203166260 261212226 3227 5792 28 119 941611 0 II872624 8 30 00 0.3430.4430.656065000.532 0.6550.5570.5440.5000.468 QUICHEQUECHUA6190691040149NEN0N. 1 1 2039 SUAECDPAUVEX NRAPURELA ANDUBURAMO SSABANA CENTRAL CEN 260147168019 192214 9486 112 36295440 113 46664133 0 124916 96 12 051 0 06428047750.5740.7640.4741 044720.2250.4240.2320.522 'SAWA*RAMA SEEPIC RAMA CAYYAMOR AMA 165 37 19 _ . 12 .... 0 6 0 0 ...... ___0.676 0.324 ittSWA.-SNIPING*SHIRIBO. 12 PRUPRO CD R UCAYALICUMEN00 UCAYALI 043169167 129 7048 255414 415823 1711 4 O0 00 0.6500.6430.531 0.3500.3570.469_ TLINGITSUMO.TICUNAsrEaaanas, USAPROCOSNIE ALASKARTERRA8AE AMAZONCOAST 032168166165 122108 9211 6016SI 9 38454914 1 261727 8 010 10 0.4380.6760.5160.519 0.5620.3240.4840.481 TZOTZIL*T2ELTAL*TnTORACTZOTZZL* 22 1 mcXMcXMEKMCXSEE CHIAPASVERACHIAPAS CRUZ 260165 se809745 313514 44365423 1312 a7 0 0 0.5980.6020.6750.567 0.3980.3250.4020.433 YANOMAMA*YAGUA*XAVANTE*WAPISMANANAIEA* 10p 3P YENPRUBRZGUY RMA AMAZON 5GROSSO POLIVARYANOMAMA 014168247018 425521120 9 301114 34 2 109254 56 01 139 22Is 3 13 061 0 0.8360.4180.4750.553 0.1640.5620.5250.447 YARURO* WAICASAMEMA* VEINVEN APURES BOLIVAR 80L/VAR 019017 103139 19 428415 4648 4 0 13 450 230 0 0.6440.7940.995 0.2060.3560.105 2APOTECYUPA MACOITA*IRAPA* 2 AMAVEN OAXACAE 2UL/AZULIA 260017017019 129 607493 225712 1 40405111 010 771722 00 0 0.5320.2050.7600.432 0.4680.7950.5680.220 tab

*...0....AmoNXEINISIMI818118.80/11MMWeellbaditig ME5"44- 4 4 , 7A8LE 11. THE GM PNENOYYPES AND GENE FREQUENCIES. TR/8EAS OASE0 OR GROUPON TESTING WITN ANTIGMCA), ANTI...GA(8)s AND NATION REGION REP SAMPLE SIZE A+X.8+ AOI81. A.4.40.. Ailt...8., PHENOTYPES GMAK GENE FREQUENCIES GMA GAUPANTI0418(X/ SERA. P EMERILLONCAN/8AWEIKOMAATHAPASCANACAMAIO* 4 ANO* 1 FRGSUN8N2USAGUY 5ALASKA CATARNA 0782962)3256063 119 349164 11 0 17 420 6$5716 3 Se3319 3 0.220062790.1720.524 0.233067030.81070.476 0.947060170.0200.000 MAKINITANE.MACUSH14GUARANI* 22 VENGUYRR2 AMAZONASS CATARNA 062083233 122116 3440 00 0103 9153151$ 31632$16 064960.263062160.209 065040.7370.7390.791 060000.0000.045 404.2K0UO*4OYAMOI.OAYAM44MOCETOMES, Firt4FRGROL AMAZONIA 078223 79969776 30 01 06SS36SS 15696020 0.9340.2070.1660.476 0.4380.614047660.517 0.0270.0000.0050.007 MICA*YANOMAMA211110100ffloW4101861MNA WAIIC4 VIM642GUYSEE MA OROSSOYAMORROR 247129029 464137124 00 0 3.76 4371 266 4994 0.172062120.277 0.82600010.423 060000.04100.000 soouvait o, 44kit TABLE 12. Data on genes that are probably absent in Indians who haveno Caucasian or Negro admixture Tabulation restricted to samples in which non-Indian ancestry is believed to be less than 5per cent, as listed in Table I. Samples with relatively high frequencies that may be due to technical problemsare starred(*).

Lutheran Among 32 samples tested with anti-Lu a alone, 28 contained no case of Lu(a+). Theyare reported in the following seventeen references: 33, 63, 126, 128, 129, 167, 168, 169, 170, 171, 173, 175, 176,177, 178, 209, 4ind1 268. The 4 exceptions are:

ribe or Group Area Reference Sample Size Lu(a-F-)

ymara Bolivia: Yungas 169 120 1 una Panama: San Blas Is, 178 388 2 leguefio U.S.A.: California 202 58 2 alapalo (2) (*) Brazil: Mato Grosso 201 73 12 639 Among six samples tested with both anti-Luand anti-Lu b, no case of tither Lu(a+) nor of Lu(b-)was evealed (References 33, 176, 177 and 178).

Kell

Among 40 samples tested with anti-K serum alone, 28 contained no positives, andmay be considered to consist holly of hotnozygous kk cases. They are reported in seventeen references:42,64, 74, 81,90,114,132, 134, 45, 171, 175, 195, 223, 231, 262, 263, and 268. The 12 exceptionsare: ribe or Group Area Reference Sample Size K+

thabascan U.S.A.: Alaska 3 206 1 weikoma Brazil: Sta. Catarina 233 122 3 araji Brazil: Sta. Isabel 108 35 7 hol Mexico: Chiapas 64 137 4 ara Mexico: Nayarit 64 96 6 ieguefio U.S.A.: California 202 56 6 uastec Mexico: Vera Cruz 64 75 2 uichol Mexico: Jalisco 64 70 2 Kalapalo (2) (*) Brazil: Mato Grosso 201 73 17

aztec ( I) Mexico: Oaxaca 64 136 1 Nahua (3) Mexico: Vera Cruz 64 141 3 Irapa (1) Venezuela: E. Zulia 193 153 3 1,300

Among 74 samples tested with both anti-K and anti-k sera, 70 containedno case of Kell positive and may bc considered to consist of homozygous kk individuals only. Theyare reported in references: 23, 28, 33, 43, 90, 123, 124, 126, 129, 130, 131, 134, 164, 167, 168, 169, 170, 171, 173, 176, 177, 178, 233 and262. The 4 exceptions are: Tribe or Group Area Reference Sample Size K+ Cherokee U.S.A.: No. Carolina 209 78 I - Chippewa (*) U.S.A.: Minnesota 159 161 24

IGoajiro Venezuela: W. Zulia 128 119 1 /Navajo (4) U.S.A.: Arizona 63 237 4 595 30 IAll of the 30 K-positive cases were heterozygous Kkexcept two of the four Navajos, who were KK.

Wright

1 Among 41 samples tcstcd with anti-Wra serum, not one containeda positively reacting case. References: 63, 91, 1164, 167, 168, 169, 171, 173, 175, 176, 177, 178, 209, 231, and 233.

181 '184 TABLE12-Continued

Miltenberger Among 40 samples tested with anti-Mia serum, only one contained positive cases. A sample of 232 Quechuas at Calderon, in northern Ecuador (ref. 167), contained four Mia+ individuals, two of whom were related as father and son. Relationship of the other two was not ruled out. Thc other 3 9 sarnplcs arc reported in references 1 64, 167, 168, 169, 170, 171, 173, 175, 176,177,178 and 179.

Verweyst Among 37 samples tested with anti-Vw serum, only one contained a positive case-in a sample of 162 Kekchi Mayans in Cobin, Guatemala (ref. 173), ow.: individual was Vw+. The other references are 91, 164, 167, 168, 169, 171, 175, 176, 177, 178, and 2 09.

"V" (in Rh-hr ,system) Among 28 samples tested with anti-V serum, only two samples contained positivecases. Among 12Kekch s tested at CobAn, Guatemala (ref. 173), two were V+, and among 45 Totonacs (sample 2) at Vera Cruz, Mcxi o (ref. 170), one was V+. The other samples are referred to in 2 8, 1 67, 170,171, 175, 176, 17 7, 178, and 20

Berriens Among 18 samples tested with anti-Bea serum not one positive case was found. References arc: 167, 1 71, 17 176, 177, 178, and 209.

Sutter Among 15 simples tested with anti-Jsa serum, only one contained a positive case-one individual in a sample t 119 Goajiros, sample no. 3, in western Zulia State in Venezuela (ref. 12 8). Other references are: 2 8, 126, 16 171, 173, 176, 177, and 209.

182 465 TABLE 13. Tribes and groups estimated to have more than 5 per cent of non-Indian ancestry. Some tribes listed here will be found also among one or another ot tables 1-11. Such duplication resu;ts: from two or more samples having been drawn from the same tribe, whether by the same or by different workers, one or more samples indi- cating less than 5 per cent non-Indian ancestry and one or wore indicating more than 5 per cent non-Indian ad- mixture.

TRIBE OR GROUP COUNTRY REFERENCES TRIBE OR CROUP COUNTRY REFERENCES

Surinam 109 Chorotega Nicaragua 165, 175 Ank; Argentina 2 05 Choroti Argentina 182, 254 Apache U.S.A. 42, 90, 111 Colla Argentina 182, 199, 254 Arawak Surinam 58 Cora Mexico 192 Araucanian Cree Canada and (see also U.S.A. 51. 97, 143, 155, 156 "Mapuche") Chile 189, 196, 211, 2 37 Crow U.S.A. 250 Atacamenos Chile 74, 164 Cuna Panama 72 Athabascan U.S.A. 62, 87, 98, 250 Dogrib U.S.A. 98 Aymara Bolivia; Peru 223; 225 Emerillon French Guiana 114 Flathead Beaver U.S.A.; U.S.A. 154, 155 Canada 62, 98, 156 Galibi French Guiana 114 Blackfeet U.S.A.; 5, 49, 110, 139 Goajiro Venezuela and Canada 154, 155, 160, 162 Columbia 128, 134, 152 Blood U.S.A.; 49, 139 Guambiano Colombia 137 Canada 154, 160, 162 Guarani Brazil 216 Boruca Costa Rica 177 Guatuso Costa Rica 81 Botocudo Brazil 153 Guayqueri Venezuela 134, 135 Cachama Venezuela 118, 119 Haida Canada 102, 150, 264 Caddo U.S.A. 99 Hopi 42, 250 Caingang Brazil 76, 77 Huichol Mexico 192 Canela Brazil 22, 65, 146 Hupa U.S.A. 104 Caraja Brazil 106, 107 Ica Colombia 22, 134 Caramanta Colombia 11 Itonama Bolivia 1 69 257 Carib Surinam Itza Maya Mexico 1 70. 2 ou Carrier Canada 98 Catawba U.S.A. 2 10 Jicaque Honduras 1 65. 1 74 t Catio Colombia 12 Kansas U.S.A. 1 12 Catopaxi Ecuador 242 Kariri Brazil 207 Ceara Brazil 17, 18 Karok U.S.A. 104 Chaguanca Argentina 181, 254 Katios Colombia 12 Chamacoco Argentina 181, 254 Keresan U.S.A. 4 Chemehuevi U.S.A. 42 Kitamaat Canada 2 17 Cherokee U.S.A. 209, 250 Klallam Canada 102 Chiapaneca Mexico 170, 260 Kuaiker Colombia 136, 198 Chibcha Colombia 252 Kokonuko Colombia 137 Chibcha Peru 251 Kutchin U.S.A. 145 Chickasaw U.S.A. 112, 250 Kwakiutl Canada 89, 102, 217 Chinanteco Mexico 165, 187 Lacandon Mexico 171 Chirrya Bolivia 258 Lamistas Peru 269 vyan U.S.A. 96, 98, 143 Lenca Honduras 1 65, 174 Chippewa U.S.A. 52, 56, 159 Lloreos Colombia 253 Chiriguano Argentina 181, 254 Choctaw U.S.A. 57, 112, 25(' U.S.A. 102 Chol Guatemala; Chile 74, 101, 184, 1 88, 1 89, Mexico 64, 170, 171, 2 60, 2 62 196,211,238,239 Chontol Mexico 64 Mataco Argentina 7, 1 82, 254

183 TABLE 13-Continued

TRIBE OR CROUP COUNTRY REFERENCES TRIBE OR GROUP COUNTRY REFERENCES Maya Br. Honduras Quinault U.S.A. 104 and Mexico 95, 170, 171, 176, 218 Ramkokamekra Brazil 65 Mextizo Mexico 170 Micmac Canada 33, 88 Sac and Fox U.S.A. 250 Miskito Nicaragua 165, 175 Sal ish U.S.A.; Canada 62, 87, 89, Mixe Mexico 64, 165 102, 141 Mix tec Mexico 64, 221 Sapelo Canada 219 Mojave U.S.A. 2 Sarcee Canada 50 Montagnais Canada 33, 219 Sekani Canada 98 Mor Bolivia 169 Scnaca U.S.A. 70 Muckleshoot Canada 102 Seri and Yaqui Mexico 192,221 Muskhogean U.S.A. 112, 250 Shoshone U.S.A. 250 Sibundoy Colombia 198 Nab ua Mexico 221 Sioux U.S.A. Naskapi Canada 33, 183, 219 154, 157,250 Slave Canada Navajo U.S.A. 4. 6, 40, 42, 63, 87, 97, 98, 156 Stoney Canada 50, 97, 98, 155 191, 203 Subtiaba Nicaragua 165,175 Ne7, Puce U.S.A. 80 Swinomish U.S.A.; Canada 102, 103 Nootka Canada 102 Tacana Bolivia 169 Oajana Surinam 187 Tarahumara Mexico Oka nagon U.S.A.; 187,221 Tarasco Mexico Canada 102, 103 24.221 Tariana Brazil 31 Omaha U.S.A. 157 Tcwa (Pueblo) U.S.A. Ona Chile 211 4 Tinnch U.S.A. Otomi Mexico 187, 226, 229 206 Tlaxcalteca 144.xico Oyama Surinarn 257 227 Tlingit U.S.A. 62, 141, 187 Oyampi French Guiana 114 Toba Argentina 182,199,204,254 Paéz Colombia 10, 22, 131, 137 Toltec Mexico 270 Panzaleo Ecuador 241 Totonac Mexico 170 Paraujano Venezuela 16. 17.127, 134 Towa (Pueblo) U.S.A. 5, 6 Patax6 Brazil 208 Tru mai Brazil 249 Paya Honduras 165. 174 Tsimshian Canada 102,217 Pawncc U.S.A. 99 Tulalip Canada 102 Pehuciche Chile 189, 254 Tungurahua Ecuador 242 Pchuenche Chile 74, 254 Tuscarora U.S.A. 185 Penobscot U.S.A. 3 Uru Peru 267 Piegan Canw , 155 the U.S.A. 161, 163 Pijio Colombia 213 Pilagi Argentina 182, 199, 254 Wichita U.S.A. 99 Pima U.S.A. 42, 100 Winnebago U.S.A. 157 Ponca U.S.A. 250 Wintun U.S.A. 104 Pueblo New Mexico 4, 5, 6, 39, 40 Yagan Chile 75 Puruhi Ecuador 242 Yakima U.S.A.; Canada 102, 103 Quechua Ecuador; 3. 28, 93, 167, 168, Yamana Chile 74 Peru 214, 723, 244 Yaqui and Seri Mexico 192,221 Quillacinga Colombia 198 Yurok U.S.A. 104 Quilieute U.S.A.; Canada 102 Zapotec Mexico 64, 170, 187,260

184 theunknownnotTABLE uncertaintiesto exceed 14. amalgamations Samples 5 per introduced' cent. of "mixed" Asterisk in by the recent past,Indians, indicates fusion but i.e.. it apparent isof Indian feltlong-separate that 100populations for per studies centgroups. resultingIndian of genetic ancestry. from relationships recent, It known and amalgamation evolution of Americanof several Indian, . ,44. is recognized that many tribes listed in Tables 1-11 may have resulted from groups, in which non-Indian ancestry appears every effort must be made to avoid ArgentinaCOUNTRY Chaco*Chaco * LOCALITY Chorote,Chunupi, Chulupie,etc. Pilga, Toba TRIRE(S) SIZESAMPLE 545 55 REVERENCES 180182 SaltaMuyChaco * * Toba,Chaguanco.Puna,Mataco, etc. etc. etc. Chamococo, Chiri 209120194 78 205,181182180 225 Brazil MatoMataBahia,Salta, Grosso, Humahnaca R. Colonia Southernupper * Xingu '1 Arawak,Arawak,Pataxos,Quechua Carib, etc. Carib, Tupi Tupi, Trumai 5 o9?7 248,208244 249 66 VIIODloma Cniambia TolimaCaucaR.. G. doValley and Sul, ilaila Misslies 'Coyaima PijaoChibchanGuarani 439236107 213216 73 GuatemalaEcuador NearNot stated Quito Natagaima * SevenMixedFourPijao regions communities 200237281120 266263123 1 Mexico VariousChiapas,SixYucatan provinces Tabasco and Yucatan MayanVarious 6,662 250223124 218241, 24295 PeruParaguay MochaCentralChaco andhighlands Laredo NotMacaVarious stated 200800111 214265228 8 Surinarn ParamariboPa,amariboPuna REVIEWS: $9, 187, 230, 254 CaribArawakAymara and and others Quechua 500177 88 151 60