Biomedical Challenges Presented by the American Indian

INDEXED BIOMEDICAL CHALLENGES PRESENTED BY THE AMERICAN INDIAN

PAN AMERICAN HEALTH ORGANIZATION Pan American Sanitary Bureau, Regional Office of the WORLD HEALTH ORGANIZATION 1968 iNDEXED

BIOMEDICAL CHALLENGES PRESENTED BY THE AMERICAN INDIAN

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

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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 one-day session is held on a topic chosen by the Committee as being of particular interest. At the Seventh Meeting, which convened in June 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 of newly contacted Indian groups and those of groups well along in transition. This volume records the papers presented and the ensuing discussions.

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t PAHO ADVISORY COMMITTEE ON MEDICAL RESEARCH

Dr. Hernán Alessandri Dr. Alberto Hurtado Ex-Decano, Facultad de Medicina Rector, Universidad Peruana Cayetano Heredia Universidad de Chile Lima, Perú 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 4I Escola Paulista de Medicina New York, New York, U.S.A. Sao Paulo, Brasil

Dr. Roberto Caldeyro-Barcia Dr. James V. Neel Jefe, Departamento de Fisiopatología Chairman, Department of Human Genetics Facultad de Medicina University of Michigan Medical School Universidad de la República Ann Arbor, Michigan, U.S.A. Montevideo, Uruguay Dr. Marcel Roche (Vice-Chairman) Dr. Carlos Chagas Director, Instituto Venezolano de Investigacio- Chief, Brazilian Delegation to UNESCO nes Científicas Paris, France Caracas, Venezuela Dr. Philip P. Cohen Chairman, Department of Physiological Chemis- Dr. James A. Shannon try Director, National Institutes of Health The University of Wisconsin Bethesda, Maryland, U.S.A. Madison, Wisconsin, U.S.A.

Dr. René 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. Kingston, Jamaica

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

.i9 Dr. Bernardo A. Houssay Director, Instituto de Biología y Medicina Ex- Dr. Salvador Zubirán perimental Director, Instituto Nacional de la Nutrición Buenos Aires, Argentina México, D.F., México

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 Horwitz, Director

iii Special Session on

BIOMEDICAL CHALLENGES PRESENTED BY THE AMERICAN INDIAN Moderator: Dr. James V. Neel PARTICIPANTS

Dr. Tulio Arends Dr. James B. Griffin Hematología Experimental Museum of Anthropology Instituto Venezolano de Investigaciones Cien- The University of Michigan tíficas Ann Arbor, Michigan, U.S.A. Caracas, Venezuela Dr. William S. Laughlin Dr. José Barzelatto Department of Anthropology Hospital del Salvador The University of Wisconsin Universidad de Chile Madison, Wisconsin, U.S.A. Santiago, Chile Dr. Moisés Behar Dr. Miguel A. Layrisse Instituto de Nutrición de Centro América y Sección de Fisiopatología Panamái Instituto Venezolano de Investigaciones Cien- Guatemala, Guatemala tíficas Dr. Thomas A. Burch Caracas, Venezuela Clinical Field Studies Unit National Institute of Arthritis and Metabolic Dr. Rubén Lisker Diseases Instituto Nacional de la Nutrición Phoenix, Arizona, U.S.A. México, D.F., México Dr. William Centerwall Dr. Max Miller Department of Human Genetics Department 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 The University of Michigan Human Genetics Branch Ann Arbor, Michigan, U.S.A. National Institute of Dental Health Bethesda, Maryland, U.S.A. Dr. Juan Comas * Sección de Antropología Dr. Noel Nutels Instituto de Investigaciones Históricas Servico Nacional de Tuberculose Universidad Nacional Autónoma de México Ministério de Saúde México, D.F., México Rio de Janeiro, Brasil Dr. Edmundo Covarrubias Departamento de Genética Dr. Marcel Roche Facultad de Medicina Instituto Venezolano de Investigaciones Cien- Universidad de Chile tíficas Santiago, Chile Caracas, Venezuela Dr. José M. Cruxent Dr. Francisco M. Salzano Sección de Antropología Departamento de Genética Instituto Venezolano de Investigaciones Cien- Universidade Federal do Rio Grande do Sul tíficas P6rto Alegre, Brasil Caracas, Venezuela Dr. George Giglioli 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 - 2-

Page 42, Table 2. The following errors occur in names of tribes: in line 2, "Ticaque" should read "Jicaque"; in line 3, "Maya" and "Mazatel" should be separated by a comma; in line 6, "Mozatel" should read "Mazatel."

Page 43, Table 3, line 4. "Periri" should read "Pariri."

Rubén Lisker: Discussion

Page 45, Table 2. The number of cases of G-6-PD deficiency among the Huasteco should be 3 instead of 0.

Page 46, second paragraph of General Discussion. Dr. Lisker's remarks should read: "When I say there are two identical albumins, I am only saying that they have the same electrophoretic mobility. Nobody has done an amino acid sequence, so that is all we know. Whether or not it is the same mutation is unknown."

Francisco M. Salzano: Survey of the Unacculturated Indians of Central and South America

Page 62. In line 10, "Quiguisaca" should read "Quiquisaca"; in the section on Brazil, the reference to Ribeiro in line 4 should be 20 instead of 19, "Makirutare" in line 14 should read "Makiritare," and the first reference in line 24 should be 11 instead of 10; in the section on Guyana, the second reference should be "Information note in América _Indlgena, 23:158, 19_6.'j." ......

George G. Giplioli: Malaria in the American Indian

Page 107, last paragraph. In line 16, "malarioastic" should read "malario- metric"; in line 30, "Macushi" should read "Makusi."

José M. Barzelatto and Edmundo Covarrubias: Study of Endemic Goiter in the American Indian

Page 130, line 5. The figure "r=4 .72" should read "r=0.4 72." BIOMEDICAL CHALLENGES PRESENTED BY THE AqERICAN INDIAN (PAHO Scientific Publication No. 165)

CORRIGENDUM

The following substantive changes were submitted by authors of papers in this volume too late for inclusion in the printed text.

José M. Cruxent: Theses for Meditation on the Orip;in and Dispersion of Man in South America

Page 11, last paragraph, line 3. "Atrate" should read "Atrato."

Page 13, last paragraph. In lines 4 and 5, "scaly-surfaced stone artifacts" should read "crude chopping tools"; in line 22, "Intahuasi" should read "Intihuasi."

Page 16, reference 21. The publisher should be Vega, not Española.

William S. Laughlin: Discussion

Page 17, paragraph 3. The last sentence should read: "But at least the other groups that may have come in have left n-o distinguishable genetic heritage that can be sorted out."

Page 17, paragraph 6. The following sentence should be added at the end: "These last are foods easily accessible to women and children."

Juan Comas: Biological Subdivisions of the Indian on the Basis of Physical Anthropology

Page 22, paragraph 8, line 9. "Wisler" should read "Wissler."

Page 23, paragraph 8 (Stewart and Newman extract), line 2. "Radical" should read "racial."

Page 34, reference 58. The paper by Weiner appears on pages 441-480 of the volume cited.

Tulio Arends: Discussion

Page 41, last paragraph. References 9 and 10 should be followed by an additional reference: "Arends, T., D. A. Davies, and H. Lehmann. Absence of variants of usual serum pseudocholinesterase (acylcholine acylhydrolase) in South American Indians. Acta Genet. 17:13-16, 1967."

(OVER) CONTENTS

Opening Statement James V. Neel ...... I

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

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

Biological Subdivisions of the Indian on the Basis of Physical Anthropology Juan Comas ...... 22

Biological Subdivisions of the Indian on the Basis of Genetic Traits Miguel Layrisse ...... 35 Discussion Tulio Arends ...... 40 Rubén Lisker ...... 43 General Discussion ...... 46

The American Indian in the International Biological Program James V. Neel ...... 47 Discussion ...... 55 Survey of the Unacculturated Indians of Central and South America Francisco M. Salzono ...... 59 Discussion ...... 67

Medical Problems of Newly Contacted Indian Groups Noel Nutels ...... 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 Indians Thomas A. Burch, Leonard J. Comess, and Peter H. Bennett ...... 82

Hyperglycemia in Pima Indians: A Preliminary Appraisal of Its Significance Max Miller, Peter H. Bennett, and Thomas A. Burch ...... 89 Malaria in the American Indian George G. Giglioli ...... 104

v Food and Nutrition of the Maya Before the Conquest and at the Present Time Moisés Béhar ...... 114 lodine Deficiency Without Goiter in Isolated Yanomama Indians: Preliminary Note René Riviere, Dominique Comar, Mentech Colonomas, lean Desenne, and Marcel Roche ...... 120 Study of Endemic Goiter in the American Indian José Barzelatto and Edumundo Covarrubias...... 124 Discussion 1. D. Niswander ...... 133 Edmundo Covarrubias ...... 136 General Discussion ...... 137 Appendix: Tabulations of Phenotype and Gene Frequencies for 11 Different Genetic Systems Studied in the American Indian R. H. Post, I. V. Neel, and W. . Schuii ...... 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, to transition is accomplished the Indian may pre- present a wide variety of questions and chal- sent some rather special disease problems, 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 able 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 untouched-the 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 opportunities PAHO/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, with all its ramifications. The number of participants in the Session is somewhat greater than usual, thanks to the In the latter half of the program, we shall turn generous financial assistance of both the U.S. our attention to some of the more medical and National Committee for the International Bio- humanitarian problems presented by the Indian. logical Program and the Human Adaptability t, In many respects, these problems are similar to Section of the Committee for the International those of the campesino 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 at which 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 Biological Program. 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 Covarrubias have kindly clear and persistent. In addition, by taking it up agreed to discharge that function. we preserve at least temporarily one of the last great resources for the study of primitive man, one of the last opportunities to attempt to fathom

the nature of the forces to which man was re- 'The proceedings appear in Lije at High Altitudes, sponding during the course of human evolution. PAHO Scientific Publication 140.

1 THE ORIGIN AND DISPERSION OF AMERICAN INDIANS IN NORTH AMERICA

James B. Griffin

The historically effective discovery of the New tations and can be read by those who care to World by Europeans in 1492 opened the Ameri- do so. cas to large-scale colonization and to a long The temporal framework for early Indian groups period of speculation on the ultimate origin of 4 the New World peoples. For almost four hun- The time of arrival of the first human groups dred years the chronological framework was is far from settled and is likely to remain a based on biblical sources and was not adequate research problem for some time. For purposes to explain the cultural diversity of New World of this paper 1 shall refer to the period of time culture. Prominent among the speculations on from the first arrival of man to about 8000 B.c. origins were the multiple-migration hypotheses, as the Paleo-Indian period, and all cultural which provided civilized Mediterranean, Euro- complexes and skeletal material of this age, if pean, and even Southeast Asian groups to bring any, will be included within this period. A rapid a level of higher culture into the Americas and review of evidence from the United States and produce the more complex civilizations of Meso- Canada includes only one fairly homogeneous america and the Andes. Such explanations were and widely dispersed complex, a few examples also employed to explain a presumed multiracial of other early Paleo-Indian period peoples, and origin of the American Indian. a number of purported sites that are not re- As early as the sixteenth century some writers garded in this paper as soundly established. recognized the predominantly Asiatic relation- The Fluted Point Hunters are the sole recog- nizable group with a continent-wide occupancy ships of the native American populations and in the latter part of the Paleo-Indian period from postulated that the movement of people took about 10,000 to 8000 B.c. The earliest radio- place from northeast Asia to northwestern North carbon dates in association with an established America. This is, of course, the only route cultural context are about 9300 to 9000 B.c., of entry seriously considered by contemporary from sites in Arizona, New Mexico, Colorado, scholars, but there is considerable room for and Oklahoma. This is referred to as the Llano disagreement in many phases of the origin and complex, with the Clovis fluted point as a major dispersion of American Indian populations. For diagnostic tool. The Sandia finds are regarded this paper I have chosen to refrain from present- as a potential predecessor, with an uncertain age ing many reference citations to support the -probably within a thousand years of Clovis. In points of view included in the paper and from the western plains the later Folsom assemblages directly citing other views. Instead I have in- at several sites have been dated between 9000 cluded in the references a list of books and other and 8000 B.C. In the eastern United States and articles that afford a wide spectrum of interpre- Canada, fluted points are known from the South

(4 2 Atlantic and Gulf coasts to the Great Lakes and North America for which considerable antiquity Ontario and into New England, and as far 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 1 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 about 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 20,000 B.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 been Sheguiandah, Manitoulin Island, Ontario, for adequately demonstrated. Among these are the a cultural complex much before 7000 B.C.; for Desert Culture, Lake Mohave and the San a completely pebble-tool culture in northern Dieguito culture of southern California, and a Alabama of extreme antiquity; for a chopper/ number of sites and complexes from Oregon to chopping-tool complex of an interglacial 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 also in Central antiquity, but its age, except on typological America, there are indications of the presence grounds, is not known, and no archaeological of man before 8000 B.C., but many of these assemblage from Alaska is as old as 8000 B.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 there these finds from the Late Pleistocene Upper Be- is a date of 12,550 - 500 B.c. (M-1409) and cerra formation may well record human occupa- from Stratum E a date of 13,050 ± 800 B.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 Paleo-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 and at Tlapacoya in the Valley of Fluted points are known from Mexico and as Mexico. As usual, there is some 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 are high antiquity but not directly associated with J) 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 the islands. A radiocarbon date of 12,800+ 120 B.c. first inhabitants of North America. On the (GIN-97) has recently been obtained on fossil l north side of the Strait of Magellan, in Fells bone from the lower cultural level of Mal'ta Cave, there is a date of 8760 B.c. (W-915) ob- near Irkutsk, often attributed to the older Upper tained by Junius Bird. From eastern Brazil, at Paleolithic of the Irkutsk area. A date of 18,950 Lagoa Santa, there is evidence of occupation - 300 B.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 (14). date close to 6000 B.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 from the Uski I site. ciation of man and extinct fauna in the period Other sites in southwest Siberia are presumed to from 14,500 to 12,000 B.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 8000 B.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 12,000 B.c. have a strong early rela- fore 9000 to 8000 B.c., imply the arrival of man tionship to the late Mousterian stone industries 4 on that continent substanitially befor the kno.wn 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 8000 B.c. implies are sufficiently different from those of the better- that the arrival of the first human groups was known areas to the west that an easy alignment substantially before this date. If the proposed has 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 to The level of occupation is attributed to the find substantiating evidence in the rest of the lower section of deposits on Terrace II of the New World. The North American geographical Yenesei. The faunal composition represents cold spread of dated evidence and the considerable periglacial conditions, and the date corresponds diversity of assemblages shortly after 8000 B.c. to about the maximum of the last major Siberian imply an antiquity of man in North America glaciation, called the Sartan. Also in the Middle considerably greater than the known age of the Yenesei, at the Kokorevo sites, radiocarbon Fluted Point Hunters or of the occupants of dates range from about 14,000 to 11,000 B.c. Wilson Butte Cave. In summary, an age of about In the same area at the Mal'ta site, a date of 15,000 years for man in the New World is about 7000 B.c. was obtained on a Mesolithic- viewed as reasonable, with the possibility that like 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 cores, tions of Siberia is just beginning; in fact, ade- and some bifacially flaked points or knives. quate investigation of ancient man in north- There is a trend toward greater use of true eastern Siberia has only recently been initiated. blades made from prepared cores and toward the Most of the sites with an age of more than four fashioning of end and side scrapers, piecers, or five thousand years are along the southern perforators, gravers and burins, and an increase borders of Siberia from Russia to the Japanese in bone tools and ornaments. The animals on

4 which the people fed are those from arctic to Their 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 substantial part 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 woolly ments of Upper Paleolithic emphasis on blade 2, rhinocerus and mammoth. 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 man and shelter. This gradual expansion northward into North America awaited the presence of into new territories from northeastern Siberia arctic-alpine tundra species, during the late to Alaska would have taken place without resis- Pleistocene, on which man could live as 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, it is in favorable areas such as elevations overlooking certain that only a small part of the land mass passes followed by game animals in moving was glaciated and that most of the area was from one feeding ground to another. *$ occupied by xerophytic arctic tundra or alpine This early population spread is believed to tundra. A long tongue of steppe or periglacial have been diverted south along the west side of steppe extended from southwest Siberia east- the McKenzie Valley. A number of recent ward between the Central Siberian Plateau and papers (9, 11, 28) have emphasized the difficulty Lake Baikal as far as Yakutsk on the Middle of 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 Arctic United 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 B.c., with corridor between these sheets is not so firmly two periods of submersion of the highest part established that sound datings for these events of the shelf corresponding to major ice-melting are available. The position adopted in this paper phases of the retreat of the Wisconsin ice. The is 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 by a few thousand years, about 19,000 to 15,000 tundra vegetation similar to that of the Siberian years ago. last glacial domi- arid-steppe tundra. During the If the early hunters came into the United nance, between 23,000 and 10,000 B.c., 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 B.c. 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 time for penetration into 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. If the corridor was closed between

5 21,000 and 10,000 B.c., archaeologists are faced an early post-Valders invasion of the east by with at least as impressive dilemmas in the form animal forms now associated with western of an absence of sound data representing man in prairie environments, and this would correlate North America 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 cool prairie 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 retreat of the Wisconsin ice they penetrate this far south before the last would have had an effect upon the way of life Wisconsin advance, or did they arrive with the of ancient man through the shift of climatic "reopening" of the corridor ? zones, vegetation, and animal life. The expan- sions of the Canadian continental ice sheets The Fluted Point Hunters of North America effectively obliterated the vegetation and animal By 10,000 to 8000 B.C. the people of the Late life from much of Canada. The expansion of the Paleo-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 previous the Atlantic. Most of these populations were 4- periods into assembiages distinctive to late Wis- strongiy 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 '4 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 extensive meier in Colorado, Graham Cave in central and effective grasslands supported the large Missouri, and others prove that the meat diet grazing and browsing animals of the late 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 mammoth 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 the full glacial was not likely to stay restricted to a few classes environments by the time of the early Fluted of 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 4le 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 activities that were causing some shifts in hunting and collecting part 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 Paleo- 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 soils left 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 and the relative homogeneity of the implements prairie provinces, continues to reflect the exis- recovered implies a rather rapid spread of these tence of a hunting economy with bison as an early hunting people, and apparently into areas important supplier of food, tools, and clothes. not hitherto occupied. There is also the implica- At some locations there is clear evidence of mass tion that there would have been continuing con- killings, evidently the result of communal drives. tact between neighboring bands, perhaps for Indications of variability of animal food comes group hunting or other food-procuring activities from sites where giant beaver, pronghorn ante- at favorable seasonal locations or at locations lope, elk, deer, raccoon, coyote, and smaller favorable for shelter during the winter seasons. mammals as well as bison were part of the food Such collective activity would have permitted supply. At other sites there are indications of exchanges of new cultural developments-in grinding and milling stones, burins, sandstone terms of sources of food, raw materials, manu- abraders, and whetstones. facturing techniques, hunting technology-and West of the Rocky Mountains from around of people. It is doubtful that individual bands 8000 B.C., and continuing for many millenia, would have been isolated from other groups for archaeologists recognize the Desert Culture, 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-sharing These 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 America and 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 deduction of agriculture around 1 B.C., many re- veloping adjustment to the essentially desert gional cultural developments occurred as the environment, which supported only small bands Indian groups became more familiar with local and in which population density remained low resources and developed the knowledge for up to the historic period. It was not, however, a successfully exploiting them. As they did so, static complex, for many significant changes successful adaptations to particular environments took place in the technology, some of them rep- tended to restrict band and group activity to resenting almost continent-wide shifts in tool these environments and to produce a higher forms, and new weapons, and shifts in the level of exchange of 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 distinguishable and lakes, in upland forested or alpine areas, regional cultural traditions. and where minor shifting climatic patterns One of the best-documented cultural continui- allowed the expansion of desert bands into some- ties from the Paleo-Indian period Fluted Point times better watered areas or the penetration of Hunters to Iater 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 the same basic form rivers, 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 4 · tools. New tools appear, such as the specialized in otherwise exploiting the food supply asso- Cody knife, and new techniques such as the ciated with the streams and coastal areas. 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 Canadian It was to be a long time, however, before the

7 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 States a gradual transition is recognized in the Fluted Point, as a collateral contemporane- several areas from the Fluted Point Hunters ous variant, or as the result of the northern and complex 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 development 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- In southern California the San Dieguito 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 is the 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 of 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 of its own thousand years resulted in an unusually dense through the development of a series of wood- pópulation for a hunting-gathering population working tools and ground and polished stone in the oak-forest area of central California. forms. While regional specializations are present Southwestern Alaska was sparsely populated 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 were B.c. as a southern extension of the Desert Cul- the first Aleuts, as is implied, it would suggest ture. Certainly the general hunting-gathering that the Bering Strait area was occupied by pattern is similar, and there are 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. there is a considerable variety of assemblages The most important feature of the Mesoameri- reflecting inland and coastal developments with can scene is the early domestication of plants, continuing ties with Siberia, and also influences from Tamaulipas to Chiapas, which has been from northward-spreading groups primarily demonstrated where systematic efforts have 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 eastward to Marked population increase in some areas such provide the first successful occupation of the as the Valley of Mexico, the Valley of Oaxaca, eastern Arctic. The first Eskifio bands reached the coastal lowland of southern Vera Cruz and northern Greenland by 2000 B.c. Eskimo cul- Tabasco, and the Pacific Coast of adjacent tural traditions have a considerable time depth, Chiapas and Guatemala is observed by 1000 B.c.,

8 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 America was Summary east of the Rocky Mountains, and 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 North differentiation 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 Paleo-Indian period. There are no occurred from 30,000 to 40,000 or more years indications in the prehistoric record of any later ago, others believe it was from about 25,000 to substantial migrating 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 cultural 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 were additional 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 Mamut Fojil de Santa Isabel lztapan, Mexico, y America. Amer. Midland Nat. 79:247-250, 1968. Artefactos Asociados. México, D. F., Instituto Na- 9. HAYNES, C. V., JR. Fluted projectile points: cional de Antropoligía e Historia, 1955. 61 pp. Their age and dispersion. Science 145:1408-1413, (Dirección de Prehistoria, Publicación 1) 1964. 2. AVELEYRA ARROYO DE ANDA, L. Antigiiedad del 10. HAYNES, C. V., JR. Muestras de C-14, de Hombre en México y Centroamérica: Catálogo Tlapacoya,-Estado de México. Mexico, D. F., Insti- 4 Razonado de Localidades y Bibliografía Selecta tuto Nacional de Antropología y Historia, Boletín (1867-1961). México, D. F., Universidad National 29, 1967, pp. 49-52. Autónoma de México, 1962. 72 pp. (Cuadernos del 11. HOPKINs, D. M., ed. The Bering Land Bridge. Instituto de Historia, Serie Antropologíca, 14) Stanford, California, Stanford University Press, 3. CUSHING, E. J., and H. E. WRIGHT, JR., eds. 1967. 495 pp. Quaternary Paleoecology. New Haven and London, 12. IRWIN, C. Informe preliminar sobre las exca- Yale University Press, 1967. 443 pp. vaciones en Valsequillo. Mexico, D. F., Archivos del Instituto Nacional de Antropologia e Historia, 4. GOODLIFFE, E. M. Un sitio pleistocénica en 1965. Tlapacoya, Estado de México. México, D. F., Insti- 13. JENNINGS, J. D., and E. NORBECK, eds. Primi- tuto Nacional de Antropología e Historia, Boletín 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 IV: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 the lower levels of Wilson Butte Cave, South región de Chapala, Mexico. México, D. F., Instituto Central Idaho. Tebiwa (Idaho State Museum) 8:57, Nacional de Antropología e Historia, Boletín 18, 1965. 1964, pp. 1-6.

9 de Tlapacoya, México. México, D. F., Insti- 17. LORENZO, J. L. La etapa lítica en Mexico. tocénico e Historia, Boletín México D. F., Instituto Nacional de Antropología tituto Nacional de Antropología e Historia, 1967. 49 pp. (Departamento de Prehis- 29, 1967, pp. 37-41. de la Cuenca de toria, Publicación 20) 26. MOOSER, F. Tefracronología los últimos treinta mil años. México, 18. MAcDoNALD, G. F. Debert: A Paleo-lndian México para Nacional de Antropología e His- Site in Central Nova Scotia. Ottawa, National Mu- D. F., Instituto 4- 1967, pp. 12-15. seum of Canada, 1968. 207 pp. (Anthropological toria, Boletín 30, and Sunken Conti- Papers, 16) 27. WAUCHOPE, R. Lost Tribes and Method in the Study of the 19. MAcNEISH, R. S. Preliminary archaeological nents: Myth of Chicago investigations in the Sierra de Tamaulipas, Mexico. American Indian. Chicago, University Amer. Philo. Soc. Trans. 48, 1958. Press, 1962. 155 pp. in the New World: 20. MAcNEIsH, R. S. Ancient Mesoamerican Civi- 28. WENDORF, F. Early man Nat. 100:253-270, lization. Science 143: 531-537, 1964. Problems of migration. Amer. 21. MARTIN, P. S., and H. E. WRIGHT, JR., eds. 1966. Man in the Western Pleistocene Extinctions: The Search for a Cause. 29. WEST, F. H., ed. Early Anthropo- New Haven and London, Yale University Press, American Arctic: A Symposium. College, 10(2), 1967. logical Papers of the University of Alaska 22. MARTíNEZ DEL Río, P. Los orígenes ameri- 1963. American canos. 3a. ed. Mexico, D. F., Páginas del Siglo XX, 30. WEST, R. C. Handbook of Middle 1953. Indians. Vol. I. Natural Environment and Early Texas Press, 1964. 23. MASON, R. J. The Paleo-Indian Tradition in Cultures. Austin, University of Eastern Noril Arnerica. C i't AnrPL : 570 pp. 227-278, 1962. 31. WILLEY, G. R. An Introduction to American North and Middle America. 24. MAYER-OAKES, W. J. Life, Land and Water. Archaeology. Vol. 1. 1966. Proceedings of the 1966 Conference on Environ- Englewood Cliffs, New Jersey, Prentice-Hall, mental Studies of the Glacial Lake Agassiz Region. 526 pp. 414 pp. 32. WRIGHT, H. E., JR, and D. G. FREY, eds. The Winnipeg, University of Manitoba, 1967, .- Prince- (Occasional Papers, Department of Anthropology, 1) Quaternary of the United States. Princeton, 25. MIRAMBELL, L. Excavaciones en un sitio pleis- ton University Press, 1964. 922 pp.

10 THESES FOR MEDITATION ON THE ORIGIN AND DISTRIBUTION OF 1' MAN IN SOUTH AMERICA

José M. Cruxent

This paper does not constitute an extensive even 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 on be 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 just South 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 of one positive fact: even today we cannot be sure the origin of American man goes back to the that 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 should archaeology and other sciences, I believe that have issued a papal bull explaining that the South America was peopled not only overland 4 Indians were in fact human and could receive but 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 man could otherwise have crossed so many rivers, have been suggested, and most of them have swamps, and lakes with their entire families- been discarded as fantastic. Today we feel that women, children, and the aged-to occupy the we have long outlived that era, that our 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 Atrate 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 difficult to the Atlantic and back again. In this labyrinth it is for a present-day archaeologist to consult there is no sun, only a weak green light. At and study this copious literature, which not only every step is a river or a marsh. Whether is hard to find but is sometimes available only through the low ground or along the mountain in archaic languages. But I wonder whether crests, walking is extremely difficult; hunting prejudging all those ancient studies without would have been practically impossible. In the

11 "winter" or rainy season no one could have Normans. Apart from the reasons I mentioned moved. And there are many other Dariens, and above, there is the general discredit to which many rivers to cross. From my travels in South these theories are subject because they have been America I suspect that Paleo-Indian man must put forward by untrained persons guided by have had to go at least part of the way by water ignorance or by a desire to sell books. The scorn (and I make bold to suggest that he may have of 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 explain many of the critically, is understandable. But there are 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 rising water levels such studies as those of Hui-lin Li (11) or since Paleo-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 the a 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 Falcón 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 sparsely populated at the tion that was found off the Jamaican coast. I time. But if we accept the possibility of a water have heard that a Mexican publication, which route, we must also accept the possibility that I have not seen, mentions a small Roman statue some groups reached South America some other unearthed in that country. The newspapers are way than through Darien. forever reporting discoveries of ancient objects The "autochthonous" theory of American of European and African origin-discoveries, man, which originated in a misreading of Qua- however, that are seldom controlled or studied ternary fossil remains and in the belief that by professional archaeologists. I bring all this up Paleo-Indian immigration would have been im- merely to inquire whether it is proper for us to possibly difficult, seems now to have been re- consider such information a priori false and of jected-among other reasons because there is 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 be 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 this respect that, even to which we must find an answer, not only from though I cannot prove the contrary, I am not the 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 in so 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 enough atten- 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 navigators as 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 capons that housewives fatten Once again, therefore, I postulate navigation in their cellars'" (20). This probably happened as a possibility that could have spared the Paleo- on many other occasions, and not only at the Indian innumerable obstacles on his way and time of the Conquest. When I was exploring that furthermore would have permitted migra- the Orinoco Delta region I came upon a French- tory movements without requiring the existence man (he eventually confessed that he had es- of an ice-free corridor east of the Rocky Moun- caped from Devil's Island), living with an tains. Indian wife and mestizo children; it was only by his slightly lighter skin and his beard that I Origin and antiquity of man in South America was able to identify him. I have 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- ,j As can be seen, the cultural influence of the neled from north to south across the Isthmus of intruder is slight or nonexistent. His biological Panama. Evidence of this migration is limited, influence I cannot say-that 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 even that 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, no transatlantic historic times until the fifteenth century, boats routes have been proposed. However, extraordi- hugged the coasts, for the fear of the open sea nary similarities have been shown between the was great and navigation was rudimentary. But stone implements of western Africa and north- any storm might pull the boat out to sea, and eastern South America (5). These parallels are 'i then the crew had to resign itself to the currents much closer than between Paleo-Indian material and the wind. When it is recalled that some of from Siberia and from North America. the remote Pacific islands were peopled by the As to the antiquity of Paleo-Indian man- time the Europeans came, the logical conclusion using this term for a hunting and gathering cul- that man willingly or unwillingly made long ture that coexisted with animals now extinct journeys across the oceans cannot be avoided. and is known by its scaly-surfaced stone arti- The most fortunate encountered islands or conti- facts-no accepted chronology yet exists. We nents; some learned in time how to get back may estimate that the Paleo-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 date in South America canoes that traveled the Caribbean from island or Venezuela of 15,000 to 20,000 years ago. This to island, support the idea that sea journeys are latter date is calculated on the basis of finds for possible even in small, primitive boats. We also which the following ages have been obtained: s have the many incredible transatlantic voyages El Jobo complex, 10,000 years; Las Lagunas and of sportsmen (16) and contemporary political El Camare, more than 16,000 years; and Muaco, refugees who have brought small, crammed 14,740 to 16,580 years (21). 1 do not know boats across the ocean in search of freedom and of any earlier dates anywhere else in South a new life in America. I do not believe that America. 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 may well 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-weight 13,000 to 14,000 years old. For Argentina we - vessels such as gourds or skins, which when have a date of 7,970 - 100 at Ayampitin- empty could have been used as floaters. Intahuasi, and for Brazil a date of 7,500 at

13 Sambaqui de Maratua (18). Current excavations which I accept for both Meso-Indian and Neo- in Chile showing man in association with masto- Indian times. I may cite here-purely for refer- dons and a variety of other animals promise to ence purposes, since I am not sure of their produce extremely important documentation validity for the Caribbean-Hester'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 Paleo-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+60 years; Mor- western and southern South America, 8,000 to dan, 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-Indian 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 typology of the Alejan- not only logical but overly conservative and I drina complex can be related to that of the ' expect future studies to at least confirm them. El Camare complex, with large plano-convex knives, scrapers, chisels, and other tools. This Principal lithic complexes material has not yet been fully studied. The artifacts are made of excellent-quality flint and Lanning and Patterson (13) have proposed a are in strict accord with the typology of flint three-group typology for the South American industries, which is based in part on the nature . lithic complexes: (1) bifacial industries, (2) of the substance. The same is true of the Man- burin or graver industries, and (3) chopper in- zanillo and El Camare implements, dustries. An examination of the principal Vene- the first of which are of silicified wood and the second zuelan complexes shows that the typology of of quartzite. I am one of those who believe that country presents certain characteristics that that a 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 collections in the Venezuelan terials would have to produce different typolo- (. Institute for Scientific Research, Dr. Lanning gies. 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 projectile use of objects as implements in certain circum- points). He could not relate the Chuqui, Tor- stances. This may be the answer in the case of tuga, Red Zone, or Exacto (Ecuador) complexes Taima-Taima, which is completely atypical; ,. with those of northeastern South America, and on the other hand, it is just as likely that we call he found some of the Lagunas and El Jobo 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 orl 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 to jandrina people lived, for no fossil remains South America: the chain of Caribbean islands of 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 great position of having no erudite citations to make hunters, that if large animals were not to be in support of my arguments.) From this experi- found on land they may well have been found ence, I do not think we can discard the possi- at sea-perhaps manatee and seals, both of bility that there were Paleo-Indian groups with- which were abundant in the Caribbean in those out stone implements any more than we can ,times (7, 23). ignore the likelihood that they knew how to Nothing 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 known Paleo-Indians without a lithic culture, which I whether they reached Venezuela, though some call "primary culture groups," could well have slight contact between the Alejandrina complex, been the ancestors of certain groups of forest -br an older one of the same sort, and the El gatherers and primitive agriculturists in South Camare and Las Lagunas complexes would not America. Lithic culture evolved through a series have been impossible. Later groups may also of phases before arriving at triangular peduncu- have had some influence on the early Meso- lated points; it is significant that this tradition Indian or late Paleo-Indian complex of Canaima survives to the present day in our llanos and in Venezuelan Guiana (21), the Rio Claro com- savannas but that I have not found it in jungle 'plex in Brazil (4), and the Gran Sabana and or thickly forested areas. Paragua material. This could indicate that a diversity of diets I hope in the near future to be able to clear existed from the start or else that it came about up this problem of another route to eastern at the end of Paleo-Indian and the beginning South America in Paleo-Indian times. This of Meso-Indian times, when the great vertebrates 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 not know explanation of migratory routes in general. whether the jungle-dwellers descended from an- Before concluding, I wish to point out that other group of great hunters or from a group many times we fail to take into account a hy- that entered the New World already carrying a pothesis that should not be ignored in drawing primary culture. It is also possible that some conclusions. I am referring here to the possibility groups abandoned the flake industry in adapta- that there may have been Paleo-Indian groups tion to an ecological area that did not allow its without a lithic industry. I do not understánd use. Geography is not determining, but it un- why we always imagine Paleo-Indian man with doubtedly affects many aspects of the life of a stone implement in his hand; we are even primitive peoples, who obtain from their habitat presumptuous enough to demand a specific type the elements essential to their needs and their of implement, an officially accepted one. We security. must not forget that until very recently projec- In conclusion, I believe that many of the gaps tile points were the only implements recognized; we observe in Paleo-Indian migration can be all others were considered "blanks." For many logically explained by taking navigation into years I have been accumulating experience with account. I do not reject the idea of terrestrial an archaeologist's eye in many Indian commu- migrations; I wish only to complement it. In nities. This in vivo archaeology has given me any case, I am optimistic about the possibility many answers that would have been hard to of solving the problem of the origin and dis- come by from purely academic studies. (Of persion of man in South America, because most course, this often puts me in the unfortunate good archaeologists do not regard it as settled.

15. REFERENCES

1. BOMBARD, A. Naufrage Volontaire. Paris, 1958. Columbian transatlantic travel by Arab ships. Har- 2. CASAMIQUELA, R. G., J. MONTANE M., and vard 1. Asiatic Studies 23, 1961. R. SANTANA A. Convivencia del hombre con 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 antropología versity Press, 1964, pp. 23-81. física ante el origen del hombre americano. Sáo 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 preceramic. Occasional Papers Idaho State Univ. Mus. 21:78-82, 1967. 5. CRUXENT, J. M. Les Cultures lithiques Paléo- 4 Indiennes du Vénézuela: un curieux phenomene de 15. MENDEZ CORREA, A. A. Nouvelle hypothese * convergence typologique et fonctionnelle avec les sur le peuplement primitif de l'Amérique 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. 17. OSGOOD, C., and G. D. HOWARD. An Archeo- 6. CRUXENT, J. M., and 1. RousE. Arqueología logical Survey of Venezuela. New Haven, Yale Uni- de Venezuela. 2 vols. Washington, ' cronológica versity Press. (Publications in Anthropology, 27) Unión Panamericana, 1961. 18. PETo-r y (GAr-TA.,. mnerica indígena. 7. Enciclopedia Universal ilustrada Europeo- Tomo 1. El hombre americano. Los pueblos de Americana. Barcelona, Espasa, 1924, Vol. 24, p. 218. América. 2a ed. Barcelona, 1962. 8. GIMPERA, B. P. L'Amérique: Paléolithique et 19. REMAN, E. The Norse Discoveries and Ex- mésolithique. In André Varagnac, Paris, L'Homme plorations in America. Berkeley, University of Cali- avant l'écriture, 1959, pp. 165-187. fornia Press, 1949. 9. GRIFFIN, J. B. The origin and dispersion of 20. ROMOLI, K. Balboa of Darien: Discoverer of American Indians in North America. In Biomedical the Pacific. Garden City, New York, Doubleday, Challenges Presented by the American Indian. 1953, p. 94. Washington, Pan American Health Organization, 21. RousE, I., and J. M. CRUXENT. Arqueología 1968, pp. 2-10. Venezolana. Caracas, Ed. Española, 1962. 10. HESTER, J. J. Late Pleistocene environments 22. SAUER, C. O. Neblina norteña. 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. Mus. 1l. HUI-LIN Li. Mu-lan-P'i: A case for pre- Nat. Hist. 85, 1945. "'r

..4o

j 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 the They 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 the intertidal zone, and so neither will I. and, very importantly, invertebrates, mollusks, A great deal of ground has been covered very octopus, and so forth. 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 can the Aleutian Islands) and the Eskimos (of the make only a few comments and perhaps add one mainland extending over to Greenland) may or two observations by way of interpretation. have originated or taken their primary form on 1 would suggest that most or all of the people the 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 not simply backed up and remained on the coast. exclude other points of entry. But at least the Of course, as the waters rose there was more other groups that have come in have left no coastline rather than less, and therefore they distinguishable genetic heritage that 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, being separated by eters from north to south. This leaves, then, an several 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 of game hunters hunting mammoth, caribou, and i the differentiation between the less Mongoloid possibly musk-ox. These people may have lived i American Indians and the more Mongoloid Es- on the Bering land bridge 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 people One was to learn to live under water; the other r lived permanently on the land bridge. They was to go back to Siberia or on to the New were there for a long period of time. They did World. Obviously, a number of them went on not simply run across in order to get to the to the New World. 4 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 the period between they were unaware that they were going to be 9,000 and 12,000 years ago. There are a great submerged. So for a few thousand years they many dates that I think can be accepted as being enjoyed the real estate they held. well replicated. 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 is one that China, as was originally observed by Franz Dr. T. Dale Stewart called to my attention Weidenreich. This similarity probably indicates yesterday. Dr. Leakey has rediscovered an old some genetic continuity. If that is so, 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 suggests a slow rate, itself-not on associated materials-at 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 in a position to say that and between the inhabitants of the New World man had to come prior to the 14,000- or 15,000- and the Old may reflect a slow rate of humanr 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 look like Neanderthal man or like, Now, turning to South America, one thing Australians does not mean that they have not I should like to mention is the trait complex been here for some time. associated with the Lagoa Santa crania of Brazil. And there is something else to be emphasizedi- You will recall that these were discovered in the -that 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 Lag6a Santa crania are dated around 8760 B.C. in different tissue systems, are proceeding at The interesting thing is that 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 cannot assume 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 perhaps a 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, which are 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 the tym- filter. All botanists, glaciologists, geologists, and panic plate. And there are several cases of inter- anthropologists are agreed that, whatever condi- parietal bones, the separate Inca bone on the tions were, they were cold. The inhabitants posterior of the skull, which reaches a high adapted to the cold for the few or several frequency in South American Indians. thousand years they lived on the Bering land Taking this example to raise a 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-e logical change in American Indians over 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 same thing for North Ameri- other varieties of zones. But they all had to pass r can, though we do not have as good a series as through a cold filter. for Lag6a Santa. Nevertheless, the amount of The last point I should like to mention is the morphological evolution does not appear to have rate of dispersion and how long it would have 4 been great. taken to get to the Strait of.Magellan. I simply We can go one step further and point out that call your attention to the fact that the Aleuts the Eskimos and Aleuts, who form a single and Eskimos are spread over a difficult expanse 4 coherent population system, demonstrate a simi- of 8,000 miles, and it took them considerably larity to Middle Pleistocene man of northern less than 8,000 years-possibly as little as 2,000

18 years-to get that way. Therefore, the trip to What I asked is this: You seem to give the South America may not 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 G§eneral Discussion America. Could this be taken as additional evidence for a north-to-south migration? Salzano: Since Dr. Laughlin mentioned the Cruxent: Yes, it seems to confirm Lag6a Santa material, I want to say a few the theory that the migratory current ;words about the prehistoric remains of Brazilian was from north to south. Indians. Moderator: Professor A few years ago a researcher from the Na- Griffin, you have a name for being tional Museum, Professor Marilia de M. Alvin, a bit of a skeptic in these matters. Would you care to did a review of the anthropometric data at hand respond at all to Dr. Crux- and found a considerable heterogeneity between ent's presentation? .¿the different Lag6a Santa sites. We cannot say, Griffin: The evidence of man in South therefore, that these people are just one homo- America, both what can be established and what geneous group. She also examined the shell- has been claimed, indicates an antiquity of man 'nmound material of different locations and they in South America greater than that in North were even more heterogeneous. America, and evidence in which I have some Moderator: Do you mean to imply that the confidence suggests that we have not found man *,heterogeneity in the Lag6a Santa material is in North America of the antiquity necessary for greater than might be found in contemporary his appearance in South America at the age Indian populations in Brazil? claimed. I think it is possible that we simply ;i Salzano: That is difficult to say, because the have not found the evidence in North America. material available for study is not very extensive I should like to comment on one point of -only about 80-odd individuals. But even in Cruxent's paper: If through man's penchant for this small sample the heterogeneity is very great. self-destruction the evidence for our astronauts I am not saying, however, that it is greater than were completely destroyed, there would be very among present-day Indians. little in the earth's atmosphere to tell the people Roche: I have a question for Professor Crux- of the future that man had actually gone around :ent. He gives a possible age for man in North the world. Cruxent's "Paleonauts," if 1 may America of from 15,000 to 30,000 years, in coin a phrase, are extremely difficult to trace in western and southern South America of 8,000 the sea waters. So that while he hinted broadly to 14,000 years, and in northern South America at the possibility of man's making voyages from western Europe or western Africa of 10,000 to 20,000 years. These ages seem to and perhaps r go from more to less. Does he consider this as across the Pacific, my position is that he may Lone more proof of north-to-south migration? have done so but I have seen no traces of the Cruxent: I do not think the last word has yet boats, or whatever they used, and thus have been said as to ages. We have not been able to no evidence for such an interpretation. date the oldest complexes we have in South I also wish to comment on the presumed con- America, in the alluvial terraces at great alti- tact of Mediterranean, European, African, or 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 in Such 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 from other the North American type. Is there any the Old because now we can clearly document evidence that they came from two 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 is no, there is and in Mesoamerica. no evidence of two separate groups that can be ; ' It is true that a great many contact areas are distinguished today. However, the record in known. For example, there are stories of the north China and Japan does suggest 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) Eskimos and Aleuts look is in the area of Minnesota and Wisconsin. But more like Japanese and Chinese may reflect some of us are extremely skeptical about the either their recency or the fact that they were appearance of Roman coins or statues or various drawn up the north Pacific rim-Japan, the other objects, because they can be reasonably Kuriles, Kamchatka-and followed the coast explained on the basis of man's pack-rat and rather than coming from northern Siberia. souvenir compiex. There is no particular group that could be And so I admit to somewhat of a skeptical identified 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 Asia is that in our views we should take into account! with any degree of certainty. I am also told that that Japanese anthropologists have again pointed the linguistic diversity in, for example, the out that they think the Ainu of Hokkaido, in eastern United States could reasonably have northern Japan, are a variant of Mongoloids taken place within a period of some 10,000 to and 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 homogeneity 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 Waterlow: Dr. Laughlin stggested that two complexes that date back somewhere around groups of people came over by different routes- 20,000 years or so. These populations seem to northern and southern-through Alaska, one have moved primarily along the northern coastal

20 areas of Beringea into the North American primitive music shows much less variation and 'interior. The evidence is that the populations evolution than primitive languages, that origins Dr. Laughlin has been talking about with his can be recognized much better through music proto-Eskimo or prehistoric Eskimo of 8,000 than through language. Practically nothing has years ago or more are distinctly different from been done in this area. For what it is worth, he the earliest movers into the New World and feels that the Latin American music he has had a cultural complex formulated primarily heard, particularly the very primitive, such as l on the Pacific side of northern Asia, at a later that of the Caribbean, has very strong relation- time. It was chiefly the first group, moving into ships with northern Asia, particularly Japan. that the New World along the northern rim, Moderator: At this point we have succeeded and I think moved populated North America in getting the Indian to the Americas, although either on migratory-bird into South America, there is something less than unanimity as to wings, across water by navigation, or plodding how 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 in the Roche: 1 want to make just a brief comment American Indian. The distifiction between in relation to Dr. McDermott's question. There and "genetic" is of course artificial, have been studies on linguistics, but there have "physical" been very few comparative studies on the music since the basis of much physical diversity is however, e of the Indians. I have been told by the head of genetic. It is a convenient distinction, , the Institute for Music Study in Berlin that and we shall adhere to it today.

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

Juan Comas

In accepting the invitation to participate in their 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 had of the pre-Columbian population of the New for .me. I w1e realize, how.ever, that m-y kno.1!- 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), Alese I present my excuses in advance and ask for Hrdlicka (1912), and Arthur Keith (1948). leniency on the part of my readers. The opposite camp-composed of those who' For a better understanding of the value and recognized the existence of obvious biological scope of the principal biological subdivisions of differences 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 cluded, 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), Wisler 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 biological Count (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 t taxonomy of the American Indian? believers in the somatic unity of the Indian re- Let us examine these questions consecutively. mained a minority, in their forefront was Mor- ton, of whom Stewart and Newman (56) say Biological variability of American aborigines as shrewdly: against classical conception of "American homotype" Indeed, so great was his influence that he was responsible in large measure for the wide acceptance of ~ The excellent works of Newman (47) and the generalization embodied in Ulloa'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 1 Most of these are well-known general works and the eighteenth century that "If we have seen therefore do not appear in the References. For additional 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 the great a century, until Hrdlicka arose as the new had been definitely discredited. While are of this opinion, N champion of the homogeneity of the Amerin- majority of anthropologists some dis- dian. In summarizing his arguments Hrdlicka we must not overlook the fact that said in 1912 (32): "The conclusions are that tinguished investigators differ. Ashley Montagu the American natives represent in the main a (44), for example, has written: single stem or strain of people, one homotype." The American Indians exhibit a certain basic homogeneity He reaffirmed this thesis in 1928 (33) in an but at the same time are evidently characterized by an to the lack of - attempt to refute those who supported the racial equally certain diversity of types. Owing is impossible to say quite how plurality of the American Indian: the necessary data it 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 are in no case of sufficient upon which to erect a satisfactory account of the Ameri- weight to permit of any radical dissociation on that can Indian. (p. 465) ; basis. (p. 481) Some years later the same author does affirm This position had the able support of Sir (45) that the Eskimos of the Arctic coast of " Arthur Keith (36): North America form part of the group of Arctic differs in appearance Certainly the American Indian Mongoloids while the other American Indians from tribe to tribe and from region to region, but un- constitute another group composed of "an unde- derneath these local differences there is a fundamental groups of North, similarity. This, too, is in favour of descent from a termined number of ethnic single, small, ancestral community. (p. 218) Middle, Central and South America." in his important volume on human Nevertheless, little by little, the physical varia- Coon, (18) devotes barely two pages to the racial bility of the American Indian had shown itself races characteristics of the American Indians. He says: to be an undeniable fact, leading to differing 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 than many peoples very clearly and objectively: area. In fact, they are more uniform who occupy an area a tenth as large. All of this indicates Much progress has been made since the early days of that a relatively small number of peoples crossed the anthropology in America when it was assumed that all Bering Strait during the last part of the Wisconsin New Indians were essentially alike. The diversity of the 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- or nificance of the diversity in terms of the evolution goloid in general and despite some of their peculiarities development of types there or of the importation of in blood groups do not necessarily merit classification as preformed types from the Old World remains to be a subspecies of their own ... The American Indians ' clarified. (p. v) differ from the Asiatic Mongoloids mainly in that they flatness, particularly in the nasal skeleton, Stewart and Newman (56) expressed the same have less facial and a more variable skin color. There is no valid evidence the conclusion of their article: opinion at 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 the fact that the principle of Indian radical than the Bering Strait. (p. 152) unity rests almost solely upon the outer appearance of cases-and quite r living Indians. In so far as the Indians exhibit in com- In spite of these sporadic 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, -I believe it can be said that the beginning of long trunk, they can be scanty beard and a relatively the second half of the twentieth century coin- said to be uniform. . ... That the Indians, on the other cides with an end to the myth of the "Indian hand, are quite variable within this racial pattern, and recognition by the especially when comparisons are made in measurable homotype" and an explicit of the exis- <á dimensions, also has been shown. (p. 33) great majority of anthropologists of somatic and osteological variability and All this would lead one to think that from tence heterogeneity among the aboriginal groups of 1951 on, the theory of the American homotype

23 America. It is then necessary to establish the to localize "the sectors and zones where the origin and the causes of this variability, bearing effects of mixture and hybridization occurred," i in mind that it in no way exceeds the limits of although 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 the mental influences, which Stewart and Newman Amerindians are due to prehistoric immigration explained 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 must try to canists who had no classifications to justify, and ac- analyze. cordingly were more willing to admit that anthropometric determinations were not always stable 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. But what seems essential to me is a somewhat First, there is the proposal of those who accept u: the immigration of diverse human types, each more caefLul anaiysis NeCwi-nan's uown view (54), which he expresses thus: of which represents one of the existing Amerin- dian "races." Stewart and Newman (56) de- The adaptive responses of bodily form to environment in warm-blooded animals have led, largely in the last scribe it very clearly, attributing it primarily to century, to the formulation of several ecological rules. those who have been concerned with American . . .Extensive testing of these rules on human materials, racial taxonomy: however, has not been performed. For this reason, I have examined the applicability of two of the best vali- . . the classifiers of Indian types operated with a dated of these rules-Bergmann's and Allen's-to the strongly hereditarian bias, out of keeping with the main 2 body forms of New World aborigines. The principle stream of biological thought. If explanations were given behind both rules is that the maximum retention of body for their classifications, 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.... Implicitly, such explanations can be achieved by larger body size, Bergmann's rule disavowed the possibility that physical changes could holds that within a wide-ranging species, the subspecies have occurred among New World peoples. (p. 29) in colder climates attain greater size than 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 t' 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 objectivity distinct Amerindian "races"; and they evaluate a series of facts, some of which I wish to men- the greater or lesser importance that some 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 Hooton-concede 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 Review 1:108-140, 1877. implies agreement that in the new habitat new C. Bergmann, Ueber die Verháltnisse der Warmeoko- nomie der Thiere zu ihrer Grosse, Gottinger Studien racial types were formed. He even tries (34) 3:595-708, 1847.

24 there are a number of exceptions to these rules: their shorter stature, then, is attributable solely 10 to 30 per cent for Bergmann's rule, calculated to their short legs. This reduction of extremity ;only from subspecies in the most contrasting length 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 to fostering body-heat retention." As to the eastern Bergmann's rule among birds and mammals is Eskimos, whose stature and sitting height are larger than this. Rensch (51) mentions "for less 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 3 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 in uncertain food supply and periodic undernutri- certain parts of the Old World [Bergmann's and tion may cancel out the selective advantage of Allen's rules] do not seem applicable. Upon larger bodies in colder climates, but this 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 elsewhere is not yet ap- British Columbia, and the Yahgan and Alakaluf parent." To admit that Bergmann's rule is not of the Magellan archipelago. With respect to the operative in Africa south of the Sahara indicates last 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, its measurements indicate they are not particularly application seems doubtful in Europe, the Near shortlegged." 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 as being the ' ences of stature and body proportions in differ- cumulative effect of a number of genes whose ent regions of Europe, and among the Veddas 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 includes this study was conducted, the primary cause of nine maps showing the distribution in America variation in stature is genetic" (p. 110). j' of stature, sitting height (living), head size Barnicot (9), referring to geographical varia- (living), cranial module (skeleton), upper face tions in stature, says: r 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 it is 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 the world shows quate, but probably [italics mine] the maps no very striking regularities. Both tall and short peoples represent reasonable approaches to the real dis- are to be found in most of the major 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 cline with adaptive significance in relation

25 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 that the New World was peopled by successive I the world. migrations or infiltrations of physically differing Harrison (27) also writes, in dealing with peoples it is very likely that the American races 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 environmentally determined, since growth is profoundly World." This, generally speaking, seems accept- affected by nutritional state, and probably by climatic able; I have earlier made observations on this ' factors as well. Further, the nature of the variation pro- matter, with particular reference to the charac- duced by both types of factor tend to be the same. (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 precisely-quantitatively and qualitatively 4. Farther on Newman writes (p. 323): "Of -the influence that heredity and environment the remaining standard dimensions, only [italics (in their broadest sense) have exercised on the mine] head form and relative head height show present somatic variability of the American distributional patterns not readily interpreted as aborigines. adanptive ones Ind-ee 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: t cessive), broad nose (dominant) and narrow "Thus Marshall Newman 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 adaptive re- 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's eco- 47)-that of Imbelloni, who uses as his basis logical rules" (p. 262). And, as expected, he , the classifications 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 entered a huge cul-de-sac offering every variety acteristics, in addition to stature. Therefore, of environment and no forerunners to mix with. A re- Newman's statement does not exactly reflect construction of what happened thereafter takes into the real facts. account that the resulting population at the time of discovery constitutted a major isolate that was homoge- ~ 5. Finally, Newman states (pp. 323-324): neous, both phenotypically and genotypically. (p. 269; "From the foregoing, it seems clear that body italics mine)

26 I wish, however, to present arguments that Other authors confirm a divergence of opin- establish categorically the true scope of this ions about the applicability and importance of "geographical and climatic determinism." I have Bergmann's and Allen's rules to the biological already said above that according to Rensch the variability 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 is very The weight/temperature relationships here demon- high. strated suggest that Bergmann's rule is applicable to In an interesting paper on the same 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 series here considered Bergmann's rule might seem to our present knowledge the rules of Bergmann be applicable, this suggestion is refuted by more extensive and Allen appear to be of historical or descrip- material [italics mine]. Defined by weight, it is not only tive interest only and certainly are 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 to some extent also appli- ecological effect of this variable of man will be discussed. cable to man," but adds: This presentation attempted to show in a sketchy fashion and Allen's rules to man the following: The application of Bergmann's have been seriously and cogently questioned (Wilber), 1) The rules of Bergmann and Allen find little sup- on the ground that the inadequate data has been im- port as causal agents in modern studies of temperature properly interpreted, and that in any event man has regulation in homeotherms. never responded to his environment in a passive manner, formal examples often cited in favor 2) The various but has always done everything within his power to generalizations do not support the of these ecological control and shape the environment to his requirements. case of the climatic determinists. One is forced to con- But while this is undoubtedly true, it should be remem- clude that the rules just do not apply causally to animals. bered that ecological rules are generalizations to which 3) In man the ecological forces supposed to be acting exceptions can be found in every group, but that by and are not doing so; Eskimos were not cold, the skinny large they do apply to most populations of a species. aboriginal Australians were. The studies of Newman and of Roberts on New and 4) The rules of Bergmann and Allen have no causal Old World human populations lend strong support to role in the formation of racial differences in man. Such the view that ecological rules apply to man as well as use of these rules on the part of some anthropologists is to other animals. a source of misinformation and conjusion [italics mine]. 5) Some human groups have met 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- populations in general and continues: "That veloped specific heat-conserving functional changes with human body-size and shape tend to follow these no gross morphological changes; the Australian aborigines are an example. rules has been demonstrated in several studies. The mean body-weight of populations in hot Garn, on the other hand, concludes as follows regions is demonstrably lower than that in tem- a detailed criticism refuting Wilber (22): 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 are exclusive or nongenetic factors in climatic adjustments, he even major causes of the differences between geographical points out: races, or that the past-century formulations of Bergmann and Allen completely solve the problems of race 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 racial Certain 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 were gram (IBP), 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 arrives at the conclusion that such 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 much mental conditions and physical stress," and a larger question of how climatic selection would proposed 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;3 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 interpretable as their pre- and Pan American Health Organizations. They -dilections decreed. Nowadays these disputes may, I hope, agreed be bypassed. The rules attest in any case that the environ- that research on high-altitude peoples ment is important as an instigator of evolutionary 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 are rules 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 are in some ways as valuable as in America on human adaptability to different the rules themselves. The lesson to 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 Aymaras). Continuing the research program so this genetic-environment interaction is effected. successfully initiated by Carlos Monge, Hurtado, We have, in fact, found evidence for the existence of and other investigators at the Institute of Andean 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 collaborators are responsible for the most recent "feedback" circuit is the simple one as follows: (1) en- studies on acclimation and adaptability to dif- vironmental stresses produce developmental modifica- A1 tions; (2) the same stresses produce a natural selective ferent ecological environments. pressure which tends to accumulate 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 unstressed course of development; conditions-that 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 nurture-is still unre- ordinated adaptive modifications has been made easy. solved. Besides Dobzhansky, whose opinion is (p. 399) 3 Published by the Central Office of the IBP (7, Mary- Recently the problems of human adaptability lebone Road, London, N.W. 1).

28 quoted above, Neel and Schull (46) had already races or breeding isolates of many authors.... Conse- stated: quently, though the actual number of races in the world exists apart from observers, the number recognized de- It is therefore practically impossible when one is dealing pends upon the aims of the observers. (p. 90) with human populations to create situations which throw a sharply critical light on the relative importance 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 possible biological >. 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 populations-the presence of different subspe- beginning-that certain contemporary anthro- cific polytypical forms, regardless of origin and pologists, especially Montagu and Coon, take causes-is recognized and accepted, all efforts the view that the existing variability and bio- at a systematization or classification 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), or the "ethnic North 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, or the skepticism of Barni- would be Caucasoids of a particular kind had cot (10) about the possibility of defining the the New World been peopled by a small band human "races" with the necessary precision. The of Upper Paleolithic Europeans." Here I wish classic description of the "human race" based on to point out that the possible presence of Cauca- the typologist approach is of historical interest soid 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 the Apparently certain cultural features cited by fundamentally similar definitions of Dobzhansky Greenman (24) as persisting on the eastern (20), Laughlin (39), Garn (23), Mayr (43), coast of North America tend to confirm this Bielicki (13), and others. I should like, how- supposition. ever, to quote here some considerations pre- The classical 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 homogeneity-this despite ',. type. Races may be conti- pearance, i.e., of the same 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 be used at dif- gations of Henckel, Layrisse, Lisker, Loria, ferent levels of abstraction from a continental race down to a local race or even their tribal subdivisions. Local Matson, Neel, Robinson, Reynafarje, Salzano, races are composed of collections of family lines which Sandoval, Sutton, Swanson, Zepeda, and others constitute breeding isolates within the larger population. have made available greatly increased data on No arbitrary standard of magnitude exists for the size or number of differences which must exist for a group 4See my own Manual of 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 the New World. Comments" able. However, for research purposes it is necessary to (Current Anthropology, Vol. 5, No. 4, 1964, pp. 321- compare the smaller, constituent subdivisions, the local 322).

29 diverse antigens, hemoglobins, transferrins, and logically and serologically typical traits. Earlier, haptoglobins that demonstrate an obvious pheno- however (p. 120), he alludes to the fact that typic and genotypic variability in these aborig- some anthropologists find the morphological, 4 ines. This confirms the present conception re- serological, and biochemical differences between garding the evolutionary process of the human Mongoloids and American Indians insufficient species and the formation of races according to for separating them into two geographical races populationist and dynamic criteria. and join them into a single, polytypic geo- In this respect, I cite the latest conclusions graphical race. I agree with him that there are of Matson and collaborators (42) on South slight grounds for this racial hypothesis. America: 2. As for local races, Garn writes: "In con- Yet as pertaining to the present study, it seems that a trast to geographical races which are geographi- 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 are not completely Mongoloid, tions themselves. Whether isolated that the present Polynesian populations are a racial by distance, mosaic and that migrants from both west and east have by geographical barriers or by social prohibitions, contributed genes to present panmictic potpourri which local races are totally or largely endogamous, and is Polynesia. (p. 188) the very small amount of gene-flow ordinarily Since this variability of blood groups observed comes from contiguous and related local races" among Indian populations does not coincide (p. 16). with other morphological variations, one is led 3. What Dobzhansky defined as microgeo- to think that Garn's conclusion (23) is correct: graphical races and what Lasker prefers to call As with classifications based on morphological traits breeding populations are what Garn calls micro- rather than on the populations themselves, artificial races (p. 18); in these he makes manifest cer- "serological races" add nothing to human taxonomy. tain differences in the composition of a local The major use of blood groups in classification is 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. (p. 51) tions, still differ from each other in numerous ways." I shall now review in some detail the racial 4. With regard to the value and usefulness of taxonomy proposed by Garn (23) with reference these concepts, Garn explains (p. 22): "Geo- to geographical, local, and micro-races, begin- graphical races, local races and micro-races offer ning with his definitions and continuing with opportunities for very different investigations his subdivision of the aboriginal populations of in 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 the answer to different questions and the solu- by Rensch, is defined by Garn as "a geograph- tion to different ically-delimited collection of similar races," and problems of ongoing evolution he adds that "the existence of geographical races in man." is due, of course, to the great geographical bar- 5. In referring specifically to the populations of the New World-that is, the Amerindian riers, chief among them oceans, that formerly Ar limited the expansion and migration of local geographical race-Garn mentions a series of races and protected them from introduction of local races that "in strict contrast with geo- different genes" (p. 14). Garn's geographical graphical races are true evolutionary units. As race is equivalent to the concept of continental populations such local races evolve or have races. On this basis he divides mankind into evolved separately." He recognizes that "withl nine geographical races, only one of which-the such a diversity of local races, it is clearly impos- A Amerindian-he attributes to the New World sible 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 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 of ranging 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, the ranging 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 a 60° N to 18 ° N). number of isolated or semi-isolated populations as is true for the several Apache and Navaho Discussion groups now." An appraisal of new techniques for interpret- Only after these admonitions does Garn list ing 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 description of York in September 1949. I should like to start biological characteristics (pp. 144-146) is quite by 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 Garn takes a firm position what different interpretations. He indicated that some (pp. 128-129): "Once, local differentiation in re-evaluation of the several morphologic, metric and the Americas was attributed to successive waves genetic techniques was in order if the best results were to be obtained from of migrations. Today such diversity is generally 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, I have 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 4' reference to the region under discussion, because race has two uses, as 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 are, at best, interim structures for dealing with genetic and phenotypic dis- demonstrates their great biological (especially tances, and should be replaced by quantitative systems. somatic and serologic) heterogeneity. It may be hoped that the comparative method will be According to Garn's own definition, I find replaced by the more accurate method of mechanism it difficult to understand how it is possible to analysis. However, neither of these hopes are likely 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 appears that É. dates back 35,000 to 40,000 years. See Krieger, in the race concept will remain in human Prehistoric Man in the New World (J. D. Jennings and biology for many decades even though it will, un- E. Norbeck, eds., University of Chicago Press, 1964, doubtedly, be a constantly changing informational con- p. 45). struct. (p. 25)

31 I have expressed a similar point of view on biological heterogeneity as a consequence of the various occasions, in discussing the usefulness diverse origin of the immigrating contingents and importance of anthropometry and osteome- who peopled the New World some 40,000 years try in any attempt to determine the variability ago and also as the result of a process of adapta- between different population groups. I reiterate tion 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 racial history has ceased to be the chief hood the biological differentiation of the Amer- excuse for field anthropometry, microevolution-espe- 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 environments-has become the core of recent studies. be determined which is the more important. Schemes of measurement are being revised in terms of the 'hereditability' of somatic dimensions and factors of 4. The most widespread and best-known sub- 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. If, 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 distinguish the local races proposed for America. I have deliberately included in this paper many quotations-too many, the reader may 5. The Amerindian groups of local races pro- think-from different workers, so as to present posed 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 1. I consider acceptable the proposal of various agreement on the other hand, may reflect prob- anthropologists for uniting all the aborigines lems 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 biological investigations (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 this able differential biological base.

REFERENCES BAKER, 1. P. T. Racial differences in heat toler- 4. BAKER, P. T. The biological race concept as a -q( ances. Amer. 1. Phys. Anthropol. n.s. 16:287-305, research tool. Amer. I. 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- A Human Biol. 32:3-16, 1960. REATEGUI, J. KOLLIAS, and R. B. MAZESS. Regulación 4 3. BAKER, P. T. Ecological and physiological de la temperatura corporal en indios quechuas adaptation in indigenous South Americans. In P. T. nativos de la altura. Anales del Instituto de Biología 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. BUSKIRK, J. KOLLIAS, and Press, 1966. 541 pp. R. B. MAZESS. Temperature regulation at high alti-

32 tude: Quechua Indians and U.S. whites during total among Indian groups of the Southwest. Amer. J. 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 G.-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 pologie physique. VI Congres International des University Press, 1966. 541 pp. Sciences Anthropologiques et Ethnologiques I:471- 9. BARNICOT, 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 and environment: 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. [. 10. BARNICOT, N. A. Le probleme de la race dans Phys. Anthropol. n.s. 21:575-589, 1963. l'etat actuel des connaissances scientifiques. Rev. 30. HIERNAUX, J. Applicabilité du concept de race Int. Sciences Sociales, 17(1):90-92, 1965. a l'espece humaine. Rev. Int. Sciences Sociales 17(1): 11. BENOIST, J. Microraces et isolats. Rev. Int. 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):120-122, 1965. la formation des races. Rev. Int. Sciences Sociales 32. HRDLICKA, A. The problems of the unity or 17(1):96-98, 1965. plurality and the probable place of origin of the 13. BIELICKI, T. Typologistes contre population- American aborigines. Amer. Anthropol. 14:11, 1912. nistes et theorie genetique. Rev. Int. 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, 1. 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, 1. Nouveaux apports á la classifica- Viking Fund, 1951, pp. 1-68a. tion de l'homme américain. Miscelanea Paul Rivet 15. BOYD, W. C. Blood groups of American Octogenario Dicata 1:107-136, 1958. Indians. American [. Phys. Anthropol. 25:215-235, 36. KEITH, A. A New Theory of Human Evolu- 1939. tion. London, Watts, 1948, p. 218. 16. BOYD, 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. 4 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. DOBZHANSKY, T. Evolution, Genetics, and 40. LIVINGSTONE, F. B. On the non-existence of Man. New York, Wiley, 1957. 398 pp. human races. Current Anthropol. 3(3):279-281, 21. DOBZHANSKY, 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. L ' 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. Int. Sciences Sociales 17(1):126-127, 1965. 26. HANNA, B. L. The biological relationships 44. MONTAGU, M. F. A. An Introduction to

33 Physical Anthropology. 3d ed. Springfield, Illinois, 54. SALZANO, F. M. The blood groups of South Charles C Thomas, 1960. 771 pp. American Indians. Amer. [. 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 anthropologist's 46. NEEL, J. V. and W. J. SCHULL. Human view of the peopling of the New World. South- Heredity. Chicago, University of Chicago Press, western 1. Anthropol. 16(3):259-273, 1960. 1958. 361 pp. 56. STEWART, T. D., and M. T. NEWMAN. An historical resumé of the concept of differences in 47. NEWMAN, M. T. The sequences of Indian Amer. Anthropol. 53:19-36, 1951. physical types in South America. In W. S. Laughlin Indian types. C. H. Evolutionary adaptation. (ed.), Papers on the Physical Anthropology of the 57. WADDINGTON, In The Evolution of Life. Chicago, University of American Indian. New York, The Viking Fund, 1960, pp. 381-402. 1951, pp. 69-97. Chicago Press, 58. WEINER, J. S. Climatic adaptation. In G. A. M. T. The application of ecological 48. NEWMAN, Harrison, J. S. Weiner, J. M. Tanner, and N. A. rules to the racial anthropology of the aboriginal Barnicot, Human Biology. Oxford, 1964. New World. Amer. Anthropol. 55:311-327, 1953. 59. WEINER, J. S. Acclimatation et differences de 49. NEWMAN, M. T. Adaptation of man to cold climat: leurs effets sur les différences 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 variation and species formation. Proc. Linnean the origins of human types. Human Biol. 29:329- Society of London, 150th Session, 4:275-285, 1938. 336, 1957. .4 52. RENSCH, B. The laws of evolution. In The 62. Yearbook of Physical Anthropology, 4. New Evolution of Life. Chicago, University of Chicago York, Viking Fund, 1949, p. 33. Press, 1960, pp. 95-116. 63. YOSHIMURA, H., and J. S. WEINER (eds.) 53. ROBERTS, D. F. Body weight, race and climate. Human Adaptability and its Methodology. Tokyo, Amer. 1. Phys. Anthropol. n.s. 11:533-558, 1953. Japan Society for the Promotion of Sciences, 1966.

34 BIOLOGICAL SUBDIVISIONS OF THE INDIAN ON THE BASIS OF GENETIC TRAITS

Miguel Layrisse

Improved techniques for the identification of differentiate 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 a intertribal 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- stricted to 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 low frequency. 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 frequencies in 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 functioned as more or less teristics. effective impediments to gene flow. Larger tribes Several attempts have been made at classifying may be divided into smaller subtribes 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 also tendency to local endogamy. For instance, in 4' 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 were contracted "American group" of a single race-the "Ameri- between individuals of the same local group. can Indian." Boyd suggests 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 local endogamy Considered as one population, American In- were observed, although under different cultural dians in general are characterized by a high fre- and environmental conditions, by Salzano, Neel, quency of blood groups O, M, R2, Fya, Dia, and and 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- ~k- hemoglobins (3, 21, 22). These characteristics teristics of the Amerindian is needed so that we

35 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 and in Peruvian tribes; and Matson et al. (20) in (b) the intratribal distribution of genetic traits. Chilean tribes. Córdova, 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 intertribal 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 e' two tribes. The Yanomama are a Paleo-Indian village populations, were examined for 7 blood- r society of hunters and gatherers with a rudi- group systems, 7 serum protein traits, and 7 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 of stances. Genes with low frequencies-MS, NS, interbreeding between the Makiritare and the R2, Lpa, and Aga-were found to have been Yanomama, the actual gene flow between them lost in several villages, while genes with high can be considered extremely limited. The com- frequencies-PGM and Se-were fixed in some parisons of blood-group frequencies between the villages. Genes with intermediate frequencies, Makiritare and two Yanomama subtribes (Sa- such as R1, Ms, Ns, Fya, and Jk%, 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 (63 provide another illustration of gene frequency 4 per cent) but only 19 per cent in the Waica and variation within the same tribe. Three villages 9 per cent in the Sanema. R,, 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 R2 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 Rl; and negative or very in the Waika and Sanema. Dia is 16 per cent low frequency of Dia . 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, differences-though less marked- traveling with gene M. It was also found that 4 in blood groups have been reported by Matson these tribes had 3 per cent of Dia. The genealogy and Swanson (17, 18) between Maya and non- of the Sacupana village gave a clue to the ex-

36 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 anthropological and medi- positive cases found among the Warao of the cal studies. Demography-including 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 genealogy-provides 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 Dhi, 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 in the Warao, for instance, is the 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, it is likely that both Dia and Dhi 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 et al. (9) showed a variation of 1 to 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 R2; the other gene frequencies It is evident that in the formation of the new had comparable results. village, as previously described, there are circumstances that may induce variation of the gene Comments frequency in the following generations-the 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- unit; it not always correspond to the biological planation of the variation of gene frequencies in appears that the population of each village rather Indian populations than other genetic forces than the whole tribe represents, in some cases, such as selection and gene flow. It is also prob- an independent gene pool. Therefore, it is not able that this dispersive process may operate studies carried out surprising that independent similarly in the formation of hybrid populations, in a tribe may show great divergence in gene explaining, in some cases, gene frequencies equal Y frequencies because they are performed in dif- to or even higher than those observed in the Vr 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 is inadequate or operating 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 studies on mortality, 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 which it possible to elucidate the role of selection in more than 70 per cent of the population was the variation of gene frequencies in Indian examined. But taking the test results of one local populations.

37 Bearing in mind this variation, I should like 3. It might be possible to make a first sub- to discuss finally the possibility of a subdivision division of the Middle American Indians: the of the American Indian populations that might northern group including Totonacs and Mayan reconcile somewhat the frequency of blood traits families, and the southern group of Chibchan with other anthropological characteristics. Dr. or Hokan affiliations. However, the only differ- Juan Comas has previously described here some ence found to exist between these is the frequen- of the attempts by prominent Americanists to cies of Dia, which tend in general to be high classify Indians. However, none of these classifi- (from 4 to 18 per cent, with a mean of 10 per cations fits the distribution of blood traits. Garn cent) in the northern group, and low (from (8), for instance, included all South American 0 to 10 per cent, with a mean of 3 per cent) in Indians except the Circum-Caribbean and Fue- the southern group. guian in one homogeneous population. It is 4. The geography of the South American known that in such a large area there are popu- continent favors the formation of microdifferen- lations with marked morphological, cultural, tiated populations. But in spite of the variations and serological differences. in gene frequencies between subtribes and vil- Even though it is very difficult to attempt a t4 comprehensive subdivision of the aborigines of lages of the same tribe, it might be possible to America on the basis of genetic traits, the fol- identify genetic peculiarities that characterize iowing taxonomic considerations can be formu- the tribe as a whole with a common genetic lated from the information available: heritage different from that of other tribes or 1. It is very likely that the Eskimo can be clusters of tribes. For example, it would be distinguished genetically from the other North interesting to investigate the relatively high fre- American Indians. They are characterized by quency of A1 among the Andean Indians, first the presence of the gene B and absence of the noticed by Newman (24), which is not accom- gene Dia. In frequency of the other blood traits panied by the presence of other non-Indian they do not differ greatly from American genes; the possible connection, on the basis of Indians. genetic traits, between the Chilean Indians and 2. The various North American tribes are the Polynesians, as mentioned by Matson et al. very difficult to analyze, partly because of heavy (20) in a recent study; and the possibility that non-Indian admixture. They have been classified the Yanomama and Warao, distinguished by the in one group characterized by the presence of absence of the gene Dia and by anthropological gene A and a low frequency of Dia; however, traits, could be classified, separately or together, microdifferentiations are observed in some tribes. as different from the neighboring tribes. The Blood and Blackfeet of Alberta show a very It is highly probable that our present hesita- high (60 per cent) frequency of A1 (6), which distinguishes this group from the rest of North tion in subdividing the American Indians on the American Indians. An exploration of the pos- basis of genetic traits will be overcome in the sible connection between Navajo, Yaqui, and near future with the increase of gene markers Lacandon tribes exhibiting the presence of and the use of mathematical models in which transferrin BO-1 would be of great interest (1, all the variables found in a tribe can be com- 26). puted.

REFERENCES

1. ARENDS, T. and R. LISKER. Unpublished infor- tribal genetic differentiation among the Yanomama mation, 1964. Indians of southern Venezuela. Proc. Nat. Acad. Sci. 2. ARENDS, T., G. BREWER, N. CHAGNON, M. L. 57:1252-1259, 1967. GALLANGO, H. GERSHOWITZ, M. LAYRISSE, J. NEEL, 3. BOYD, W. C. Anthropologie und Blutgruppen. 4 D. SHREFFLER, R. TASHIAN, and L. WEITKAMP. Intra- Klinische Wochenschrift 37:993, 1956.

38 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 hereditary blood antigens among American ' supplements definition from morphological char- Indians in Middle America: Lacandon and other acters. Science 140:1057-1064, 1963. Maya. Amer. 1. 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. ,4m. 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. ROBINSON. Distribution of Alberta, Canada. Amer. 1. Phys. Anthropol. 11:369- hereditary blood groups among Indians in South al, 384, 1953. America. Am. J. Phys. Anthropol. 27:157-194, 1967. 7. CORDOVA, M. S., R. LISKER, and A. LoRIA. 21. MOURANT, A. E. The Distribution of 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. J. 22. NEEL, J. V., and F. M. SALZANO. A prospectus w Phys. Anthropol. 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. 23. NEEL, J. V., and F. M. SALZANO. Further 'J Illinois, Charles C Thomas, 1965. 155 p. studies on the Xavante Indians. X. Some hypotheses- 9. GERSHOWITZ, H., P. C. JUNQUEIRA, F. M. generalizations resulting from these studies. Amer. SALZANO, and J. V. NEEL. Blood groups and ABH- [. Human Genetics 19:554, 1967. Le' secretor types in the Somóes Lopez and Sáo 24. NEWMrAN, M. T. A trial formulation present- Marcos Xavantes. Amer. 1. Human Genetics 19:502, ,*) 1967. ing evidence from physical anthropology for migration from Mexico to South America. In R. H. 10. GREENBERG, J. In J. H. Steward, and L. C. Thompson (ed.), Migration of the New World Faron, (eds.), Native People of South America. Culture History. Tucson, University of Arizona, New York, McGraw-Hill, 1959, p. 22. 1958. (Social Science Bulletin 27) 11. LAYRISSE, M., Z. LAYRISSE, and J. WILBERT. 25. PARKER, W. C., and A. G. BEARN. Haptoglobin Blood group antigen tests of the Waica Indians of and transferrin gene frequencies in a Navajo popu- Venezuela. Southwestern f. Anthropol. 18:87-93, lation. A new transferrin variation. Science 134:106- 1962. 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 antigen studies of four Chibchan Santa Catarina, Brazil. Amer. 1. 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. i1 1966, p. 72. I. Demographic data on two additional villages: 15. LAYRISSE, M., C. PEDREIRA, and J. WILBERT. Genetic structure of the tribe. Amer. J. 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. 1. Phys. Anthropol. 6:236, 1948. > acteristics in a malarial region. Amer. f. Human 31. WILBERT, J., and M. LAYRiSSE. The Warao Genetics 17:179-187, 1964. Indians. In preparation. 17. MATSON, G. A., and J. SWANSON. Distribution 32. ZERRIES, O. Waika. Die Kulturgeschiechtliche of hereditary blood antigens among Maya and non- Stellung der Waika-lndianer des oberen Orinoco im Maya Indians in Mexico. Amer. J. Phys. Anthropol. Rahmen der volkerkunde Sudamerikas. Munchen, 17:49-74, 1959. 1964.

39 DISCUSSION

Moderator: Dr. Comas' paper emphasized the As Dr. Comas pointed out, to determine which variability existing in the American Indian on 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 Indian lation of America derived from a small group that is increasingly occupying the thinking of of hunters-and-gatherers from eastern Siberia those of us who work with the Indian: that who crossed the Bering Strait during the latter within a relatively undisturbed tribe, there is a part of the Wisconsin glaciation and whose great deal of genetic variability from one part descendants gradually became masters of an of the tribal distribution to the next. The geneti- uninhabited continent, we can understand bet- M4 cist has tended in the past often to think of a ter the great influence of environmental factors tribe as a kind of homogeneous beanbag, but upon the American Indian. There is still the it now becomes clear that this is 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 two papers will be The great progress made lately in the study of opened by Dr. Arends. blood groups, serum protein groups, and enzyme polymorphism nullifies the idea expressed some Tulio Arends years ago: "Type one Indian and you have typed them all; so much are they alike in their blood The physical variability of the American groups."2 The view that serology has not added Indian was recognized even by the conquista- anything new to human taxonomy3 is also no dors. In 1530, Federman described a tribe of longer tenable. On the contrary, we who work dwarfs 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 will play an important queri in the eastern part of the country to be role in the reinterpretation of the American "people as black as coal."' Anthropometric Indian taxonomy and is laying the foundations studies have accumulated a great many data on for a future molecular anthropology. Boyd has stature, cranial and cephalic indexes, body pro- said: "The genetic method has, in short, made portions, and so forth, which multiply these contributions to anthropology that differences. If to the somatic characteristics are morphologic methods could added the many inherited biochemical and hardly have made, and is pro- hematological variations known today the diversity of the American Indian becomes infinite. 2 Stewart, T. D. A physical anthropologist's view of the peopling of the New World. Sout/hwest. 1. Anthropol. 16:259-273, 1960. 1Federman, N. Historia indiana. Transl. J. Friede. 3 Garn, S. M.: Human Races. 2nd ed. Springfield, Madrid, Aro, 1958. Illinois, Charles C Thomas, 1965.

40 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 establish biological subdivisions of the From a genetic standpoint, most of the Ameri- American Indians based on the study of their can Indians share certain characteristics that genetic characteristics, the best-known methods differentiate them from other racial groups of population 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 authentic South 1. It is advisable to study as many genetic American Indians are all of group O; usually traits as possible, in order to avoid biased inter- lack r and R° gene complexes of the Rh system; pretations and rule out gene drift, especially are 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 other Hb-A; and are not deficient in glucose-6-phos- events that might have modified the frequency phate dehydrogenase. 7, 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 E1a.s' 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. J. Hum. Genet. 19: gene introduced through a nonapparent ad- 554-575, 1967. mixture. 7Mourant, A. E. The Distribution of the Human 3. The "fission-fusion" phenomenon, which Blood Groups. Oxford, Blackwell, 1954. s Arends, T. Haemoglobinopathies, thalassaemia and according to Neel et al.,5, 6 is not infrequent in glucose-6-phosphate dehydrogenase deficiency in Latin America and the West Indies. New Zealand Med. I. 4 Boyd, W. C.: Genetics and the human race. Science 65:831-844, 1966. 140:1057-1064, 1963. 9 Gutsche, B. B., E. M. Scott, R. C. Wright. Hereditary 5 Neel, J. V., F. M. Salzano, P. C. Junqueira, F. Keitter, deficiency of pseudocholinesterase in Eskimos. Nature and D. Maybury-Lewis. Studies on the Xavante Indians 215:322-323, 1967. of the Brazilian Mato Grosso. Am. ]. Human Genet. 1o Motulsky, A. G. and A. Morrow. Atypical cholines- 16:52-140, 1964. terase gene E1 a: rarity in Negroes and most Orientals. 6 Neel, J. V., and F. M. Salzano. Further studies on Science 159:202-203, 1968.

TABLE 1. Characteristics common to American Indians

CHARACTERISTICS

COMMON TO ALL

GENETIC SYSTEM SUBJECTS EXCEPTIONS

ABO Group O 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-) Dieguefio (U.S.A.); Brazilian (Brazil) Hemoglobin Hb-A Nahua, Chiapas (Mexico); Oyana, Arawak (French Guiana) G-6-PD Absence of deficients Oyana, Carib (Dutch Guiana)

41 Where published reports give different char- greatly upon the condition of the sample ob- acteristics from those in Table 1, the reason is tained. 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 also here 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 this Differential characteristics of some Indian tribes could be the so-called heterogeneity of American Finally, there are some genetic markers that Indians. (The same may be said of the figures identify 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 Dehi. 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-B _ ) that distinguishes the North racial groups (Table 2). The interpretation here 0 and Central American from the South American is really difficult because different samples from Indians.l 2 the same population have sometimes given dif- Human alloalbuminemia, or the presence of ferent results. For example, when the hapto- albumins with electrophoretic migration differ- globin system of two Makiritare samples was ent from the normal, expressed by either one studied,1l results for Hpl as dissimilar as 0.610 and 0.342 were found. These results depend 12 Arends, T., G. Brewer, N. Chagnon, M. L. Gal- lango, H. Geroshowitz, M. Layrisse, 1. Neel, D. Shreffler, 1i Arends, T. Los grupos séricos humanos determina- R. Tashian, and L. Wietkamp. Intratribal genetic differ- bles por métodos electroforéticos: su distribución en Sur entiation among the Yanomama Indians of Southern América. Acta Cient. Venez. Suppl. 3:162-178, 1967. Venezuela. Proc. Nat. Acad. Sci. 57:1252-1259, 1967.

TABLE 2. Quantitative differences in some genetic systems of American Indians

GENETIC SYSTEM CHARACTERISTIC EXCEPTIONS

MNSs Gene M ~ 0.7 Jicaque (Honduras)

Rh Gene complex R2 > 0.35 Seneca, Pima (U.S.A.); Ticaque (Honduras); Miskito, Sumo (Nicaragua); Maya Mazatel (Mexico); Ticuna r (Peru); Quechua (Ecuador); Xavante (Simoes Lopes, Brazil); Waica (Venezuela)

Duffy Gene Fya : 0.6 Guaymi (Panama); Jicaque (Honduras); Mozatel (Mexico); Xavante (Brazil) ABH Secretor Gene Se > 0.8 Xavante (Brazil)

Haptoglobin Gene Hp' % 0.5 Yupa (Venezuela); North Athabascan, Tlingit, Navajo (U.S.A.) Gammaglobulin Gene Gma 1l.0 Xavante (Brazil)

Inv Phenotype Inv(a+) > 0.6

Gc Gene Gcl > 0.65 Xavante (Brazil)

42 TABLE 3. Genetic markers in some Indian tribes

> GENETIC SYSTEM CHARACTERISTIC TRIBES IN WHICH IT HAS BEEN FOUND k Diego Di(a+) North American, Central American, South American Lacandon, Transferrin Tf Bo_ 1 Navajo (U.S.A.); Yaqui, Cora, Chinantec, Tzotzil, Zapotec (Mexico) Quiche (Guatemala)

Tf Dchi Irapa, Macoita, Paraujano, Periri, Shaparu, Motilon, Piaroa, 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 Warao (Venezuela)

(homozygous form) or two components (het- 3. Some genetic markers-for 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 Dchi and B0,-, and albumin Naskapi, Mexico, system, which at present is the subject of inten- Maku, and Warao-have been found in only w sive study, has made it evident that certain some Indian tribes. The characterization 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 albumin slower than normal (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- ferrin Dchi, 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 fusion. 1. 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 Indians before these populations dis- investigated by the best serologic, immunologic, appear through cultural and economic pressures electrophoretic, and chromatographic methods or through epidemic, endemic, or sporadic / at hand in order that characteristics reported in pathological processes. the past, when modern methods and reagents were not available, may be rechecked. Rubén Lisker 2. The differentiation of populations by frequency gradients of genetic traits should 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. Since lations do not have a homogeneous genetic these populations have been under physical, constitution. However, the task of subdividing pathologic, and war pressures, it will always be them on the basis of genetic traits is not an very difficult to evaluate how much these agents easy one, especially if to taxonomy is added an have influenced the distribution of the inherited interest in interpreting or understanding the variations. reasons for the differences encountered. A gen-

43 eral cause of this difficulty is that the Amer- TABLE 1. Frequency of Diego antigen in 18 Indian indians as a group have characteristic gene tribes and 9 coastal mestizo groups of Mexico frequencies for many traits that do not allow PHENOTYPE FREQUENCY further subclassification. A good example would GROUP RANGE (%) (%) be blood group O and the Rh-negative phenotype. A reasonably "pure" Indian population Indiansa 5 tribes 2-8 should have close to 100 per cent of group O 9 tribes 12-19 individuals and no Rh-negative subjects; devia- 3 tribes 20-24 tion from this pattern would be generally 1 tribe 40 interpreted as the result of non-Indian admix- Mestizos b ture. West Coast Even so, some genetic traits do show enough Ometepec 28.2 variability in apparently unmixed Indian groups Cuajinicuilapa 26.3 San Pedro M. 23.4 that attempts at taxonomy are possible. The Pochutla 31.3 difficulty here-and I should like to stress this East Coast point-is what the classification means. I feel Saladero 4.1 sure that this consideration explains, at least in Veracruz 32.8 part, Dr. Layrisse's understandable hesitation in Paraiso 32.! subdividing Amerindians. El Carmen 38.6 Tamiahua 3.5 I want to consider briefly some examples of the difficulties I have mentioned. The Diego a More than 50 individuals studied in all but two antigen appeared to be an excellent marker of tribes. Indian groups, especially suitable for taxonomi- b More than 90 individuals studied in each location. cal purposes since it is not present in other ethnic out of the nine hybrid groups studied on the groups (except some Mongoloids) and shows coasts of Mexico have higher Diego frequencies enough intertribal variability to allow for sub- than most of the pure Indian tribes. Before divisions.' However, it soon became apparent learning the meaning of this phenomenon, one that the correlation of its distribution with would hesitate to use this marker to subdivide either geographical or cultural criteria was not Amerindians. Incidentally, 1 would say that too good,2 and furthermore, although its ab- 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 certainly no pattern to Diego frequency because individ- contribute to the gene pool have higher Diego uals with the same linguistic affiliation show frequencies than the original Indian popula- very marked differences in frequency. tions.a3 4', This is clearly seen in Table 1; seven An additional difficulty stems from the fact that in the study of Indian tribes with some 1 Layrisse, M., and J. Wilbert. El antígeno del sistema degree of non-Indian admixture, the variation sanguíneo Diego. Caracas, Sucre. in the gene frequencies of some of the traits 2 Comas, J. Significado de la presencia del antígeno does not follow the expected trend. The follow- Diego entre los Amerindios. Anal. Antropol. (Mexico) ing examples will clarify this statement. In the 2:89, 1965. 3 Lisker, R., A. Loría and M. S. Cordova. Studies on investigation of Mexican Indian groups, four several genetic hematological traits of the Mexican popu- gens in Nahuas, Yaquis, Tarahumaras, Tarascos and lation. VIII. Hemoglobin S, glucose-6-phosphate dehy- Mixtecos. Hutm. Biol. 35:359, 1963. drogenase deficiency, and other characteristics in a ma- 5 Cordova, M. S., R. Lisker and A. Loría: Studies on larial region. Am. 1. Hum. Genet. 17:179, 1965. several genetic hematological traits of the Mexican popu- 4 Rodríguez, H., E. Rodríguez, A. Loría, and R. Lisker: lation. XII. Distribution of blood group antigens in Studies on several genetic hematological traits of the twelve Indian tribes. Am. I. Phys. Anthropol. 26:55, 4d Mexican population. V. Distribution of blood group anti- 1967.

44 had evidence of Negro admixture, as judged by the slow variant known as albumin Mexico 0 the presence in them of a few individuals with was present in southern North American In- > the sickle-cell trait and/or red cell glucose-6- dians and in some Mexican Indian groups and phosphate dehydrogenase (G-6-PD) deficiency. 6 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 variants were found except in the last , these expectations was confirmed.5 It would be group.1l This pattern has already at least three also logical to expect the Hpl gene frequencies exceptions: the Huasteco in northeast Mexico of the Indian tribes with evidence of Negro and the Chol in southeast Mexico have a fast admixture to be higher than in the rest. Table variant that has electrophoretic mobility similar 2 shows that this is not the case, and that in to the Naskapi type and may be the same, and fact the Cora have the lowest Hp' gene fre- the variant recently encountered in the Yano- quency in our material. 7 mama is fast but different from the Naskapi. a The recently described polymorphism of the It is also interesting to remark that whereas albumin locus seemed to have 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.8 Albumin gether with other Indian groups, it is evident, variants are rare in all groups so far studied as stated by Dr. Layrisse, that the only Ameri- except the Amerindians, and it appeared that a can aboriginal population that clearly deviates fast 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., G. Zárate, and A. Loría. Studies on several indians, and despite the fact that the precise genetic hematological traits of Mexicans. IX. Abnormal reason for this is unknown, it is important to hemoglobins and erythrocytic glucose-6-phosphate dehy- continue the characterization of the genetic con- tribes. Blood drogenase deficiency in several Indian stitution of the Amerindians, quite apart from 27:824, 1966. 7 Lisker, R., G. Zárate, and E. Rodríguez. Studies on the question whether or not it can be used at several genetic hematological traits of the Mexican popu- this point for meaningful subclassification. I lation. XIV. Serum polymorphisms in several Indian tribes. Am. I. Phys. Anthropol. 27:27, 1967. 10 Melartin, L., B. Blumberg, and R. Lisker. Albumin 8 Melartin, L. Albumin polymorphism in man. Studies Mexico: a new variant of serum albumin. Nature 115: ' on albumin variants in North American native popula- 288, 1967. tions. Acta Path. Microbiol. Scand. Suppl. 191, 1967. 11 Weitkamp, L., J. Chagnon, J. Saave, F. Salzano, 9 Melartin, L., and B. Blumberg. Albumin Naskapi, a and J. Gall. Serum albumin variants in American and new variant of serum albumin. Science 153:1664, 1966. New Guinea indigenes. Clin. Res. 16:298, 1968.

a l TABLE 2. Comparative frequencies of genes M, Fy , Hp , and chromosome cDe+cde in groups having some cases with Hb.S and /or G-6-PD deficiency with other tribes investigated for these markers

NO. CASES GENES CHROMOSOME Hp' GROUP NO. Hb.S G-6-PD M Fya cDe + cde GENE

Huasteco 235 0 0 - - - 0.51 Nahua 141 0 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 0 0.65 - 0.79 0.50 - 1.00 0.03 - 0.12 0.44 - 0.65

45 offer at least two important reasons: one, that in the first. The third stage would be that this will provide the geneticist of tomorrow with reached by tribal groups in Mexico. The genetic a good baseline for understanding population 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 differentiation of these groups population, for which at least some Indian would follow a different pattern from the other groups are quite adequate, might significantly 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 sure we need more data on different groups, General Discussion in different ecological situations. Bier: I would like to know whether the I do not think that genetic data per se would albumin variants are antigenically identical. be very important for the differentiation of Lisker: When I say there are two different Amerindians unless they were correlated with " albumin-albumin Mexico and albumin Nas- this kind of data. For instance, the selection . kapi-I am only saying that they have the same acting on groups in the second stage is probably electrophoretic mobility. 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 it is the same mutation is un- in his book Man Adapting. The nutrition of known. these groups would become somewhat more Bier: Before doing the sequence, wouldn't it limited; 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 in these three stages is quite have been found. different and that if data from populations at Salzano: I think that the studies on Vene- these different cultural stages are mixed, a 6 zuelan Indians in different stages of cultural meaningful picture is probably impossible. evolution, and also the researches of Dr. Lisker Waterlow: Dr. Neel, is it really proved that in Mexico, are very important since a pattern all these things-albumin variants, haptoglobin appeared among the Brazilian groups that can variants, red-cell variants-are genetically de- be tested. Three stages can be visualized. The termined or is that an assumption? 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 than I 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 simple population the second stage, the groups would rely more survey. Family studies are required. And in on agriculture but would not yet have a critical every instance in which a variant has been , size. These would conform more to what Sewall described today, there have been studies that Wright calls the island model; that is, their show it is transmitted according to Mendelian populations would be small and separated by laws within family lines. And this kind of large distances. A priori, more variation would evidence is necessary before we can speak of a perhaps be expected in the second groups than variant with any assurance.

46 THE AMERICAN INDIAN IN THE INTERNATIONAL BIOLOGICAL PROGRAM

James V. Neel

Administrative background four other countries may do so soon. It will be the purpose of this presentation to attempt to As is a matter of common knowledge, in 1960 a variety of types the International Council of Scientific Unions point out the manner in which of studies of the American Indian fall within (ICSU) took the initiative in the promulgation the IBP and especially the of an International Biological Program. As orig- the framework of with human adaptability. In inally envisioned by ICSU, the objectives of this section concerned provides the Americas with a * program were the world-wide study of (1) or- fact, the Indian unique resource in IBP-oriented studies and at ganic production on the land, in fresh waters, the same time presents interdigitating medical and in the seas, and the potentialities and uses and humane challenges that are worthy of the of new as well as existing natural resources; and consideration of this Organization. (2) human adaptability to changing conditions. A Planning Committee appointed by ICSU in At all the various levels of consideration of turn established a number of subcommittees to human adaptability, it has been clear there are consider various aspects of the program and to two interrelated components: the genetic, which elaborate a more extended protocol. These vari- sets the limit of the organism's ability to re- ous subcommittees combined their efforts into spond, and the physiological, which determines response available to the organism, a series of suggestions and recommendations, the range of which the Planning Committee coordinated and given its genetic set. At one extreme in the spec- submitted to ICSU in November 1963. ICSU, trum of interests that can be brought to bear on after approving these suggestions and recom- human adaptability is environmental physiology, 'mendations, then called upon the various coun- and at the other is population genetics-and in , tries having representation in it through national between there is room for many aspects of , societies to formulate national programs. The modern medicine and anthropology. timetable for IBP, several times revised, consists From the earliest considerations of most of of a "planning phase," which officially termi- the various committees and subcommittees con- nated in mid-1967, and an "operational phase," cerned with human adaptability, it has been to continue until mid-1972. apparent that the surviving primitive popula- It has been clearly understood from the outset tions afford an opportunity to study physiologi- that the various national committees were in no cal and genetic adaptations potentially quite way bound by this international formulation-it different from those of highly civilized groups. was designed merely to provide convenient Parenthetically, although our cultural orienta- guidelines. Thus far ten countries in North, tion is such that we tend to view many of these Central, and South America have established surviving primitive groups as representing ex- committees to formulate national programs, and tremes of adaptability, the fact is that they have

47 lived under their conditions far longer than we of these problems has reached such a magnitude under ours. Viewed from their vantage point, as to call for concerted action on a hitherto 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 that we immediate problems lurks a far larger issue. k will adapt to our man-made environment as Now that man so largely controls both the en- successfully as they have done to theirs. We must vironment and his reproduction, is there any note in passing that some of the adaptations of long-range objective more important than that these groups are complex and ingenious, and we understand the genetic potentialities of the that the term primitive as commonly used is individual and the species, the manner in which something of a misnomer, amounting to "pre- this potential interacts with an environment that X 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 Geneva a Scientific Group on Re- You will recognize that I have only restated search in Population Genetics of Primitive in a different context the need to understand Groups. Its purpose was to consider studies of the genetic aspects of human adaptability. The ' populations of unusual genetic interest, with par- American Indian offers prime research material . ticular reference to the surviving hunting and for this objective. The principal lines of attack food-gathering groups of the world, to advise can be conveniently summarized under three on the most suitable populations for such headings (see 12). studies, and to draw up appropriate research 1. Population structure and natural selection designs. The pertinence of this to IBP-related in 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 report selection 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). This report, inci- tion has guided human evolution and will con- dentally, was updated in the summer of 1967 tinue to guide the evolution necessary for a suc- and a revised version has just been issued as cessful genetic adjustment to a 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-related groups in primitive cultures. Otherwise stated, there is has been the rate at which these primitive groups no evidence that the developments of the past are disappearing. It can be said without exag- few thousand years have yet resulted in any very geration that ours is probably the last generation significant modifications of man's genetic po- privileged to study such groups in a relatively tential. It follows that one way of understanding T pristine state. If there are important scientific the genetic impact of our changing environment. lessons to be learned-and the WHO document is from the baseline supplied by detailed knowl- would seem to develop that case adequately- edge of primitive man in his primitive ecosys- then there is an urgency to this problem not tem. The simple fact is that at present we know shared by many other areas of research. 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 of us who are based on the results of artificial selection with concerned with human biology view our 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 to us than to the crease, trace chemicals in foodstuffs, air pollu- animal species involved. tion, water pollution, the so-called stress diseases, What the geneticist terms "population struc- cultural alienation, to mention only a few. Each ture" places limits on the effectiveness of natural

48 or artificial selection. Thus, hand in hand with cians. Although the precise nature of the data efforts to understand selective pressures must go collected depends upon local circumstances, un- efforts to understand the genetic structure of der optimal conditions we assemble as complete ykpopulations. To the geneticist, this means pat- demographic and genealogical data as possible; terns of mating, fertility, and survival; amount conduct physical, dental, and anthropological of inbreeding; migration patterns; mutation examinations; 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 another determinations. The effort to understand thropological literature, in the main what is better man's interaction with his environment aavailable simply lacks the precision and depth carries far beyond the identification of genetic necessary to the precise statistical treatments of traits-we are fortunate to have the collaboration the geneticist. of such groups as the Communicable Disease Even with all the facilities of modern science Center 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- tions-problems of logistics and communication mama, which in any event must be regarded as being what they are. Even so, from our own preliminary. Not at all surprisingly, both the experience of the past several years and that of population 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 relevant and quantitatively 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- infertility is rare, practically all women con- tion, Dr. Francisco Salzano of the University of tributing to the next generation; in which effec- Rio Grande do Sul, Brazil, and 1, with several tive fertility seems intermediate, and infant and collaborators, undertook a pilot study on the childhood mortality also intermediate; and in úXavante 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 basic surprising 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 'ritical 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 colleagues-in addi- chance may have played a greater role in much t,ion to Dr. Salzano-Drs. Miguel Layrisse and measurable human genetic variation than here- Tulio Arends of the Venezuelan Institute for tofore 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 ladder-that is, in relatively three different occasions. The composition of the densely populated agricultural societies. jeam varies from year to year, but ideally it 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 substantial portion of the mathematical theory each may be unusual in one or several respects. of population genetics is currently 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 4 complex that precise mathematical formula- factor out the significant common denominators k tions do not appear imminent, and we are being that have guided human evolution. Already forced to involved computer simulation pro- parallel studies are in progress on such groups as grams to explore the genetic consequences of the Cayapó 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 clear ton and colleagues, as well as in other parts oPf 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 I mentioned above that a substantial portion questions discussed earlier in this session by, of the mathematical theory of population ge- Professors Griffin, Cruxent, and Laughlin, the netics is being reworked. One of the most pro- date and place of arrival of the Indian in thée 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 rnan's concealed genetic variability, as subdivision of mankind. In view of what can be revealed by modern biochemical techniques. deduced concerning popuiation density in north- Many of the newly recognized variant types ern Europe 20,000 years ago, we may surmise occur in populations in such frequencies that we that the total number of immigrants across th~e speak of the underlying systems as genetic poly- Bering Bridge, whether they came in one or in, morphisms. The maintenance of such poly- 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 the of course be forever conjectural, but let us as- nature of these forces unknown, however, but sume that the tribal population(s) concerned under the previous formulations the mortality had the range 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 otherf sidered (7, 9, 25). Understanding how all this than by the Bering Bridge is negligible.) Once- variability is maintained is 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 seems self- 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 opportunity to measure ship at this 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 has just 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 three levels of miscegenation: (1) essentially 4variety of specific systems. In theory, for any none; (2) known, but probably accounting for 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 be greater than 5 per cent. The tabulation contains 4, defined by a point in a space that has one less data on the 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. Further Rh, 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 of have been computed for each study with com- Iteach population vis-a-vis the others can be plot- puter programs written in Fortran and available 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 as laborious as this of surveying populations for gene frequencies in description sounds (6, 22). But the advent of the the vague hope that someday the information 4 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 on which consider- lation distance (see 3). able progress may be expected in the near future. Although considerable progress has been made, 3. The response of genetic systems to a rapidly there is still a great deal to be done in the way changing environment. As I mentioned earlier, of characterizing the genetic attributes of the we recognize that our species is all over the relatively pure Indian groups, as a basis for 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 sized the need for continuing efforts at the a position to begin to document with any pre- \genetic characterization of all the human strains, cision what biological adjustments really take so that this type of investigation of the Indian place. We have probably lost forever the oppor- fits readily into a larger framework. The pub- tunity to study what occurred in Caucasian lished data on the Indian are widely scattered, populations. We are about to lose the opportu- no survey having been attempted since those of nity to document what happens in Negro and Mourant (10, 11) and Salzano (18). Dr. Rich- Mongoloid (including Indian) populations. 4 ard Post, Dr. William Schull, and I have now The transition of surviving primitive Negro i completed a compilation of all publications prior or Indian groups will be telescoped into a far to November 1967 that deal with the principal shorter time period than was true for Caucasian genetic polymorphisms known to occur in the groups. Thus, some Indian groups will be called Indian.1 The rate of discovery of new poly- upon to move from the Stone Age to the Atomic morphisms being what it is, any such compila- Age in a few hundred years, whereas the transition 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 reasonable to argue that here is an papers on which the tabulation is based attest opportunity to study changes in population to the strong interest in this subject. Since the structure and adjustments to new stresses that data on unmixed Indian groups have in most we must not lose. See Appendix of this volume. As for the previous two sections, so here the

51 intellectual challenge is to convert an easily Some moral issues enunciated but somewhat diffuse series of prob- It has seemed appropriate, in a presentation lems into specific 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's we increasY structure in transitional 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 Caingang of considerations, it also seems appropriate to con- Brazil (19, 20), of Layrisse and colleagues on sider briefly what these studies, 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 Quiché 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 in a 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 sev- on the Papago (5). 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 portion of this session will attest. The relations between the early settlers of his country f _ work of Roche and colleagues in particular will and the Indians without deep shame f;r chne illustrate just how rapid and subtle may be the barbarism heaped upon . people who were changes introduced by minimal acculturation. If driven to de.-.-i -lie land they occupied? The' genetic factors are important in the suscepti- world is watching my country as it agonizes bility. to the thyroid hyperplasia and diabetes over the Negro problem-it might equally well mellitus now so common in certain Indian be watching the Americas for signs of a 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 favorable material for the programs of our governments for the health, study of these genetic factors. economic advancement, and education of thep Some unpublished studies of Drs. Warren Indians? How can we translate the results of, Eveland, William Oliver, and myself may help our scientific investigations into concrete action point to some of the nuances of this transition. programs, programs that must be carefully re- Six E. coli strains were isolated by Dr. Eveland lated to other government measures? It is a from each of 77 Yanomama stools, each stool dubious favor to lower infant mortality among from a different subject. An effort is in progress the Indians without parallel economic measuresá 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 fail to (see references in 23), so easy to attribute un-J type out. By contrast, experience indicates that critically to their heritage of violence and lack only a very small fraction of E. cohi strains iso- of familiarity with our gadgets, will yield to lated in the United States would fail to type. education until the frustrations that contribute., What remains to be determined is whether this to accident-proneness are relieved. is a unique finding in a long-isolated population In a world that seems to be groping for per- or whether similar studies on other such groups spective, the Indian provides a reference point will reveal the same pattern, with the inference from which to view the fantastic disruptions of a predictable shift in the intestinal flora with that modern men, intrinsically still litdtle differ- acculturation. ent in all essential biological attributes from

52 Indians, have brought about. There are those data are grossly inadequate and we who call who will take umbrage at my characterization ourselves scientists must adhere to the null of us representatives of Western culture as "in- hypothesis, the more so since its various alterna- trinsically still . .. Indians." I am aware of the tives can be so conveniently misused by those so-called intelligence tests that purport to show who would evade their social responsibilities. the inferior intellectual qualities of the American Be this last digression as it may, each time I Indian, just as I am aware of similar results with return from the field there is a period of culture the American Negro. These results can be and shock as I realize how greatly in a short period have been used as an excuse for less-then-equal of time we have contrived to disrupt our eco- schooling. But in both instances it is a matter of system, and how profound is our ignorance of a culturally deprived and alienated group-per- the long-range results of this disruption. Now haps also subjected to early nutritional defi- in this time of greatly intensified concern over ciencies, whose role in impaired intellectual per- these problems, studies in depth of the Indian, formance we are just beginning to recognize- being judged by tests designed by and standard- within or without the framework of IBP, will ized on a very different group (see also 4). By surely contribute not only to his well-being but ,these remarks I do not mean to dismiss the also to our own perspective and, eventually, the possibility of intellectual differences between necessary adjustments toward which we are ethnic groups, but only to insist that to date the evolving.

REFERENCES

1. ARENDS, T., G. BREWER, N. CHAGNON, M. 11. MOURANT, A. E., and A. C. KOPÉC. The ABO GALLANGO, H. GERSHOWITZ, M. LAYRISSE, J. NEEL, Blood Groups: Comprehensive Tables and Maps of D. SHREFFLER, R. TASHIAN, and L. WEITKAMP. World Distribution. Springfield, Illinois, Charles C Intratribal genetic differentiation among the Yano- Thomas, 1958. 276 pp. (Royal Anthropological mama Indians of Southern Venezuela. Proc. Nat. Institute Occasional Publication 13) Acad. Sci. 57:1252-1259, 1967. 12. NEEL, J. V., and F. M. SALZANO. A prospectus 2. BEARN, A. G., B. H. BOWMAN, and F. D. for genetic studies of the American Indian. Cold KITCHIN. Genetic and biochemical considerations of Spring Harbor Symp. Quant. Biol. 27:85-98, 1964. the serum group-specific component. Cold Spring 13. NEEL, J. V., F. M. SALZANO, P. C. JUNQUEIRA, · Harbor Symp. Quant. Biol. 29:435-442, 1964. F. KEITER, and D. MAYBURY-LEWIS. Studies on the 3. CAVALLI-SFORZA, L. L., and A. W. F. EDWARDS. Xavante Indians of the Brazilian Mato Grosso. Analysis of human evolution. In S. J. Geerts (ed.), Amer. 1. Hum. Genet. 16:52-140, 1964. Genetics Today, vol. 3. Oxford, Pergamon Press, 14. NEEL, J. V., and F. M. SALZANO. Further 1965, pp. 923-932. studies on the Xavante Indians. X. Some hypoth- 4. FISCHER, J. L. Letter to the editor. Perspectives eses-generalizations resulting from these studies. 4 in Biol. & Med. 11:325-328, 1968. Amer. ]. Hum. Genet. 19:554-575, 1967. 5. HACKENBERG, R. A. An anthropological study of 15. NEEL, J. V., and N. A. CHAGNON. The demog- demographic transition. Milbank Memorial Fund raphy of two tribes of primitive, relatively un- Quart. 44:470-493, 1966. acculturated American Indians. Proc. Nat. Acad. 6. HANNA, B. L. The biological relationships Sci. 59:680-689, 1968. among Indian groups of the Southwest. Amer. 1. 16. NEEL, J. V., and W. J. SCHULL. On some Phys. Anthrop. 20:499-508, 1962. trends in understanding the genetics of man. Per- 7. KING, J. L. Continuously distributed factors spectives in Biol. & Med. 11:565-602, 1968. affecting fitness. Genetics 55:483-492, 1967. 17. ROSENBLAT, A. La población indígena de 8. LEWONTIN, R. C. Population genetics. Ann. América. Buenos Aires, Institución Española, 1945. Rev. Genetics 1:37-70, 1967. 18. SALZANO, F. M. The blood groups of South 9. MILKMAN, R. D. Heterosis as a major cause of American Indians. Amer. 1. Phys. Anthrop. 15:555- heterozygosity in nature. Genetics 55:493-495, 1967. 579, 1957. 10. MOURANT, A. E. The Distribution of the Hu- 19. SALZANO, F. M. Studies on the Caingang man Blood Groups. Springfield, Illinois, Charles C Indians. I. Demography. Human Biol. 33:110-130, Thomas, 1954, 438 pp. 1961.

53 20. SALZANO, F. M. Some genetic aspects of the Accidental deaths among British Columbia Indians. demography of American Indians. In J. Sutter (ed.), Canad. Med. Assoc. J. 94:228, 1966. Human Displacements, Entretiens de Monaco en 24. STEWARD, J. H., and L. C. FARON. Native Sciences Humaines, Premiere 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 Navaho popula- The number of balanced polymorphisms that can tion. Human Biol. 25:295-317, 1953. be maintained in a natural population. Genetics 55: 22. SANGHVI, L. D. Comparison of genetical and 469-481, 1967. morphological methods for a study of biological 26. World Health Organization. Research on difference. Amer. 1. Phys. Anthrop. 11:385-404, Population Genetics of Primitive Groups. Geneva, 1953. World Health Organization, 1964. 26 pp. (Techni-í 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 high mortality to a much greater Dr. Neel, I should like to ask you to take over extent than any innate susceptibility. again the problem raised by Dr. Waterlow a Now, how to pin down that kind 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 encounters tuberculosis or measles or pertussis, 3 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 populations are regard to those adaptive processes of which 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 susceptibility to disease, which will be able literature, based largely on studies in twins, discussed this afternoon. In other words, how on disease susceptibilities in man. There is also possible is it in practice to obtain the kind of a large literature of studies in experimental information that would determine whether the animals, 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 our gations, as with standardized challenges under discussion for this afternoon. It is relatively controlled conditions. simple to obtain evidence for the genetic nature Waterlow: What is worrying me, Dr. Neel- of the kinds of traits that Dr. Layrisse, Dr. and I am very ignorant of this subject-is that Arends, and Dr. Lisker have presented. It is the markers you are studying may in a sense be clearly much more difficult to obtain evidence 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 ' about some of the medical problems of Indian proteins, the isoenzymes such as the phospha- groups in transition. We have recently had the tases, dehydrogenases, and so on, are also experience of being in the middle of a measles genetically determined. 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 the strong impressions we came relation to adaptability. What I am not clear r away with-and Dr. Centerwall will present about is how far your program of the kind of some of the data-is that there may have been things you study is complete in itself. It rather a great deal of uncritical thinking in the past looks as if geneticists had their own list of about the constitutional susceptibility of the proteins that they call markers, whereas we Indian to this, that, and the other disease. We biochemists are thinking 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 life at high could spend a great deal of time. Probably the altitudes. most important development in human genetics Now, one of the better opportunities to get in the past decade has been the realization of at a question like that exists in Ethiopia. In the o the tremendous amount of genetically deter- highlands and lowlands of Ethiopia are groups mined biochemical diversity in our species. I, who on the basis of the genetic markers we for one, think that one of the outstanding chal- have been discussing appear to be very similar. lenges in human genetics is to understand the And yet one group lives at a very considerable significance of this diversity. What is it all there altitude and one group not far above sea level. for? It is so great that we are having difficulty There is thus an experiment of nature, as itt in seeing how it can all be maintained at once were, in which a group has been subdivided and by selective pressures. The problem is to sort out the resulting subgroups placed in diverse en- which of these polymorphisms are like vestigial vironments. organs and which are meaningful to the species. The situation does not seem to be quite so Now, to come more directly to how we get simple in the Americas. We have not yet found at this. It is quite clear that those polymorph- the "control" for Andean man. Once there are isms arose in primitive man. And we feel that such controls it is possible to investigate whether many clues to their function may come from they are physically more similar to high-altitude primitive man. One of the things we attempt to man than other Indian groups-that is, whether do in our field work is collect all our data in there has been self-selection of groups able to family relationships. We can then analyze family live at high altitudes. There is also the hope of material to see whether certain polymorphisms finding marital exchanges between the high-' are associated with increased fertility or with and low-altitude groups. decreased infant mortality. This is a long, slow Dubos: Would you elaborate on the state- business. Parallel studies on civilized populations ments that were made last year about the ana- are also needed. The numbers necessary to estab- tomical differences in the heart between man lish a significant association are tremendous. born at a high altitude and man born at our And, in fact, we almost have to think in a 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 elaborate on To sum up, although we are interested in them. As I recall it, what we were shown was these traits as "markers," for which purpose we evidence that changes appear very early on in need not understand their physiological signifi- the heart and lungs. But I did not see any cance, we are also just as interested in under- convincing evidence 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- it is a wide-open question whether this is mining whether adaptation to high altitude is a genetic or an environmental adaptation. genetically determined or a purely phenotypic Griffin: 1 have a suggestion on the general adaptation? I think this is relevant to our dis- problem Dr. Neel is working on. There was a cussion last year. somewhat comparable dispersal, by a quite dif- Moderator: There is a great deal of interest ferent 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 were a 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 see-for example, the barrel chest of the among "isolated" human groups. Andean Indian-is genetic and how much is Moderator: Yes, I agree that the Australian '

56 aborigine does represent the group most nearly and it is a very different world, the world of comparable to the Indian. However, the disrup- primitive man-you 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. It is extremely difficult now to find village before you get the same feeling for the groups of any size that have not been 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. forced amalgamation of previously separate 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. Waterlow's and Dr. tural levels. Whether, on the other hand, there Neel's remarks, I would like to ask Dr. Neel are subtle 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: Along the lines of Dr. Dubos' chemical terms, the concept of the regulation of question, Dr. Neel, are the people who are readouts at the translational level as well as at conducting the studies of lower-animal behavior the transcriptional level is beginning to emerge. also making these 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 standpoint. 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 is traits 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 as less contaminated. The primitive population mammal at the present time. However, what would be less contaminated by synthetic and might be called the comparative polymorph- non-natural factors; relative to our own situa- ology of the primates is under very active investi- tion, however, it may be more contaminated. gation. A great deal of effort is now going into "'This comes to affect the whole question of exploring 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 for any generalizations, at the level of translation. There may be all but it is a fact that some polymorphisms en- kinds of factors in terms of the phenotypic countered 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: I would remind you that the kinds of associations you refer to. But, again, I geneticist has had quite a hand in molecular come back to the problem of numbers. It is ,biology. going to be very difficult to get large numbers Obviously, there is at present no answer to on 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 us has not only touched on but developed for us who have done field work have emerged with the foremost problems of man's adapting to va feeling that here in a very different world- the new conditions he is creating. All sorts of

57 statements are continuously being made, such as 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 ". . . cannot become adapted to crowding"; a general statement on this issue. "Modern man has become adapted to pollution" Moderator: I merely want to agree that we or ". . . cannot adapt to pollution." And, in are not going to solve the problem this morning fact, these problems, which are so crucial to the and to agree with one other thing Dr. Dubos whole development or survival of our civiliza- has implied-that there are great difficulties in tion, have not as yet been formulated in suffi- clear formulations in this field. How do we set ciently clear terms for us to determine whether 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? I1 we are simply playing on the polymorphic rich- think we are in accord that this is an important ness of the human race. area. And 1 think we are in accord with respect The problem is so large and so important that to the methodology: we have a long way to go.

,y'

58 SURVEY OF THE UNACCULTURATED INDIANS OF CENTRAL AND SOUTH AMERICA'

Francisco M. Salzano

Acculturation and biological change context, is that related to the biological changes Before trying to make the survey mentioned that occur and sometimes are decisive for the in the title it is important to indicate the prob- direction and speed of acculturative modifica- lems that arise when an attempt is made to tions. define a specific group as "acculturated" or Ribeiro (20) recognizes for Brazilian Indians "unacculturated." The term acculturation is three stages on the road to acculturation. The generally employed by American anthropolo- first would be represented by the isolated tribes gists, while their British colleagues prefer the -those living in regions not yet occupied by the word culture contact and the Cuban scholar Brazilian society and having only accidental and F. Ortiz the term transculturation, which em- rare contacts with our civilization. Some simply phasizes the reciprocal character of most contact avoid such contacts, while those living near situations. Though much criticized and modi- pioneer settlements may have suffered violence fied, the most widely used definition is that of and be frankly hostile to "foreign" people. In Redfield, Lindon, and Herskovits (19): "Accul- either case, they can obtain all they need from we< turation comprehends those phenomena which the region where they live and from their work; result when groups of individuals having dif- they are therefore able to maintain complete ferent cultures come into continuous first-hand cultural autonomy. The second stage would be contact, with subsequent changes in the original that of populations having intermittent contacts cultural patterns of either or both groups." with civilization. They live in regions already A number of basic problems exist in the inter- reached by the expanding frontiers of the na- pretation and analysis of these situations; they tional society; though they maintain a certain are reviewed by, among others, Beals (1) and cultural autonomy, they already have needs that Oliveira (16). For instance, we have sometimes can only be satisfied through economic relations conflicting views of culturalists and sociologists, with civilized people. The third stage is repre- and in some specific cases the intrinsic nature of sented by groups in permanent contact with the process and its consequences are difficult to more numerous and more differentiated members of the civilization. They have lost their '> identify. A new dimension, which so far as I am aware has not been properly analyzed in this cultural autonomy to a large degree, being entirely dependent on others for the supply of 1 Research supported in part by the Rockefeller Foun- many tools and products. They still preserve dation, Conselho Nacional de Pesquisas, Conselho de some of their traditional customs, but already Pesquisas da Universidade Federal do Rio Grande do show profound changes due to ecologic, eco- Sul, U.S. Public Health Service, and Wenner-Gren Foundation for Anthropological Research. nomic, and cultural pressures.

59 What concomitant modifications occur in the sonal communications of many colleagues. The biology of these groups? Only a very limited rapidity of change in these populations and also start has been made on quantifying the observa- the discovery of new groups indicate that such tions 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 sur- 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 many Even 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 Cayapó group of central lations were checked in the most 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 incidence of tuberculin reactors. reviews concerning the older ones, see Rosenblat Since the Amerindian populations apparently (21) and Steward (23). had no contact with tubercle bacilli in pre- Columbian times, the question arises, From The survey what groups did they get the infection? Their CENTRAL AMERICA .- contacts with neo-Brazilians are recent and intermittent, and it is reasonable to suppose that Political subdivisions: Bahamas, Belize, Bermuda, Costa Rica, Cuba, Dominican Republic, El Salvador, the source of infection was the Gorotire, another Guadeloupe (and dependencies), Guatemala, Haiti, Hon- Cayapó group from whom they separated at the duras, Martinique, Mexico, Netherlands Antilles, Nica- beginning of this century but with whom they ragua, Panama, Puerto Rico, Virgin Islands, West 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, of the diffusion of cultural traits. The speed Nicaragua, Panama. these epidemiological changes is such that it is Indian population: 8,000,000 (10 per cent of the 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 (1.3 per 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- Information about the Indians: Mainly Maya antl date important problems of human biology. Chekchi, who came from Mexico and Guatemala. Unacculturated Indians: Probably none. 9. Source of the data and their limitation Rejerence:

In the next section I shall 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. Inlormation about the Indians: Most important groups first-hand contact only with Brazilian Indians, appear to be Bribrí (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 Indlians belong to groups

60 of Chorotegamangue (Nicoya Peninsula) and Guatuso NICARAGUA borders). (northern and western Total population: 1,631,000 (1966). UnacculturatedIndians: Probably none. Indian population: 80,000. 9, 24. References: Information about the Indians: Most important groups Negro mixture), EL SALVADOR are Miskito (with a large component of Sumu, Rama, and Ulva. Total population: 3,000,000. Unacculturated Indians: Stone (24) mentions the Ulva Indian population: 100,000. tribe between the Mico and Siquia rivers, and the Sumu Information about the Indians: Most important group (3,000-4,000 in 1948). is Pipil (about 80,000) of Nahuatl origin, concentrated Rejerences: 9, 18, 22, 24. in various towns in Sonsonate Department. Also smaller groups of Nahua and Cakchiquel distributed over several PANAMA departments. Total population: 1,314,460. Unacculturated Indians: Probably none. Indian population: 65,000-100,000. References: 9, 24. Information about the Indians: Main tribes are Guaymi

GUATEMALA (about 36,000), Cuna (some 22,000), and Chocó (6,000). Two other small groups are Bocotá (200) and Teribe Total population: 4,438,000 (1965). (400). Guaymi are found in Chiriquí, Bocas del Toro, Indian population: 2,500,000. and Veraguas provinces; Cuna mainly in San Blas Prov- Information about the Indians: Main groups in center ince, both mainland and some islands; Chocó in Darién and west: Maya, Cakchiquel, Tzutujil, Uspanteca, Achí. Province. West: Mam, Aguacateca, Jacaltec, Kanjobal, Chuj, Ixil. Unaccultulrated Indians: Some Chocó groups of Pan- North: Kechkí, Pocomchí, Pocomam, Lacandón, Mopán. amanian-Colombian border (?). East: Chortí, Lacandón Chol, Chontal. Southeast: Xinca. References: 9, 24, 26. Unacculturated Indians: Some groups living in inaccessible places could provide information on the influ- SOUTH AMERICA ence of a non-European culture on genetic and other subdivision: Argentina, Bolivia, Brazil, Chile, traits. Political References: 9, 18. Colombia, Ecuador, French Guiana, Guyana, Paraguay, Peru, Surinam, Uruguay, Venezuela. HONDURAS Total population: 175 million. Countries with significant number of Indians: All Total population: 2,400,000. except Uruguay. Indian population: 110,000. Indian population: 10 million (6 per cent of the area's Information about the Indians: Main groups in west: total population). Chorotega, Miquirano, Guajiro. On coast: Moreno. Other: Unacculturated Indians: Perhaps 980,000 (0.6 per cent Opatoro, Jicaque, Paya, Zambo, Miskito, Sumu. The of the total population). Miskito have mixed with Negro groups. UnacculturatedIndians: Probably none. ARGENTINA References: 9, 18, 24. Total population: 22,254,102 (1965). MEXICO Indian population: 130,000. Information about the Indians: Main groups in Chaco Total population: 40,913,000 (1965). region: Mataco, Chulupi, Chorote, Chiriguano, Pilagá Indian population: Estimates vary widely. A conserva- (totaling about 30,000). Andean region: Quechua and tive number would be 5,000,000. Colla (some 18,000). South: Araucanian. Other regions: Information about the Indians: Main gloups in north: Mocobi, Toba. Kikapú, Tepehuano, Nahuatl, Pame, Huastec, Otomí. Cainguá, Unacculturated Indians: Perhaps some Chaco groups. Northern Pacific: Cora, Huichol, Yaqui, Seri, Papago. References: 8, 9. Center: Mazahua, Matlatzinca, Tarasco, Mazatec, Mixtec, Popoloco, Totonac, Zoque, some of previous groups. BOLIVIA Southern Pacific: Chol, Tzeltal, Tzotzil, Tojolabal, Maya, Lacandón, Amuzgo, Tlapanec, Chatifio, Chinantec, Cho- Total popurlation: 3,800,000. cho, Chontal, Huave, Mixe, Triche, Zapotec. Indian population: 2,450,000. Unacculturated Indians: Lacandón (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 Rejerences: 4, 9, 18, 25. tribes belonging to 15 linguistic families. Some of them

61 are as follows: Beni Department: Sirionó, Baure, Jora, COLOMBIA Chacobo, Tacana, Chimane, Movima, Yari, Sirineire, Total population: 17,787,000. Cayubaba, Itoma; Department of Santa Cruz: Guarayo, Indian population: 300,000. Borosese, Chamacoco, Zamuco, Lengua, Tapieti, Otuqui, Information about the Indians: Main groups in Cauca .S Guatose, Yuracaré; Pando Department: Araona, Toro- region: Guambiano, Páez, Guanaco, Yanacona, Coco- noma, Pacaguara, Machicanga; Department of La Paz: nuco (approximately 80,000). Nariño: Quillacinga, Chama, Chimane, Guarayo, Leco. Southeastern region Kwaiker, Inga, Kamsá (about 60,000). Guajira: Guajiro (Bolivian Chaco): Tarija Department: Chiriguano, (50,000). Magdalena: Kogi, Ika, Chimila, Yuko (15,000). Mataco, Toba, Chulupi, Guaycuru, Chanese (about Antioquia: Cuna (2,000). 14,500); Quiguisaca Department: Chorote, Chiriguano Unacculturated Indians: Perhaps 100,000. (some 3,800); Cochabamba Department: Sirionó, Yura- References: 2, 9. caré (2,000). General total for the region: approximately

50,000 Indians. ECUADOR Unacculturated Indians: Almost all forest-dwellers. References: 9, 28. Total population: 5,084,000. Indian population: 2,000,000. Information about the Indians: Main groups in coastal

BRAZIL region and east: Colorado, Jívaro (13,000), Yumbo, Auca. Sierra: Quechua groups: Quindivana, Saguatoa, 41 Total population: 90,193,000. Salasate, Poato, Ilumane, Peguche, Quinchuqui, Quilotoa, Indian population: 100,000. Zumbagua, Pilaloe, Siquisilie, Puruhay, Cañaris, Sara- noiiji ,iNato,aboui ile idians: Tribes or groups ciassi- guro. fied by Ribeiro (19) and others as isolated or in inter- Unacculturated Indians: Perhaps 250,000 (50,000 in mittent contact (excluding those with fewer than 50 the jungle). individuals): Agavotokueng, Apalai (100-250), Araras, References: 9, 15, 17, 28. Asurini (250-500), Atruahí, Barawana, Boca Negra,

Canoeiros, Cayabi (250-500), Diore (500-1,000), FRENCH GUIANA Gavioes (1,500-2,000), Guajá (100-250), Ipewi, Javai Total population: 36,000. (250-500), Kabixí, Kalapalo (100-250), Kamayurá Indian population: 1,200. (100-250), Katawian, Kren-Akoro (2,500-3,000), Information about the Indians: In coastal region: Kuben-Kran-Kegn (250-500), Kuben-Kragnotire (400), Galibi' (600), Arawak (100), Palikour (100). In interior: Kuikuro (100-250), Makú, Mandawáka, Maopityan, Wayapi (130), Mereyo (60), Waiyana (200). Makirutare or Mayongong (500-1,000), Mehináku Unacculturated Indians: About 900 (Galibi, Palikour, (50-100), Mentuktire (500-1,000), Mudjetire (50- Waiyana). 100), Nahukuá (50-100), Nambicuara (500-1,000), 1, Rejerences: 3. Pakaánovas (500-1,000), Parakanan (250-500), Pari- kotó, Pianokotó, (250-500), Puruborá (50-100), GUYANA Salumá, Sikiana, Tapayúna, Tirió (2,000-3,000), Trumai, Txicáo, Txukahamae (200), Urubus-Kaapor Total population: 647,000. (500-1,000), Waimirí, Waiwai (100-250), Waurá (100- Indian population: 27,840. 250), Xavante (2,000-3,000), Xikrin (250-500), Ya- Information about the Indians: Main groups in coastal baána, Yanomama (2,000). For location, see references region: Arawak, Carib, Warrau, Akawaio (about 10,000). 10 and 19. The former presents many other names, but In interior: Carib, Akawaio, Patamona, Arekuna, Makusi, it is not always clear whether tribes or tribal subdivisions Wapisiana, Waiwai. are meant. Unacculturated Indians: Perhaps 2,000. Unacculturated Indians: Perhaps about 40,000. Re/erences: 3, 23 (p. 158). ReJerences: 11, 20.

PARAGUAY

CHILE Total population: 2,030,000. Indian population: 37,570. Total population: 8,567,000 (1965). Information about the Indians: They speak languages Indian population: 130,000. of 6 linguistic families and live in 17 locations. Main Injormation about the Indians: Main groups in North- groups: Chiripá, Mbia, Tabitera, Guayaki, Guarayú, ern region: Quechua and Aymara. Southern region: Chiriguano, Tapieti, Maká (about 600), Chulupi (more Araucanian (Mapuches). than 10,000), Chorote (approximately 100), Lengua Unacculturated Indians: Probably none. (5,000), Sanapana (3,400), Guana (400), Angaite (400), Rejerences: 9. Toba (1,400), Chamacoco, Morotoco.

62 Unacculturated Indians: Perhaps 10,000. has to do with the identification of the accultura- References: 9; data supplied by Department of Indian tion process per se; the other is the proper place- Affairs. ment of the group studied into "acculturated"

PERU or "unacculturated" categories. Because of the lack of detailed studies in most of the tribes Total population: 11,800,000. Indian population: 4,800,000. surveyed here, and of reviews placing them in InJormation about the Indians: Main groups in North- the appropriate category, the numbers obtained west: Aguaruna, Jívaro, 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, Chama, Huitoto, With these reservations in mind it is now pos- Ticuna, Cocoma, Capanahua, Cashibo, Shipibo. Center: Campa, Amuesha, Amahuaca, Marinahua, Cashinaua. sible to examine the numbers arrived at. The Southeast: Sirinayri, Machigüenga, Piro, Mashco, Hua- population of Central America was estimated at rayo. Carneiro (5) also mentions following tribes from 77 million and the number of unacculturated montaña: Yaminahua, Remo, Marobo and Iñapari. Indians at about one million (1.3 per cent). In Unacculturated Indians: Perhaps 450,000. South America the proportion is even lower: Reierences: 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 Total population: 360,000. we may still have about two million largely 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), Arawak (1,500). In interior: tive data in a number of populations living in Waiyana (200), Trio or Tirió (400). UnacculturatedIndians: Perhaps 2,000. different ecosystems. As a matter of fact, we Reference: 3. simply do not have experienced persons to un- dertake the study of even, say, 10 per cent of this VENEZUELA number in the immediate future. Total population: 8,880,000. With acculturation changes occurring so Indian population: 62,700. quickly, we need to work fast to establish on a Information about the Indians: Layrisse and Wilbert firm basis the biological and nonbiological forces (9) list the following: Guajiro (8,429), Paraujano

"r (1,348), Yupa (2,057), Barí (1,000), Yaruro (1,427), acting at this cultural level and determining the Guahibo (5,397), Piaroa (1,886), Panare (412), Yaba- fate of these populations. This information is rana (64), Makiritare (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 Sapé (100), Pemon (2,700), Warao (11,700), Carifa (2,776), unknown (1,000). problems. The acculturation process is generally UnacculturatedIndians: Perhaps 30,000. painful and distressing. In fact, a large propor- References: 6, 7, 10, 29, 30. tion of the groups affected by it simply disappear physically. Data on populations living at dif- Discussion ferent cultural levels can provide clues to an As was mentioned previously 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. Kroeber 3. BUTT, A. J. The Guianas. Bull. Intern. Comm. (ed.), Anthropology Today. Chicago, University of Urgent Anthrop. Ethnol. Res. 7:69-90, 1965. Chicago Press, 1953, pp. 621-641. 4. CAMARA, F. Rescate etnográfico nacional en 2. BURGos GUEVARA, H., and G. PESÁNTEZ REINOSO. México. Bull. Intern. Comm. Urgent Anthrop. Plan nacional indigenista de Colombia: 1966-1969. Ethnol. Res. 7:99-104, 1965. América Indígena 27:751-781, 1967. 5. CARNEIRO, R. L. Little-known tribes of the

63 Peruvian Montaña. Bull. Intern. Comm. Urgent aniversario, 1. México, D. F., Libros de México, Anthrop. Ethnol. Res. 5:80-85, 1962. 1965, pp. 79-134. 6. COMISIÓN INDIGENISTA NACIONAL (VENEZUELA). 18. PERICOT Y GARCíA, L. América Indígena. Tomo Nuestro Indio. Caracas, Ministerio de Justicia, 1960. 1. El hombre americano. Los pueblos de América. 26 pp. 2nd ed. Barcelona, Salvat. 1962. 1182 pp. 7. COMISIÓN INDIGENISTA NACIONAL. Informe de 19. REDFIELD, R., R. LINTON, and M. J. HERSKO- la Comisión Indigenista Nacional del Ministerio de VITS. Memorandum on the study of acculturation. Justicia de Venezuela, correspondiente al año de Amer. Anthrop. 38:149-152, 1936. 1964. América Indígena 25:131-145, 1965. 20. RIBEIRO, D. Culturas e línguas indígenas do 8. FOCK, N. Urgent ethnographical tasks in the Brasil. Educajfo e Ciencias Sociais (Rio de Janeiro) Argentine Chaco. Bull. Intern. Comm. Urgent 2:5-102, 1957. Anthrop. Ethnol. Res. 5:162-166, 1962. 21. ROSENBLAT, A. La población indígena de 9. Guía de la población indígena de América. América. Buenos Aires, Institución Cultural Es- Boletin Indigenista 21:170-266, 1961. pañola, 1945. 296 pp. 10. LAYRISSE, M., and J. WILBERT. Indian societies 22. STEWARD, J. H. (ed.) Handbook of South of Venezuela. Their blood group types. Caracas, American Indians. Bull. Bur. Amer. Ethnol. 143 (5 Instituto Caribe de Antropología y Sociología, volumes, 1946-1950). Fundación La Salle de Ciencias Naturales, 1966. 23. STEWARD, J. H. The native population of 318 pp. (Monograph 13) South America. Handbook of South American 11. MALCHER, J. M. G. indios. Grau de integrafao Indians 5:655-668, 1949. na comunidade nacional Grupo linguistiro. Locali- 24. STONE, D. Urgent tasks of research concerning zafao. Rio de Janeiro, Conselho Nacional de the cultures and languages of Central American Proteçao aos Indios, 1964. 266 pp. (Nova série, Indian tribes. Bull. Intern. Comm. Urgent Anthrop. Publicaçco 1) Ethnol. Res. 5:65-69, 1962. 12. NEEL, J. V., and F. M. SALZANO. A prospectus 25. SWANTON, J. R. The Indian tribes of North for genetic studies on the American Indian. In P. T. >A America. Bull. Bur. Amer. Ethnol. 145:1-726, 1952. Baker and J. S. Weiner (eds.), The Biology of 26. TORRES DE ARAÚZ, R. Los grupos humanos de Human Adaptability. Oxford, Clarendon Press, 1966, Panamá. In Homenaje a luan Comas en su 65 pp. 245-274. aniversario, 1. México, D. F., Libros de México, 13. NEEL, J. V., and F. M. SALZANO. Further 1965, pp. 135-142. studies on the Xavante Indians. X. Some hypotheses- 27. VILLA RojAs, A. Departamento de Investiga- generalizations resulting from these studies. Amer. ciones Antropológicas del Instituto 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. Anuario Indigenista 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. OBEREM, U. 1962. Dringende ethnologische partamento de Investigaciones Antropológicas, 1965- Forschungsaufgaben 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. O índio e o mundo dos in Venezuela. Bull. Intern. Comm. Urgent Anthrop. brancos. Sao Paulo, Difusao Européia do Livro, Ethnol. Res. 5:86-96, 1962. 1964. 144 pp. 30. ZERRIES, O. Die volkerkundliche Forschungs- 17. ORBE, G. R. Destribalización de indígenas en situation in Suidost-Venezuela. Bull. Intern. Comm. Ecuador. In Homenaje a Juan Comas en su 65 Urgent Anthrop. Ethnol. Res. 5:97-111, 1962.

64 Appendix ALPHABETICAL LIST OF TRIBES CITED 1

Achí (Gua) Chama (Bo, Pe) Huambisa (Pe) Agavotokueng (Br) Chamacoco (Bo, Par) Huarayo (Pe) Aguacateca (Gua) Chanese (Bo) Huastec (M) Aguaruna (Pe) Chatifio (M) Huave (M) Akawaio (Gui) Chayhuita (Pe) Huichol (M) Amahuaca (Pe) Chekchi (Be) Huitoto (Pe) Amuesha (Pe) Chimane (Bo) Amuzgo (M) Chimila (Co) Ika (Co) Andoa (Pe) Chinantec (M) Ilumane (E) Angaite (Par) Chiriguano (A, Bo, Par) Ifiapari (Pe) Apalai (Br) Chiripá (Par) Inga (Co) Araona (Bo) Chocho (M) Ipewi (Br) Araras (Br) Chocó (Pan) Itoma (Bo) Araucanian (A) Chol (M) Ixil (Gua) Araucanian (Mapuche) (Ch) Chontal (Gua, M) Arawak (Fr G, Guy, S) Chorote (A, Bo, Par) Jacaltec (Gua) Arekuna (Gui) Chorotega (H) Javai (Br) Asurini (Br) Chorotegamangue (CR) Jebero (Pe) Atruahí (Br) Chortí (Gua) Jicaque (H) Auca (E) Chulupi (A, Bo, Par) Jívaro (E, Pe) Aymara (Bo, Ch) Chuj (Gua) Jora (Bo) Cocoma (Pe) Coconuco (Co) Barawana (Br) Kabixí (Br) Colla (A) Bare (V) Kalapalo (Br) Colorado (E) Barí (V) Kamayurá (Br) Cora (M) Baure (Bo) Kamsá (Co) Cuna (Pan, Co) Boca Negra (Br) Kanjobal (Gua) Curaraye (Pe) Bocotá (Pan) Katawian (Br) Curipaco (V) Bora (Pe) Kechkí (Gua) Kikapús (M) Borosese (Bo) Diore (Br) Boruca (CR) Kogi (Co) Kren-Akoro (Br) Bribrí (CR) Galibi (Fr G, S) Kuben-Kragnotire (Br) Gavioes (Br) Kuben-Kran-Kegn (Br) Cainguá (A) Guahibo (V) Kuikuro (Br) Cakchiquel (ES, Gua) Guajá (Br) Kwaiker (Co) Campa (Pe) Guajiro (H, Co, V) Cañiaris (E) Guambiano (Co) Canoeiros (Br) Guana (Par) Lacandón (Gua, M) Capanahua (Pe) Guanaco (Co) Lacandón Chol (Gua) Carib (Guy) Guarayo (Bo) Leco (Bo) Carifia (V) Guarayú (Par) Lengua (Bo, Par) Cashibo (Pe) Guatose (Bo) Cashinaua (Pe) Guatuso (CR) Machicanga (Bo) Cayabi (Br) Guayaki (Par) Machiguenga (Pe) Cayubaba (Bo) Guaycuru (Bo) Maká (Par) (V, Br) Chacobo (Bo) Guaymi (Pan) Makiritare

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

65 Makú (Br) Patamona (Guy) Tirió (Br) Makusi (Guy) Paya (H) Tlaponeco (M) Mam (Gua) Peguche (E) Toba (A, Bo, Par) Mandawáka (Br) Pemon (V) Tojolabal (M) Maopityan (Br) Pianokotó (Br) Toronoma (Bo) Marinahua (Pe) Piapoco (V) Totonaco (M) Marobo (Pe) Piaroa (V) Triche (M) Mashco (Pe) Pilagá (A) Trio (or Tirió) (S) Mataco (A, Bo) Pilapoe (E) Trumái (Br) Matlatzinca (M) Pipil (ES) Txic5o (Br) Maya (Be, M, Gua) Piro (Pe) Txukahamae (Br) Mayongong (Br) Poato (E) Tzeltal (M) Mayoruna (Pe) Pocomam (Gua) Tzotzil (M) Mazahua (M) Pocomchí (Gua) Tzutujil (Gua) Mazatec (M) Popoloco (M) Mbia (Par) Puinave (V) Ulva (N) Mehináku (Br) Puruborá (Br) Urubus-Kaapor (Br) Mentuktire (Br) Puruhay (E) Uspanteca (Gua) Mereyo (Fr G) Miquirano (H) Quechua (A, Bo, Ch, E) Waimirí (Br) Miskito (H, N) Quillacinga (Co) Waiwai (Br, Guy) Mixé (M) Quilotoa (E) Waiyana (Fr G, S) Mixtec (M) Quinchuqui (E) Wapisiana (Guy) Mocobi (A) Quindivana (E) Warao (V) Mopán (Gua) Warrau (Guy) Moreno (H) Rama (N) Waurá (Br) Morotoco (Par) Remo (Pe) Wayapi (Fr G) Movima (Bo) Mudjetire (Br) Saguatoa (E) Xavante Salasate (E) (Br) Xikrin Nahua (ES) Salumá (Br) (Br) Nahuakuá (Br) Sanapana (Par) Xinca (Gua) Nahuatl (ES, M) Sapé (V) Nambicuara (Br) Saraguro (E) Yabaána (Br) Seri (M) Yabarana (V) Ocayna (Pe) Shapra (Pe) Yagua (Pe) Opatoro (H) Shipibo (Pe) Yaminahua (Pe) Orejon (Pe) Sikifna (Br) Yanacona (Co) Otomi (M) Siquisilie (E) Yanomamo (Br, V) Otuqui (Bo) Sirinayri (Pe) Yaqui (M) Sirineire (Bo) Yari (Bo) Pacaguara (Bo) Sirionó (Bo) Yaruro (V) Páez (Co) Sumu (H, N) Yuko (Co) Pakaánovas (Br) Yumbo (E) Palikour (Fr G) Tabitera (Par) Yupa (V) Pame (M) Tacana (Bo) Yuracaré (Bo) Panare (V) Tapayúna (Br) Pano (Pe) Tapieti (Bo, Par) Zambo (H) Papago (M) Tarasco (M) Zamuco (Bo) Parakanan (Br) Tepehuano (M) Zapotec (M) Paraujano (V) Teribe (Pan) Zoque (M) Parikotó (Br) Ticuna (Pe) Zumbagua (E)

66 DISCUSSION

Moderator: Dr. Salzano, I wonder 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 your response 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 find those numbers. Where any tuberculin reaction. So I do not think that are these 1,000,000 concentrated? in this particular instance cross-reactivity is a Salzano: As I said, these are very preliminary problem. estimates. They make allowance for the very Cohen: I was using that only as an example. large groups in Guatamala and Mexico on Unless one knows what the spectrum of sensi- which we do not know enough for me to set a tivity of a population is to a test that has been specific figure. But I am sure that there should developed for a different population, one has 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, and 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 in the group of central Brazil? Are you sure the Indians with positive tuberculin tests, and whether or .·> do not have histoplasmosis or some other condi- not there was positive sputum? Second, in the tion that would cross-react and you would group from which it was thought that the appear to be getting a tuberculin that may be infection might have come, what was known unrelated to the exposure to the disease? about the amount of tuberculosis caused by Salzano: 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 in relation to the nonspecific Moderator: That is just about to be discussed reaction but also problems of reading and the by Dr. Nutels.

67 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 one ure 1). This region in the geographical heart rapid contact-for 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 Kamaiurá, this paper is simply to mention facts and report Kalapalo, Kuikuro, Waurá, Iawalapiti, Aueti, on my personal experience, not to interpret them. Trumai, Juruna, Aupatse, Nahukwá, Matipuhy, i In considering the deficiencies of such a study Kajabí (since 1952), Suiá (since 1960), Txu- it is necessary to take into account the difficulties karramai (since 1962), and Tchic5o tribes (5). that prevail in this enormous and primitive Table 1 shows the age distribution in the groups that have been examined. region, lacking in means of transportation and These fifteen tribes are autonomous. Some communication, where a dispersed and almost groups now dispersed are to be found among extinct population still live as in the Stone Age. them. There are others, not yet contacted, living According to various authors cited by Wagley in or around the Park. The tribes belong mainly (20), the native population of Brazil at the time to four linguistic groups: 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, is reduced 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 this decline, infectious diseases-at 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 men-have perhaps mission is needed by all strangers wishing to been the most efficient (13). Many are the enter. Communications within the Park are documented instances of extermination of the limited by the fact that it has only one airplane, indigenous population 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 B6as, 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 1. 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

0; Kamaiurá 89 16 11 11 9 18 24 Kajabí 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 Suiá 67 11 7 11 6 18 14 Iawalapiti 34 2 8 3 3 6 12 ,1A Trumai 22 5 3 2 3 4 5 Aucti 40 9 8 6 3 14 Kuikuro 118 20 19 10 8 38 23 Waurá 62 17 14 1 7 15 8 Txukarramai 170 36 21 19 16 33 45 Tchicáo 53 9 4 5 9 22 4 Total 959 186 144 94 99 221 215

69 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 seems rather peculiar that I found no sign of it, not even Karl von den Steinen expedition when the first congenital, and that it 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 tuberculosis are totally non- 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 culosis is of the greatest significance, since wherever Indians come into direct contact with white men, terrible Lieutenant Ramiro Noronha, in 1920; Percy devastation results. Fawcett, also in 1920; Heinrich Hintermann, It may also be that measles, scarlet fever, and small- in 1924; the Rondon Commission, this time di- pox are likewise unknown in the Xingu area, although rected by Captain Vicente Vasconcelos, also in we are not so sure about their nonexistence at the time 1924; Percy Fawcett again, in 1925 (his sec- of our visit as we are in the case of the previously Only in the case of smallpox can ond expedition, during which he disappeared); mentioned diseases. we assure its nonexistence, because we found no one G. M. Doyott, in 1928; Vincent M. Petrullo, in displaying smallpox scars. 1931; and the National Service for the Protection In 1946 the white man took firm hold of the of Indians, in 1940 (1). region, though not in large numbers. Before In 1898 Karl E. Ranke, a German physician entering the area that later became the Xingu and demographer who had participated in the National Park, they were all given a medical Meyer expedition, reported on the diseases he examination. Between 1947 and 1950, in my had observed (12). His account is still relevant, capacity as a doctor in the Roncador-Xingu and I feel it is worth quoting in full: expedition, I imposed a sort ot moral and sani- The diseases observed in 10 Indian villages among tary quarantine on every person entering the approximately 800 to 1000 Indians are the following: able to prevent venereal many healed fractures and one old haunch luxation but, region. Thus we were according to the anamnesis, not congenital; one case of diseases and to protect the Indian women from clubfoot, congenital; a very frequent skin disease, de- the "civilized" men who would have competed scribed as "Tinea imbricata," from the Malayan Archi- with the native males. pelago and the Southern sea; numerous furuncles, most At the time of their first contact with the Ron- of them in the gluteal region; two cases of idiocy; one cador-Xingu expedition of 1946, about 25 Kala- case of apparently parasitical tumor in the liver, which 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 also in the Kamayurá tribe, years old; and a fairly mild enteritis among newborn killing 12 persons this time. In June 1954 there infants. The occurrence of leucomas and staphylomas was an epidemic of measles in the upper Xingu was extremely frequent in the Xingu area. Among the Bakairi from Kulisehu, none had been spared. (7). Neither from the recollections of the elders With the help of a Bakairi from Paranatinga, the nor from the traditions 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 area, I learned the history of Indian who came in contact with the Roncador- their existence. Once, after the second expedition to the Xingu area, 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. welcomed, 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, others 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 endemic malaria (Atnopheles darlingi, Plasmo-

70 dium vivax and P. falciparum); Tinea imbri- Pirquet modified: cutipuncture) and since 1960 cata; gastroenteritis; a purulent lung abscess of with purified tuberculin (PPD-Rt 23-1 UT), I unknown etiology; furunculosis; childhood um- have investigated tuberculosis not only in the bilical hernia; arthritis deformans; helminthia- Park but also in other Indian villages, 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 Kajabí Indians came to the Park sis alba; gravidic striation; Pulex penetrans; from the Teles Pires River, in the valley of the conjunctivitis; pigmentary and cellular nevi (6); Tapajós River, bringing with them Blastomy- leucoma; pinguecula; pterygium; pupillary seclu- cosis cheloidiana (Jorge L6bo'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 original area. malarial hematological disturbances (17); de- In 1960 the Villas Bóas brothers established formation of the auricular pinna due to a fight- contact with the Suiá tribe. This small group ing game called huca-huca; atrophy of the lower of approximately 80 Indians had had only limbs due to the ritual custom of prolonged con- sporadic encounters, over a three-day period, striction in infancy; scapulo-humeral luxation; with the first Von den Steinen expedition in tegumentary leishmaniasis (4); reversible pa- 1884. At that time Von den Steinen estimated ralysis of the lower limbs due to ritual ingestion their number at 150. No other encounter was of mucuna (Dolichos pruriens and/or Dolichos recorded, even by Von den Steinen himself on urens); toxoplasmosis, recently discovered (2); his second expedition in 1887 (19). The pres- filariasis (2); and various arboviruses, recently ence of the Suiá group on the upper Paranajuba discovered (2). River was known because of their periodic 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 my greaty feared. A few days after the Villas B6as personal experience in dealing with groups of contact 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 main field of my culin survey (PPD-Rt 23-1 UT). Though at observations. Beginning in 1952, at first with that time we did not have much suitable equip- X-ray and sometimes with old tuberculin (Von ment and had had little experience with PPD,

TABLE 2. Reactions to tuberculin test (PPD-Rt23-1 UT) in Indian tribes of Xingu National Park, Brazil, July 1960

TESTS READ

0-4 MM 5-9 MM 10 MM + NO.

TRIBE TESTED lTOTAL NO. % NO. %0 NO. %

Kamaiurá 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 Suiá 42 42 34 81.0 1 2.4 7 16.6 Iawalapiti 37 37 37 100.0 Trumai 17 17 17 100.0 Aucti 15 15 15 100.0 Kuikuro 3 3 3 100.0 Waurá 2 2 1 50.0 1 50.0 Total 389 389 373 95.9 5 1.3 11 2.8

71 we found eight positive reactions-one weak TABLE 4. Reactions to tuberculin test (PPD-Rt23-1 UT) and seven strong-among this population, for by age group among Suiá Indians of Xingu National Park, Brazil, August 1966 an infection rate of 19 per cent. We also, on

that occasion, gave the PPD test-again for the TESTS READ first time-to the other tribes of the upper M 0-4 5-9 10 Xingu, finding infection rates of 3.2 per cent MM MM MIM + among the Kajabí and 5.3 per cent among the AGE NO. Meinako. The former, as I have said, had had GROUP TESTED NO. % NO. % NO. % 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 Sao Paulo (8). 10-14 13 11 84.6 - - 2 15.4 The other tribes were found to be nonallergic. 15-19 5 4 80.0 - - 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 Suiá, 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 Sula solution to the Cen- vealed BK in their sputa. Moreover, we dis- tral Tuberculosis Laboratory in the State of covered other strongiy positive cases presenting Guanabara, where they are being studied by radiologic forms of the type that do not usually Professor Milton Fontes Magarao and his staff reveal BK in direct sputum analysis, such as, (Tables 3, 4, and 5). for example, two ganglionary forms. All were treated with INH, PAS, and In 1965, 180 Indians of the Txukarramái SM (Table 8). In 1967 there came to the knowledge of the tribe, fleeing the pressure of so-called pioneer Park administrators the fact that a tribe of com- fronts (in this case a group of garimpeiros, pletely isolated Indians-the Tchikao-were be- prospectors of precious stones and metal), sought ing pressed by another 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 the left bank of the few contacts they are known to have had con- Xingu River. When observed in 1962, this tribe y 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 Txukarramai 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 Suiá Indians of Xingu National their sputa (Table 7) (10). Three persons re- Park, Brazil, July 1967

TESTS READ TABLE 3. Reactions to tuberculin test (PPD-Rt23-1 UT) among Suiá Indians of Xingu National Park, Brazil, 0-4 5-9 10 June 1960 IMM MM MM + AGE NO.

GROUP TESTED NO. o0 NO. % NO. %0 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. % NO. % NO. % 15-19 6 4 66.7 - - 2 33.3 20-29 18 14 77.8 - - 4 22.2 Adults 21 17 80.9 - - 4 19.1 30 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 TABLE 6. Reactions to tuberculin test (PPD-Rt23-1 UT) TABLE 8. Findings of tubreculin tests, X-rays, and among Txukarramai Indians of Xingu National Park, bacilloscopy in Txukarramái tribe of Xingu National Brazil, 1962 Park, Brazil, August 1966

TESTS READ TUBERCULIN TESTS

0-4 5-9 10 PULMONARY 0-4 5-9 10 MM MM MM + SHADOWS MM MM MM + TOTAL AGE NO.

GROUP TESTED NO. % NO. % NO. % Minimum - - 1 1 Moderate 2(1') 2 Adults 19 17 89.4 1 5.3 1 5.3 Advanced 1* 1 1* 3 Children 20 20 100.0 - - - Pulmonary Ganglion 1 1 2 4 Total 39 37 94.8 1 2.6 1 2.6 Pleural Total 2 2 6 10

women and utensils. In order to prevent indis- No. X-rayed 105 11 32 148 Percentile of contact with mem- criminate and undisciplined pulmonary shadows 1.9 18.2 18.7 6.7 bers of our civilization, the administrators were able to resettle the whole 54-member tribe *Positive sputum. 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 different forms such as the so- results. called infant form in adults and the rapidly In 1967 we repeated the research among the evolving acute and miliary forms, similar to Txukarramai 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 Suiá and Txukarramai 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 Suiá 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. Reactions to tuberculin test (PPD-Rt23-1 UT) TABLE 9. Reactions to tuberculin test (PPD-Rt23-1 UT) by age group among Txukarramai Indians of Xingu by age group among Txukarramai Indians of Xingu National Park, Brazil, August 1966. National Park, Brazil, July 1967

TESTS READ TESTS READ

0-4 5-9 10 0-4 5-9 10 MM MM MM + 1MM MM MM + AGE NO. AGE NO. i GROUP TESTED NO. % NO. % NO. % GROUP TESTED NO. % NO. %O NO. %O

0-4 36 31 86.1 1 2.8 4 11.1 0-4 34 32 94.0 1 3.0 1 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 20-29 28 18 64.3 1 3.6 9 32.1 20-29 33 16 48.5 1 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 Total 166 112 67.5 6 3.6 48 28.9

73 4

FIGURE 2. X-rays of the four known BK-positive cases in the Xingu area

74 tuberculosis among these Indians, in its clinical, TABLE 10. Reactions to tuberculin test (PPD-Rt23-1 age group among three tribes of Mato Grosso radiological, and even epidemiological aspects, UT) by State, Brazil, September-October 1965 can be equated with that of peoples with a long experience with BK. TESTS READ one time undergone a so- Had these tribes at 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. TESTED NO. % NO. % NO. % Is it possible that the BK responsible for the GROUP disease among them is less virulent than that 0-7 584 559 95.7 4 0.7 21 3.6 responsible for the disease among whites? This 8-14 634 537 84.7 37 5.8 60 9.5 hypothesis may be answered by the result of 15-49 746 525 70.4 71 9.5 150 20.1 50+ 145 98 67.6 19 13.1 28 19.3 Professor Magarao's research mentioned above. In any case, we believe that the Park is an Total 2,109 1,719 81.5 131 6.2 259 12.3 appropriate environment for planned and controlled scientific research on the response of primitive populations to the introduction of 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 1 consider it useful to conclude with Table 10, with white civilization for at least a hunderd which shows the rate of tuberculosis infection years (9).

REFERENCES 1. BALDUS, H. Bibliografia Crítica da Etnologia de 1954 apresentado ao Diretor do S.P.I. (pp. 131- Brasileira. Sao Paulo, Comiss5o do IV Centenário da 141). Rio de Janeiro, Serviço de Proteçao aos Indios, Cidade de S5o Paulo, Serviío de Comemoraçoes 1955. Culturais, 1954. 8. NUTELS, N., and L. V. DUARTE. SUSA-1960: 2. BARUZZI, R. G. Determination de l'Etat de Cadastro tuberculinico na área indígena. Separata Santé d'un Groupe d'lndiens lsolés et des Mesures da Rev. SNT 5(19):259-270. Rio de Janeiro, 1961. Prophylatiques Necessaires a leur Preservation, Par- 9. NUTELS, N., J. A. NUNES DE MIRANDA, 1. BEJGEL, que Nacional do Xingú, Brésil. Memoire présenté a 1. YAMASAKI, and A. FRAGA HAUTEQUEST. Atividades l'Institut de Médicine Tropicale "Prince Léopold". do SUSA em 18 aldeiamentos indígenas 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, O. L. RAMOS, and P. L. PONTES. Ocorrencia Tuberculin reactions, X-ray and bacteriological de blastomicose queloideana entre índios Caiabí. studies in the Cayapó Indians of Brazil. Tubercle Rev. Inst. Med. Trop. Sao Paulo 9(3):135-142. 48(3), 1967. 4. CARNERI, 1., N. NUTELS, and J. A. MIRANDA. 11. OLIVEIRA, R. C. DE. O processo de assimila~ao Epidemia de leishmaniose tegumentar entre os dos Terena. Rio de Janeiro, Museu Nacional, 1960. indios Waurá do Parque Nacional do Xingú. Rev. 12. RANKE, K. E. Beobachtungen fiber Bevolke- Inst. Med. Trop. Sao Paulo 5(6):271-272, 1963. rungsstand und Bevolkerunsbewegung bei Indianern 5. GALVÁO, E. Cultura e sistema de parentesco das Central-Brasiliens. Correspondenz-Blatt der Deut- tribos do Alto Xingú. Antropologia (Río de Janeiro) schen Gesellschajt #iJr Anthropologie, Ethnologie No. 14, 1953. und Urgeschichte XXIX(11), 1898. 6. MATTOS, R. B. Acuidade visual para longe e 13. RIBEIRO, D. Convivio e contaminaçao. Efeitos jrequencia de discromatopsia em indios brasileiros. dissociatívos da depopulaq5o provocada por epi- Descri¢ao de alguns aspectos oftalmológicos em demias em grupos indígenas. Separata de Sociologia indios examinados. Tese apresentada para o concurso (Sao Paulo) XVIII(1), 1956. e culturas indígenas do de docencia-livre da cadeira de clmnica oftalmologica 14. RIBEIRO, D. Línguas da Escola Paulista de Medicina. Sao Paulo, 1958. Brasil. Separata de Educaqao e Ciéncias Educacionais. 7. MOTA, J. L. DA. A epidemia de sarampo no Rio de Janeiro, Centro Brasileiro de Pesquisas Xingú. Condensaaqo do relatório em 10 de outubro Educacionais, 1957.

75 15. RIBEIRO, D. Política indigenista brasileira. Rio 18. STEINEN, K. VON DEN. O Brasil Central. de Janeiro, Ministério da Agricultura, Servigo de Tradujao de Catarina Baratz Cannabrava. Rio de Informaj6es Agrícolas, 1962. Janeiro, Companhia Editora Nacional, 1942. (Origi- 16. RICH, A. R. Patogenia de la tuberculosis. Ver- nal edition Leipzig, Brockhaus, 1886) sion castellana por el Dr. O. C. Croxatto. Buenos Aires, Alfa, 1946, p. 651. 19. STEINEN, K. VON DEN. Entre os aborígenes do Brasil Central. Tradu5ao de Egon Schaden. 17. SILVA, M. P. DA. Contribui•Fo para o estudo do Sepa- sangue periférico e da medula óssea em indios do rata renumerada da Revista do A4rquivo 1:XXXIV- Alto-Xingú. Tese de doc¿ncia-livre de hematologia LVIII. S5o Paulo, Departamento de Cultura, 1940. apresentada ao Departamento de Medicina. Sao 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 the north 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 and Brazil. 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 are present here today: Dr. James Neel, 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, the anthropologist area extending from the Equator to 5 degrees whose work, study, and life among these In- dians since 1964 2 was so essential to our mission; and myself, a pediatrician with interest in and some experience with the diseases and 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 expedition was in essence a cooperative venture of the University of Michigan and the Venezuelan Institute for Scientific Research, and in particular Drs. Layrisse and Tulio Arends of the 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 100 villages scattered throughout the region. They are very primitive forest-dwellers who practice a slash-

1Investigations supported in part by U.S. Atomic Energy Commission Grant AT (11-1)-1552. 2 Chagnon, N. A. Yanomamo Warfare, Social Organi- zation and Marriage Alliances. Ph.D. thesis, The Univer- sity of Michigan (1966).

77 and-burn agriculture along with hunting and be at the mission in connection with a study gathering. Although some mission stations have on thyroid physiology. been established along the Upper Orinoco and The boy never developed the characteristic its tributaries since early in the past decade, the morbilliform 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 vaccinated at once stockade, with families living in hammocks with no gamma globulin coverage. It turned out under the shelter of the enclosure. In general that the boy indeed did have measles and that the people are healthy and handsome, well pro- visitors to the mission meanwhile had returned portioned and strong but almost pigmoid in to their own villages carrying the virus in its stature. Their cheerful, friendly countenances incubating form with them. Thus started a full- bely the aggressiveness and hostility for which scale epidemic, which, by the time our 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 along a hundred 1967 demonstrated that this was a virgin-soil miles of waterways and tributaries and adjacent tribe and because measles was known to have inland regions involving no fewer than 15 penetrated into the nearby regions, practically villages. surrounding the Yanomama territory by 1967 Within a few weeks we had exhausted our (see map), we brought along with us into South vaccine supply. Observations on these Yano- America 2,000 doses of donated Edmonston- mama Indians of the effect of the wild measles strain attenuated live measles vaccine, plus and of the measles vaccine with and without gamma globulin.3 Half of these doses we the concomitant gamma globulin form the diverted to the mission stations on the Brazilian basis of this report. Since it was neither con- side, where measles was beginning to penetrate ceived nor designed as a scientific study in itself the region, and the remainder went with us. We and since our data-collection responsibilities did had intended to vaccinate and to advise the not permit a prolonged stay in any one village, missionaries on the procedure toward the end our observations are of necessity disjointed and of our scientific studies. We did not anticipate fragmentary, but still, we believe, of sufficient encountering measles during our two-to-three- interest and value to warrant presentation. month expedition. It was thus a fortuitous occur- Most of the Indians and also the several rence that our arrival into the region was almost Brazilians we observed became very ill and pros- simultaneous with the introduction of measles trate 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, Director of the Venezuelan Insti- the measles epidemic. The complication of otitis tute for Scientific Research, who happened to media appeared to be practically nonexistent. We were particularly impressed with the 3We are most grateful to Parke, Davis and Co., devastating effect of measles in its near-total Phillips Roxane Laboratories, and Lederle Laboratories for their donation of the vaccine. involvement of the Indian community. When

78 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 for necessary nursing better organization and provision of medical care. We have even seen several instances where care, it was only 2 per cent. three generations of Indians were simultane- The use of measles vaccine permitted us 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 and measure the effects in a fatalistic depression was usually one of at their height, we found no real complications panic. Sometimes the well members of the but considerable reaction, especially in terms of village even abandoned the sick, and normal febrile response. That this response was asso- community structure and function were lost. ciated with the production of protective measles Village fragments often fled to other villages antibodies was borne out clinically. To our for haven, thus increasing the spread of the knowledge, no persons vaccinated prior to the disease. exposure to measles later developed the disease. In three villages with approximately 170 cases This protective value of the vaccine showed up of measles there were 29 deaths, a rate of 17.1 in peculiar ways, as for example in the case of per cent. Most of these were due to associated an unvaccinated mother who became very ill pneumonia, complicated by dehydration. This with measles while her previously vaccinated despite some measure of applied, though less and healthy youngster was playing about and than optimum, nursing care. In a fourth village, pestering her for attention she could not give. the authors were present for two days at the This is just the reverse of the situation com- peak period of measles reaction involving nearly monly seen in civilized societies. 40 Indians, a majority of them extremely ill. Searching the literature for reports of reac- Vigorous and dramatic treatment included fluid tions of other peoples to similar vaccine both administration, cooling measures, aspirin, and with and without concomitant use of gamma long-acting antibiotics. No deaths occurred dur- globulin, we observed that these Indians in gen- ing our stay, but we did hear later that subse- eral seem to react with a greater febrile response quently two Indians died (approximately 4 per than has been reported in North American Cau- cent mortality). casians, 6 Europeans,7 Icelanders, 8 Africans 7, 9 1 These and other observations supported our and Micronesians. ' Again, however, as with feeling that differences in medical and nursing care play the major role in the measles-mortality 5Pert, 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. should be recalled that the mortality from 6 Krugman, S., J. P. Giles, A. M. Jacobs, and H. Fried- measles among Caucasians in the United States man. Studies with live attenuated measles virus vaccine. only half a century ago was in excess of 3 per Am. I. Dis. Child. 103:353-363, 1962. 7 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- 231, 1966. -of this reduction occurred in the period before 8 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.4 Other evidence that the death rate in Hlth. Org. 30:753-762, 1964. highly susceptible groups can be substantially 9 Meyer, H. M., Jr., D. D. Hostetler, Jr., B. C. Bern- heim, N. G. Rogers, P. Lambin, A. Chassary, and J. E. reduced by elementary organization and medical Smadel. Response of Volta children to live attenuated 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. '0 Brown, P., M. Basnight, and D. C. Gajdusek. 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: Am. . Dis. Child. 103:224-226, 1962. 115-122, 1965.

79 the wild measles, we are unable to define accu- become "virgin-soil" 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 Panum re- 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 for some 65 wide variety of diseases is 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 1 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 to me 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 limited with less than optimal clinical results in the evidence for a greater innate susceptibility of management 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 the will not settle for anything less than the best. primary susceptibility of the Indian to certain This 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 presentations represent very fine contributions to our General Discussion accumulating knowledge on this subject. In previous meetings of this committee, several of Moderator: As I am sure you gathered from us-among them Dr. Neel and I-expressed the that capsule presentation, there were some prob- belief that evidence that a genetic predisposition lems in conducting ourselves 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 this experience adequately. seems to me the evidence in both these studies is in that direction. As Dr. Nutels said, Dr. Nutels in his presentation gave data on it is conceivable that the strain of mycobacteria how the mortality rate in an Indian group can either may not have be lowered by even what I think he would agree been tuberculosis or may is less than adequate medical care. Certainly be attenuated, but 1 do not think that is very this was our experience-that 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 hall to a third have tuberculosis infection and disease. 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 that after having to do with the culture, and in 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 must munity against infection with tubercle bacilli. remember first of all that it may be a very By this I mean that practically all members of severe disease in any adult. Adults with measles a group exposed under the proper circumstances, 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 of infected. We know of course that infection leads measles-and the complications are what people to disease in only a minority of those infected; die of-can be very high, depending upon the the 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 large simply 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, 1 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 measles, 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- Tuberculosis infection can be spread and turated societies. What I am really saying is that probably usually is spread by the airborne route. our knowledge of the pathogenesis of both in- This means several things. One is that the infec- fections now gives us perfectly credible alterna- tious agent-in 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 not known, but it is an appreciable Centerwall: By simple inspection and physical interval-a matter of an hour or two. And the examination. The nutritional status of the chil- 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 was active 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 protein deficiency 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 aboriginal devoted 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 1

Introduction logic study (5) has confirmed the higher prevalence of "clinical gallbladder disease" among The etiology of gallbladder disease is uncer- Pima Indians than was established by identical tain, though the problem has excited consider- criteria among the predominantly Caucasian able speculation and controversy. Diet (11), community of Framingham, Massachusetts (7) female hormones (10), obesity (16), and dia- (Table 1). betes (13) have all been said to be associated purpose of this paper is to report pre- with cholelithiasis. In addition, differences in the The liminary data on a partly completed epidemio- frequency of gallbladder disease among various based on chole- racial groups have been noted. On the basis of logic study of gallbladder disease cystography in a population sample of Pima autopsy data Caucasians have been reported to Indians, a Uto-Aztecan tribe living in the desert have a higher frequency of gallstones than of the southwestern United States. Negroes (6), and several studies based on hospital 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 1 Presented by Dr. Burch. the National Institute of Arthritis and Metabolic

TABLE 1. Prevalence of documented gallbladder disease in 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 2.0 40-49 779 7 0.9 45 2.2 50-62 725 23 3.2 57 7 12.3 a Total 2,336 31 1.3 153 5.9 a

Females 30-39 1,037 24 2.3 45 16 35.6 a 40-49 963 58 6.0 51 20 39.2 a 50-62 873 88 10.1 62 21 33.8 a Total 2,873 179 5.9 158 57 36.0 a

ap < 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 River scout 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 that all houses were taken on each respondent. If the gallbladder was 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 of gallbladder stimulant and either the upright or the chest, cervical spine, hands, feet, pelvis, the decubitus film was repeated. thigh, and calf; a 12-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 perform a repeat gers); a urinalysis; and determinations of serum 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-cell agglutination 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 selected cholecystogram 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 they excluded 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 Iopanoic acid manufactured by Winthrop Labora- 3 Distributed by General Electric X-Ray Department, tories. New York. Milwaukee.

83 I SALT LAKE CITY 1

¡uchesne

U T~ A - H i

UNITED PUEBLOS

® R N E~W M E X A R I ~Z 0 N A N E:[WJ M E X C O

T. CHE,.....

PHOENIX :::: CARLOS,

T.c-,/

FIGURE 1. Indian reservations of southwestern United States. Gila River Reservation is 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 DOCUMENTATION MALES FEMALES 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 this report without knowledge of the age or sex CFSU Clinic 35 44 79 Number with gallbladder 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 0 2 2 the new interpretation was used. Two nonvisualizations 0 7 7 One nonvisualization 1 4 5 Results Abnormal cholecystogram at 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 female Pima Indians. In males the disease frequency increased with age, with 66.7 per cent bladder 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 cholecystograms. 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 Unit Clinic, the TABLE 2. Prevalence of gallbladder disease in 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. 0% Morbidity from gallbladder disease Males 15-24 5 0 0.0 In Table 4 the clinical importance of gall- 25-34 6 0 0.0 bladder disease among Pima Indians is shown. 9 1 11.1 35-44 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 37.5 or had a history of such symptoms noted in their medical records. Among the females, however, Females gallbladder disease resulted in considerable mor- 15-24 11 1 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 a previous cholecystectomy and 45-54 28 19 67.9 an additional 25 (27.2 per cent) had had gall- 55-64 19 12 63.2 65 and over 27 25 92.6 Defined as an affirmative answer to the question "Have you ever had pain in your stomach or vomited Total 136 92 67.6 after eating certain kinds of food?"

85 TABLE 4. Morbidity from gallbladder disease in Pima Indians

MALES FEMALES TOTAL

NO. %70 NO. % NO. %

Total examined 56 100.0 136 100.0 192 100.( With gallbladder disease 21 37.5 92 67.6 113 58.' Postprandial abdominal pain or nausea 6 10.7 64 47.1 70 36.5 Cholecystectomy 5 8.9 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 a gallbladder disease and possible associated con clinical problem among Pima women is best ditions 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 there was an association Discussion with diabetes. These data must be interpreted cautiously, These data, though as yet incomplete, indicate since the present study indicates that nearly that the gallbladder was an extremely frequent a third of the Pima males and nearly half of the site of pathology among Pima Indian females. Pima females without documented clinical Gallbladder disease was responsible for numer- gallbladder disease actually had gall- ous hospitalizations, which often culminated in bladder pathology. a major surgical procedure with its attendant In order to advance the study of cholelithiasis morbidity and slight mortality. Unfortunately, and to further explore the problem of gallstones we have no data on complications that may have in Indians, it would be desirable to supplement compounded the morbidity from gallbladder the epidemiologic studies currently being per- disease in the members of this sample. formed among the Pima with selected biochemi- Gallbladder disease was less common among cal 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 biliary formed. 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 contribut- changes found in pathologic bile have recently ing to gallbladder disease in the Pima-such as been reviewed (1). Since other groups of Ameri-

86 can Indians appear to have a high prevalence of It has been reported that peptic ulcers are rare the disease, it is possible that the bile of Ameri- among southwestern Indians (9). This observa- can Indians, perhaps as a result 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 bladder 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 if it 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 21 (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 with that of per cent of the 56 males and 47.1 per cent of the other racial groups, might add significantly to 136 females. Nearly a third of the males and khowledge 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 for clearing 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 the United States Public Health Service for their encour- absorption of dietary fats and of the synthesis agement and assistance. Special thanks are due to Drs. and excretion of cholesterol might give clues to W. J. Held and Jon E. Gustafson, the radiologists at the the etiology of gallbladder disease in this group. Phoenix Indian Hospital who read the cholecystograms.

REFERENCES

1. BOUCHER, I. A. D., and J. W. FRESTON. The 5. COMESS, 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 in contrast to Framingham, 2. BUMIN, J. J., T. A. BURCH, and W. M. O'BRIEN. Massachusetts. New Eng. 1. 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 cholelithiasis in the Negro factor in American Indians. Bull. Rheum. Dis. 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 gallstones in the mouse. Amer. 1. Physiol. Observations in the Framingham Study. 1. Chronic 209:473-478, 1965. Dis. 19:273-292, 1966. 4. CAPPER, W. M., T. J. BUTLER, J. O. KILBY, and 8. HESSE, F. G. Dietary study of Pima Indians. M. J. GIBSON. Gallstones, gastric secretion, and fla'u- Amer. 1. Clin. Nutr. 7:532-537, 1959. lent dyspepsia. Lancet, i:413-415, 1967. 9. HESSE, F. G. Incidence of cholecystitis and

87 other diseases among Pima Indians of southern 14. MILLER, M., T. A. BURCH, P. H. BENNETT, Arizona. IAMA 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 RoorT, P. A. SALMON, W. O. GRIFFEN, and O. H. tests. in Pima Indians of Arizona. Diabetes 14:439, WANGENSTEEN. Production of gallstones by pro- 1965. longed administration of progesterone and estradiol 15. SIEVERS, M. L. and J. R. MARQUIS. South- in rabbits. Surg. Forum 10:246-249, 1959. western American Indians' burden. Biliary disease. 11. KAMEDA, H. Gallstone disease in Japan. A lAMA 182:570-572, 1962. report of 812 cases. Gastroenterology 46:109-114, 1964. 16. VAN DER LINDEN, W. Some biological traits 12. LAM, R. C. Gallbladder diseases among Ameri- in female gallstone disease patients. A study of body can Indians. J. Lancet 74:305-309, 1954. build, parity and serum cholesterol level with a 13. LIEBER, M. M. Incidence of gallstones and discussion of some problems of selection in observa- their correlation with other disease. Ann. Surg. tional hospital data. Acta Chir. Scand. 269, Supple- 135:394-405, 1952. ment 1, 1961.

88 HYPERGLYCEMIA IN PIMA INDIANS: A PRELIMINARY APPRAISAL OF ITS SIGNIFICANCE

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

the There have been several reports in recent lation group, geographically isolated from over years indicating that certain groups of North other. Some 3,300 Indians aged 5 years and claim to American Indians have a high prevalence of reside in this community; of these 3,041 diabetes mellitus (5, 14). It has also been sug- be half- to full-blooded Pima. gested that the frequency and types of 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 those ance test at the National Institutes of Health 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 and related variables in a North load2 (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 fluoride and another specimen was placed in a plain glass tube. Glucose concentra- The Pima Indians have resided for at least tion was measured on the AutoAnalyzer, using several centuries in the region of the Gila River the modified Hoffman method,3 in plasma ob- in south-central Arizona, which is typical of the tained from the fluoride tube. Serum creatinine hot, dry Sonoran desert of the southwestern was determined on the additional serum sample. United States. The Pima were traditionally an Before drinking the carbohydrate load each agricultural group, and they or their forebears, to void, and two hours later a with the aid of irrigation water from the Gila subject was asked which was tested River, have farmed its valley for about two urine sample was collected, 4 When- thousand years (4). The majority of them now for protein and glucose with dipsticks. of reside on the Gila River Indian Reservation, to ever protein was found in a concentration the south and east of Phoenix, Arizona. approximately 30mg/100ml or more (> 1 +), a The inhabitants of the reservation reside in quantitative determination of the urinary pro- two main regions, one of which receives its tein concentration in a sample of the urine was medical care from the U.S. Public Health made together with determination of the urinary Service Indian Hospital in Phoenix and the Company, Elkhart, Indiana, or other from the PHS Indian Hospital in Sacaton, 2 Glucola, Ames l)excola, Custom Laboratories, Baltimore, Maryland. Arizona. The latter region contains several 3 Technicon AutoAnalyzer Method File N-20, Tech- villages that form an independent natural popu- nicon Instruments Corporation, Chauncey, New York, 1965. Company, Elkhart, Indiana. 1 Presented by Dr. Miller. 4 Labstix, Ames

89 creatinine concentration. Among those aged 15 and their outcome among the Pima women years and over a subsample was examined more aged 25 to 44 years on 1 January 1966 (1). 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.5 have been collected systematically since 1964. A prospective study of the outcome of the Results pregnancies among the Pima women living in the same community was initiated in October Table 1 shows the number of Pima subjects 1965. The modified glucose tolerance test was in the sample with hyperglycemia by age and performed on these women one to three days sex and the number with previously diagnosed after parturition, and data on the course of diabetes who have received specific 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 65 to 74 (26.2 per cent) and in the 55-to-64-year-old "The subsample consisted initially of a one-in-two females (40.8 per cent). aged 30 years and over but was ex- sample of those venous tended, after examination of the first 1,100 persons, to If "diabetes" is arbitrarily defined as a include all those examined aged 15 years and over. plasma glucose level of 160mg/100ml or more Though this is not a truly random sample, we believe two hours after a 75-gram glucose equivalent that it is unlikely to be significantly biased in relation to any findings associated with plasma glucose levels. load, the majority of cases of the disease were

TABLE 1. Carbohydrate intolerance among Pima (4/8-8/8) Indians of the Gila River Reservation aged 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 O O O 0.0 15-24 146 O 4 4 2.7 25-34 86 O 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 O 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 DISTRIBSUTION OF TWO HOUR POST LOAD VENOUS GLUCOSE LEVELS IN PIMA INDIANS tolerance test. The combined prevalence of previously and newly diagnosed "diabetes" reached a maximum of 45.9 per cent in the males aged ¡ - FEINALESP 65 to 74 years and 68.8 per cent in the females ¡í aged 55 to 64. In all age groups, especially those of 35 years and over, the prevalence of "diabetes" in the females exceeded the rate in the males, the differences being significant in each decade between 35 and 64 years. Thus, both male and female Pima Indians had an extra- ordinary frequency of "diabetes"-higher than that reported in any other population studied. PLASMA GL-COSEdM4100 NL Table 2 and Figure 1 show the distribution of FIGURE 1. Age- and sex-specific cumulative per- the two-hour plasma glucose results on all Pima centage distributions of two-hour post-load venous Indians examined regardless of whether they plasma glucose levels in Pima Indians had previously diagnosed diabetes, provided they values had a bimodal distribution. Using an did not have concurrently treated diabetes-i.e., iterative maximum likelihood procedure, these had not taken specific hypoglycemic medication distributions were dissected mathematically and within the period in which the two-hour glucose each of the observed distributions in those aged determination could have been directly influ- 25 to 74 years-for example, that shown in encéd.? Such a determination was made on 1,776 Figure 2 for males aged 55 to 74-proved to be of the subjects. The age- and sex-specific cumula- a statistically satisfactory fit for two overlapping tive frequency distributions of the plasma glucose values indicated that there was a relatively log1o normal (Gaussian) distribution curves. In those below 25 years, however, only a unimodal steep rise in the cumulative frequency distribu- distribution was found, as is shown in Figure 3 tion in the lower range of the glucose values in for males aged 15 to 24, which was a satisfactory all age groups. There was not, however, a uni- fit for a single log,, normal distribution (13). form increase in each successive age group at Hence the heterogeneity inferred from the all percentile levels. Below the median (fiftieth cumulative percentage distribution in the older percentile level) in both sexes the increases were groups appeared to result from the presence of relatively small, but the upper percentile glucose values were much higher in the older age groups of both sexes than in the younger subjects. This 3 MALES 55 74 suggests that, as age increased, the proportion of subjects with hyperglycemia increased markedly. _ 5- From this the inference was drawn that the plasma glucose levels in all except the youngest 5 o10- groups of each sex were not homogeneous but 5- consisted of two or more components. Further l S analysis indicated that in each sex and in all decades from 25 to 74 years the frequency distribution of the logarithms of the two-hour glucose 50 PLASMA GLUCOSt(.mg//o ml) 6 For patients taking tolbutamide or phenformin this period was 24 hours; for patients taking insulin or 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 i4 el O N4 V O tI 08 t, O r t, 0 I u I _ N t oE IEXOR s 1 --- 1 z

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92 Among those with diabetes concurrently under

20o MALES 15-24 treatment, retinopathy was observed in 20.0 per cent of the males and in 19.6 per cent of the

2 females. -> 15+ Proteinuria ii o 10- Other evidence that hyperglycemia among the

z Pima Indians was associated with vascular com- owZ v plications was investigated by examination for 5- evidence of renal disease. Table 4 shows the relationship of proteinuria ( 1 +) to the two- hour post-load plasma glucose level. The fre- 50 100 200 40 o PLASMA GLUCOSE (/-100m.L) 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 was a 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 distributions were 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 the was possible to make more quantitative esti- higher curve was relatively high. mates of proteinuria by determining albumin In order to determine whether the subjects in and creatinine concentrations in the two-hour the two distributions consisted of those with and urine samples. Since the daily excretion of without diabetes mellitus in the clinical sense, creatinine in the urine is relatively constant the relationship of other parameters of this (11), the albumin/creatinine (A/C) ratio will disease to glucose tolerance levels was 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 the pres- hour post-load glucose levels in excess of 200mg/ ence of microaneurisms, preretinal and vitreous 1OOml 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 and over who proteinuria was both more frequent in any de- had one or more satisfactory retinal photo- gree and also much more severe. graphs. The nephrotic syndrome associated with se- No definite evidence of diabetic retinopathy vere intercapillary glomerulosclerosis is recog- was observed in male or female Pima Indians nized as one of the important complications of under 35. In those above this age, it was found the renal disease of diabetes mellitus. The essen- only in those with the higher plasma glucose tial feature of this syndrome is the occurrence levels. In those with two-hour plasma glucose of excessive proteinuria, usually exceeding 3 levels of 200mg/100ml and over, retinopathy grams a day. It is noteworthy that of those with was approximately twice as frequent as in those plasma glucose values of 200mg/100ml and with levels between 160 and 199mg/100ml. over, 11 of the 39 with proteinuria on whom the

4 93 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/100ML) AGE GROUP WITH CONCURRIENTLY BY SEX 0-100 101-159 160-199 200+ TREArED DIABETES TOTAL

Males 15-34 years No. examined 43 43 0 8 1 95 No. affected 0 0 0 0 0 0 Percent 0.0 0.0 0.0 0.0 0.0 0.0 35-54 years No. examined 26 35 9 27 9 106 No. affetted 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 cent 0.0 0.0 9.1 15.8 20.0 6.3 Total No. examined 84 119 20 54 20 297 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 cent 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/lOD ml while not on medication. He had, however, lost weight and had carcinoma stomach with metastases, and in 1965 had a fasting blood glucose of 165 mg/100 ml.

94 0, TABLE 4. Proteinuria and two-hour post-load plasma glucose levels in Pima (4/8-8/8) Indians examined 1965-1967 (Ž1+ on dipsticks 4)

AGE GROUP PLASMA GLUCOSE LEVEL (MG/100ML) ~~~~~~AGE ~GROU~~p~~~WITH CONCURRENTLY 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 Percent 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 Per cent 5.9 7.8 15.6 17.9 37.0 10.7 Females 15-34 years No. examined 109 176 16 17 6 324 No. affected 4 7 2 5 0 18 Per cent 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 cent 4.4 4.2 6.5 24.0 24.0 10.0

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

PLASMA GLUCOSE LEVEL (MG/100ML) 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 69.2 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 concurrently excess of 250mg/100ml 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 2 00mg/100ml 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 glomulonephritis nine years pre- among the Pima Indians was strongly associated viously. with proteinuria and impairment of renal function, especially among the older subjects of both Serum creatinine levels sexes. In order to determine the frequency of subjects with significant reductions in renal Vascular calcification function, serum creatinine levels were examined. Evidence of disease of the peripheral vascula- Elevated levels were defined for this purpose as ture was evaluated in soft-tissue radiographs of 1.5mg/100ml and over in males, and 1.3mg/ the thigh (femoral artery) and the calf (tibial lOOml and over in females. Such levels were arteries), since vascular calcification has been found mainly among subjects with two-hour shown to be more frequent among diabetics plasma glucose levels of 200mg/100ml and over than among nondiabetics (3) and to develop and among those with concurrently treated dia- concomitantly with increasing duration of diabetes (Table 6). They occurred predominantly betes among the younger subjects with diabetes among the older subjects in the same groups in (15). Tables 7 and 8 show the frequency of 4* 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/100ml who had than in the females, was age-related, and was elevated serum creatinine levels were examined encountered 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 heart observed 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/100ml, was a tibial arteries was not found in any Pima Indians known diabetic of many years' duration. He had below the age of 35 years. In the older groups

96 TABLE 6. Elevated serum creatinine and two-hour post-load plasma glucose levels in Pima (4/8-8/8) Indians examined 1965-1967 (Males >1.5 mg/100 ml: Females> 1.3 mg/100 ml)

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 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 55 and over No. examined 19 64 17 26 16 142 No. affected 0 1 2 1 5 Per cent 0.0 1.6 5.9 7.7 6.3 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 ák 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

a See text.

97 TABLE 7. Medial calcification of the femoral artery and 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 59 49 3 7 O 118 No. affected O O o O O O 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 No. affected O O 1 2 1 4 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 O O o O O O 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 O O O 2 O 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 2 O 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 -- 4

98 TABLE 8. Medial calcification in the tibial arteries and 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 J 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 4: 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 Percent 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 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 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. affccted 0 3 2 12 7 24 Per cent 0.0 1.8 5.3 12.8 12.1 5.7

99 both sites were 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 with quéncy of "diabetic" pregnancy was therefore femoral artery involvement, the tibial arteries some 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 similar genital 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 had in the latter was 25.5 per cent, which is in the received a modified 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/100ml, those with either clini- mother generally two to three days postpartum, cal diabetes requiring hypoglycemic treatment or plasma glucose values of 160mg/100ml or but in 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 the consistent 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 "cNORMOGLYCEMIC` 'PREDIABETIC" 1'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 Congenital 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 mortality No. of births 815 391 47 1253 .4 No. of deaths a 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 TABLE 10. Infant birth weight and maternal post diagnosis was sometimes based upon less-than- partum two-hour post-load plasma glucose levels in ideal criteria. The rate was approximately twice Pima (4/8-8/8) Indians examined Oct. 1965-Dec. 1967 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 rates among Pima MATERNAL > 4000 GRAMS subjects with hyperglycemia were substantially PLASMA AT BIRTH higher than among those with "normal" plasma GLUCOSE NO. OF

(MG/ 100ML) PREGNANCIES NO. 57 glucose levels.

0- 63 17 0 Post-mortem study 64- 80 49 2 81-100 90 5 Of the Pima Indians who died since our 101-126 76 5 initial study in 1963, autopsy data were available _, 127-159 33 9 on 34 subjects with "diabetes" and on 19 with- 160-199 6 out. Evidence of nodular intercapillary glomu- 2l lerosclerosis was reported in six of the "dia- 200+ 3 betics" and in none of the others. Seven of the Total 274 24 34 diabetics had myocardial infarcts, compared to 2 of the 19 "nondiabetics." Tuberculosis was Mortality found in 7 "diabetics" and in only 1 "nondia- betic." Pyelonephritis, howover, was reported as Death certificates have been collected on the frequently among the "nondiabetics" as the Pima Indians of the Gila River Indian Reserva- "diabetics." Amputations had been performed tion since 1964, and autopsy data were obtained on 2, and mucormycosis was present in 1 of on persons who have died since 1963. About half those with "diabetes." of those who died received post mortem examinations. Over a four-year period there were 53 Discussion male and 49 female deaths among persons aged The data presented show that hyperglycemia 15 years and over who had available glucose is extremely frequent among the Pima Indians. estimations or whose diabetic status was known, As complete glucose tolerance tests were not per- excluding those due to accidents or violence formed routinely and as only a single venous (Table 11). glucose determination made two hours after a For comparative purposes the subjects were 75-gram glucose equivalent load was available classified as "diabetic" on the basis of a clinical on all subjects, the usual criteria for the diagno- history of diabetes and/or high glucose values. sis of diabetes mellitus could not be applied. Age-standardized death rates for subjects with In order that comparisons might be made with "diabetes" in both sexes were found to be higher other studies, however, the frequency of subjects than those of the "nondiabetics," although the who had received specific treatment for diabetes was determined, as was the frequency of subjects TABLE 11. Four-year mortality, excluding accidental who had two-hour venous plasma glucose levels and violent death (1964-1967) in Pima Indians aged 15 years and over of 160mg/100ml and over. It was felt that this level, which corresponds to a whole-blood sugar MALES FEMALES of approximately 140mg/100ml, was suitable for comparison with several studies in which some- Total no. deaths 53 49 No. with diabetes 27 34 what similar testing methods were used. It was Age-standardized rates/ found that known diabetes and hyperglycemia, 100 persons as so defined, occurred more frequently among Diabetics 11.2 9.8 the Pima Indians than in any other population Nondiabetics 7.8 4.4 thus far studied. Unusually high frequencies of

101 diabetes have been found among the Cherokee also been performed on the same subjects, and Indians of North Carolina, but the prevalence the 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 greater than had that myocardial infarction occurred twice as fre- been found among Alaskan Eskimos (9) and quently 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 among Pima Indian subjects on To determine whether hyperglycemia in Pima whom glucose determination had been made, Indians was associated with the chronic vascular either in the course of routine medical care or complications that are characteristic of diabetes for research purposes, was shown to be greater in other populations, analyses were made to among 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 Joslin tion of their plasma glucose level or previously Clinic patients, which is mainly related to the known diabetes. Retinopathy, as shown on fun- chronic vascular complications (2). Though dus photographs, and impaired renal function, similar processes appear to be responsible for 4 as indicated by elevated serum creatinine levels the excessive Pima mortality, the number of .4 or heavy proteinuria (A/C ratio >3.0), were deaths is so far too small for deducing the rela- found exclusively among those with two-hour tive importance of the various types of vascular glucose levels in excess of 200mg/100ml or with complications. Serial studies of renal function concurrently treated diabetes, except in rare 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 all such a definition in the future. degrees was more frequent among those with The frequency of "diabetic" pregnancy among high glucose levels or 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 2 00mg/100ml. of significantly increased perinatal 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 .4 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 among hyperglycemia and are ameliorated by the spe- 4 such subjects. cific hypoglycemic medications, the substantial 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 of diabetes mellitus jects over the whole range of glucose levels but among the Pima Indians. While only 40 per cent was again more frequent among those of the of the subjects with glucose values of 160mg/ same age and sex with glucose levels in the upper 1OOml and over had received such therapy, they range or with concurrently treated diabetes. included 60 per cent of the females and 50 per Standard twelve-lead electrocardiograms have cent of the males aged 55 to 64 years with such

102 levels. This observation suggests that substantial differing tribes with high and low prevalences numbers of the younger persons with hyper- of diabetes at present could contribute to our glycemia will become symptomatic in the future. knowledge of the etiology, especially if identical Before the systematic study of the Pima methods of evaluation could be used in several Indian community was begun, diabetes mellitus of the groups. Such investigations could be was locally recognized as an important health enhanced by parallel study of groups of the problem but the true frequency of hypergly- same race in Central and South America. As cemia and of the vascular manifestations were these groups are subjected to change and as their unknown. Although some other southwestern life expectation is increased, it is possible that Indian tribes are recognized as having a high others may go the same way: in them diabetes frequency of the disease, others are said to have mellitus may emerge as 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 the reasons for < the differences could be identified, would be an Acknowledgments important contribution to our understanding of We gratefully acknowledge the help and assistance of the etiology of diabetes mellitus. many persons who have contributed to this study-in /, A definition of the geography of the disease in particular, Dr. E. J. Ballintine for his evaluations of the the southwestern American Indian seems to be retinal photographs, Dr. F. A. Rose for the soft-tissue needed as a matter of some urgency, since these radiographic readings, and Miss P. E. Sperling for the emerging groups are likely to be subjected to majority of the laboratory determinations. We also thank the Division of Indian Health, the Tribal Council, radical and rapid change. Some may lose their and members of the Gila River Indian community, identity; others may adopt ways widely differ- whose collaboration and cooperation made this study ent from those of their forebears. The study of possible.

REFERENCES

1. COMESS, 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 Eskimos. fAMA 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 Treatment of 3. FERRIER, T. M. Radiologically demonstrable Diabetes. Amsterdam, Excerpta Medica Foundation, arterial calcification in diabetes mellitus. .dust. Ann. 1965, pp. 722-726. Med. 13:222-228, 1964. 11. PETERS, J. P., and D. D. VANSLYKE. Quantita- J' 4. HAURY, 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. Y. 5. JOHNSON, J. E., and C. W. McNUTT. Diabetes Diabetes 16:346-351, 1967. mellitus in an American Indian population isolate. 13. RUSHFORTH, N. B., P. H. BENNETT, A. G. Texas Rep. Biol. Med. 22:110-125, 1964. STEINBERG, T. A. BURCH, and M. MILLER. Studies 6. KENT, G. T., and J. R. LEONARDS. Mass screen- of plasma glucose measurements in the Pima Indians in a metropolitan area using finger ing for diabetes of Arizona. I. Application of a composite of two blood glucose after a carbohydrate load. Diabetes Gaussian distributions as a model for plasma glu- 14:295-299, 1965. cose distributions. In preparation. 7. McDoNALD, G. W., and G. F. FISHER. Diabetes J. M. ROBEY, D. F. prevalence in the United States. Public Health Rep. 14. STEIN, J. H., K. M. WEST, 82:334-338, 1967. TIRADOR, and G. W. McDoNALD. The high preva- 8. MILLER, M. Diabetic pregnancy and foetal lence of abnormal glucose tolerance in the Chero- survival in a large metropolitan area. In B. S. Liebel 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. Natural course and prognosis of Foundation, 1965, pp. 714-717. juvenile diabetes. Diabetes 5:445-450, 1956.

103 MALARIA IN THE AMERICAN INDIAN 1

George G. Giglioli

Malaria prevails, or has prevailed up to re- being made in respect to the different ethnic cently, over very wide areas of the Western groups that might be present. Given the magni- Hemisphere, lying between latitudes 50 ° N and tude of the task, the remote, scattered, and eco- 40° S and from the shores of the Atlantic Ocean nomically unimportant Indian communities to those of the Pacific. The American Indian were-to a great extent-missed, overlooked, or ranges from Canada 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, 4 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; the Carib, formerly very data are conspicuously absent from the literature, numerous and widely distributed, but now re- and none of the principal malaria eradication duced 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 antimalarial Demerara, 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; the Arekuna, of the upper therapy, not on epidemiology. Excessive hopes Cuyuni and Mazaruni mountains and savan- were founded on the new insecticides with long- nahs; the Makusi, of the North Rupununi 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, efforts in the sparsely inhabited it may be noted, however, that few environ- .« countries of the American tropics were concen- ments present greater stability and uniformity trated on the more populous areas, no distinction than the equatorial rain forest, which covers

104 most of the Guianas and Amazonia, where northeastern 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 this been recorded only among the Bush Negroes of vast region that the majority of American In- Surinam-descendants of runaway slaves who dians live still fully exposed to the ravages of have lived, for many generations, isolated in uncontrolled and untreated malaria. the Guyana forest, where other races, equally The study of racial differences in respect to exposed, 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- parasites presents many difficulties; for valid laria. 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 are Van der Kuyp (12) found a very high P. falci- rare, and this is probably the reason why studies parum rate in both children and adults; the on this aspect of malariology are few and the mortality at all ages was low, however, and they conclusions drawn somewhat tentative and were 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 acquired adults. The actual parasite rate averaged 70.5 practically total immunity to Plasmodium vivax per 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 of 30 per cent, from 698 to 901, with an average tolerance to the effects of parasitization. Rai- annual death rate of 21, a birth rate of 39 per chenow (10), in 1929, working among the 1000 inhabitants, and infant mortality of 113 Hausa of the Cameroons, reached the conclusion per 1000 births! Creole Negroes from the that toxin resistance was inherited but parasite Surinam coastland, living in similar surround- resistance was acquired by continued exposure ings, showed the same or even higher infection and reached its peak in early adulthood; there- rates and suffered severely. after, the parasite rate and the degree of para- The Amerindians of Surinam, living under sitization remained low and more or less stable. the same conditions as the 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 Holoendemic malaria is regarded as the 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 (108 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 the en- young children, but the adults enjoy good larged adult spleen (2.2 Schuffner units), surpassing anything we encountered in Bush Negroes, are evidence health so far as malaria is concerned. Holo- 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 size of the swollen spleen are peculiar of Indian malaria, been recorded in other rather restricted situa- as distinct from the Bush Negro and Creole malaria one tions, as, for instance, among the aboriginals of meets everywhere in Surinam.... Besides a strong

105 splenic 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 malarial infection 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 in their seminomadic life which enables them to carry malignant Clinical experience in sugar-estate hospitals malaria infection from the Bush to the Savannah. showed that the majority of adults went through American Negroes, descendants of slaves life afflicted by large-often very large-spleens brought from Africa between the sixteenth and and failed to come to terms with the infection; the nineteenth centuries, who have lived for they suffered frequent attacks of fever, and generations under conditions of more or less anemia of some degree was practically uni- moderate malaria endemicity, still enjoy some versal. 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 the its 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 races living in the same mainly in Bengal, Bihar, Orissa, the Central i environment (2). Provinces, and Madras, where malaria is mostly, In Guyana, people of six races have been though not exclusively, hypoendemic or meso- living for many generations in the same general endemic, and seasonal in its 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 to the the Chinese constitute small minorities and live suffered when they were transferred hyperendemic Guyana coastland. mostly in Georgetown. The rural population of the coastlands comprises large numbers of Further inland, along the tidal course of the 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 spraying of the Arawak and Akawai tribes and by per- operations were introduced, malaria used to be sons of mixed race, usually with Amerindian, stable and hyperendemic throughout the coast, European, and Negro blood in very variable from the Berbice Estuary to the Pomeroon proportion. There are few permanently settled River (7). Systematic surveys of schoolchildren Negroes, but a considerable fluctuating popula- were conducted between 1937 and 1946, separate tion of this race, mainly adult males, is employed records being kept for the two main races. The in the mining and timber industries. At the time dis- parasite rate showed no significant racial varia- when malaria was hyperendemic in these tion; splenic reaction in the Negroes was, how- tricts, the Negroes came not only from the in- ever, decidedly and consistently more moderate, tensely malarial coast but also, and in high more mildly malarial and decreased rapidly with age so that spleens of proportion, from the Leeward and Windward Islands and from

1 According to the last census, the population of malaria-free Barbados. Whatever their origin, Guyana in 1960 totaled 560,406, comprising East Indians, their reaction to malaria infection followed the 267,840; Negroes, 183,980; Mixed, 67,189; Amerindians, 25,450; Chinese, 4,074; Europeans, 3,218; and Others, same pattern: they contracted the infection soon 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- Akawai, Arakuna, and Carib tribes. On the ously, all ages being affected; mortality and Rupununi Savannahs and the Pakaraima Pla- morbidity were certainly graver in children, but teau, the inhabitants are 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; the largest spleens recorded, 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. falci- 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 increased traffic with the with the following racial incidence (4): coastlands. Thus, the savannahs of the Ru- pununi, 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 visited the 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 malarioastic survey 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 these and 46 per cent of the adults had large spleens, syndromes in Negroes might be due, to some which 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 area 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 tolerance ment; it was said that before malaria appeared to the infection, suffering from its effects at on the savannah, in 1930, old and very old all ages. people were numerous. (One rancher who came In the far interior the population is widely to the area in 1922 had known two Macushi

107 ancients who remembered quite well the British incidence of the disease has been drastically military expedition to Pirara in 1843.) During reduced. The rapid increase that has followed this 1943 survey, many settlements in the Indian in all the tribes is a clear indication of the reservation along the northern Kanuku Moun- disastrous effect of malaria on this population tain foothills were found deserted and some of group over the past years; in 1968, their number the houses burned, as is customary after the is estimated at 32,000. death of an inmate. This high susceptibility and reactivity of the This high susceptibility of the Guyana Indians Amerindian to malaria is without doubt related to malaria is without doubt one of the major to the recent introduction of the infection in the causes of their decline over the past 150 years. Western Hemisphere. It is generally accepted Smallpox, measles, yellow fever, 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. cas the new arrivals found, wherever they The early Dutch colonists sought the alliance landed, native anopheles capable of acquiring of the Indians-particularly the Caribs, then a and transmitting the disease; given the pro- very numerous and warlike tribe-relying on longed infectivity of malaria subjects, the lack them for the procurement of Indian slaves of curative medicines at the time, and the wide captured in raids on villages and Spanish mis- diffusion of anopheles, it is not surprising that sion settlements on the upper Cuyuni and serious early outbreaks 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 relied on In spite of tradition, it appears that even the Indian help in subduing slave rebellions and in Peruvian Indians were ignorant of the properties the capture of fugitives. From 1778 to the end of cinchona bark. Richard Spruce, the botanist, of slavery, annual official receptions were 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. (11). It was he who was responsible for the The figures that have been given, however, are collection and shipment of seeds and young 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 of any inter- found that the cascarilleros (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 powerful and feared but regarded his explanations with skepticism tribe, practically vanished; only a few survive at and suspicion, for they were convinced that the the present day on the upper Barama and bark was used for the extraction of a valuable Barima rivers. In 1848, after proper 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 aboriginal or semi-aborigi- the spread of malaria to even the most remote nal state (3). and isolated Indian communities. The Wai-Wai Systematic malaria-control operations were Indians are an entirely primitive tribe that has inaugurated in the interior in 1947; though its main settlements in Brazil on the Mapueira eradication has not as yet been achieved, the River, which flows, through the Trombetas, to

108 the Amazon, some 400 miles to the south. A few localities where conditions for abundant 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 rain entire family, he spends a large part of his forest. Cassava graters and feather ornaments existence elsewhere-working on his farm, have been traditional items of trade between usually 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- ment-as a guide, a porter, or a boathand, bleed- siderable numbers of Wai-Wai from across the ing balata (wild rubber), collecting Brazil nuts, border. In 1966 a group of 80 paid their first and so on-and when so engaged he is usually visit to the mission, after a journey of several accompanied by his family. When away from weeks; of these, 14 were found infected with their permanent home, the Indians camp under malaria, P. falciparum, P. vivax, and P. malariae temporary shelters, ranging from a few large all being identified. palm leaves stuck into the ground to an open So far I 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 of In 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 are frequent. The large villages to be found in square miles; all of them, however, had worked the more civilized areas are due to missionary at, 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 gravel pits sunk in the and split palm trunks for walls and floors, and valley 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 few migration 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, for better hunting or fishing, 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 a eradication 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 transmission takes place not in villages and forest is eminently a place disease, prevailing in houses but at camping sites, on the farms, and

109 FIGURE 1. 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. falciparum 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. jalciparum, 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 in the forest. The main villages can be sprayed sory in proclaimed areas and regulating its with insecticides without excessive difficulty, but shipping, transportation, and sale in the interior. :he existence of remote, isolated houses is very Epidemiological evaluation preceded, 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 sprayable surfaces. Under such condi- greatly reduced the frequency of the disease; the tions, insecticidal methods can reduce, but not reservoir of infection within the population was eradicate, malaria, and their application is diffi- therefore relatively small. cult and expensive. Systematic spraying of 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 [lseful practice, for the Indian always travels square miles and a 1964 population of 39,550, with his hammock and the insecticide residue malaria disappeared within six months (the resists occasional washing. parasite rate in 1960 had been 8.3 per cent); Mass chemotherapy, through systematic, peri- 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. vivax infection was discovered cal treatment of P. falciparum infection needs among the Caribs of the upper Barama River- at least three days, and for P. vivax 14 days are a very inaccessible locality that could receive required. Pinning down an Indian for regular chloroquinized 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 outbreak at the (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 at top of map). iresults were by no means satisfactory, and ma- From there the infection spread rapidly to the 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 reached its 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 1961 to December 1965 an tion of 16,850; P. vivax was the only parasite attempt was made to eradicate these residual involved. The outbreak has been controlled by foci of malaria in the interior by means of DDT house spraying and the reissue of chloro- chloroquinized salt (9); in order to assess fully quinized salt, these techniques being used in the value of this new technique, DDT house combination for the first time. Only 15 cases spraying was suspended. This campaign 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 June 1968. of 48,500, mainly Amerindians. The campaign An identical incident occurred among the was conducted under strict administrative and Akawai Indians of the Wenamu and upper scientific control; specific legislation was enacted Cuyuni, along the Venezuelan border, the first making the use of chloroquinized salt compul- cases being discovered in April 1966, one year

111 after the cessation of antimalaria operations. in the forest during the rainy season, from April This again is a very inaccessible area, with a to November. Among 700 bleeders and their small population of Akawai Indians who can dependents (estimated at 2,800), 24 cases 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 proposed to issue to was responsible for all cases discovered. Indians all bleeders, along with the other rations they crossing overland from the Wenamu to the receive, 20 pounds of salt medicated with chloro- Kamerang, an affluent of the Mazaruni, spread quine (0.43 per cent chloroquine base) and the infection to the upper Mazaruni and its pyrimethamine (0.043 per cent), and, in addi- tributaries, a sparsely inhabited area with an tion, to give to every individual who can be estimated population of 3,000 Indians and "caught" before leaving for the forest a two-day diamond miners. During 1966, 280 cases were treatment with a combination of Fanasil (Sul- discovered in this sector. Many infected persons furthomidine) and pyrimethamine. The result reached the coastlands, but were intercepted by of this new approach will not be known till the systematic examination of plane passengers from end of the year. the interior. DDT house spraying and chloro- These various developments in the course of quinized salt (issued free to Indians) have the Guyana campaign illustrate clearly the brought the situation under control, though peculiar difficulties encountered in malaria con- progress has been slow, owing to the very trol and eradication operations among the Amer- difficult terrain. indians. As I have said, in the early planning of It is 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 economically more important and duction of chloroquinized salt accounted for 57 politically more vocal. 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 malaria among the Indians who inhabit the savannahs of the Ru- Indians assumes a new dimension, for it is pununi, close to the Brazilian border, chloro- evident that eradication will remain a drearni quinized salt failed, after a favorable start until means have been discovered to extend during 1961, owing to the introduction of a control to even the most remote of Indian chloroquine-tolerant P. falciparum strain from settlements. 1 have shown how rapidly and the neighboring Rio Branco Territory (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 at pres- adequate. DDT house spraying, interrupted ent, medicated salt appears to be the most since 1960, was reintroduced in 1962, and as promising, since it can find its own way far soon as systematic spraying operations were beyond the reach of organized public health established by the Brazilians across the border, agencies. We do not yet possess, however, the beginning in January 1965, malaria transmission ideal drug for this purpose. Those in use today, on the savannahs was interrupted. Silvatic 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 occasional cases of P. falciparum emergence and favor the diffusion of drug- malaria continue to occur, but exclusively among tolerant or drug-resistant infections, introducing Indians engaged in bleeding balata, who camp therapeutic problems which may be serious.

112 I have shown how, in Guyana, chloroquinized for the fish and game on which they subsist and ;alt was completely successful against all forms which tend to abound in certain seasons and not f native malaria parasite, only failing where in others; the traditional method of preservation md when a chloroquine-tolerant strain was in- employed is smoking. The entirely primitive Iroduced from Brazil. Our campaign, it should Wai-Wai Indians, for instance, who reach be emphasized, was inaugurated at a stage when Kanashen Mission from the depth of the the reservoir of infection had been greatly de- Brazilian virgin forest, take to the use of salt pleted by DDT spraying. Under such conditions, with great readiness as soon as it is made the probabilities of the presence of resistant available. strains were much reduced. With the drugs at Obviously, spreading the use of salt to the Present available, we believe that medicated salt more remote tribes will take years, and the should not be used under conditions of high endemicity. same can be said for eradicating malaria where Indians still in the aboriginal stage of civiliza- primitive Indians exist; at the present stage of tion are not acquainted with commercial salt malariological knowledge and antimalaria tech- and manufacture salt substitutes by washing nique, it appears likely that treatment and pre- jertain soils or from the ash of certain palms. vention may be extended more easily to the Natural deposits of salt are not likely to occur primitive and semiprimitive Indians through throughout the vast equatorial rain forest. On the established channels of intertribal barter 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 medication have ieasoning their food but a valuable preservative to be found.

REFERENCES 1. BOYD, M. F., and W. K. STRATMAN THOMAS. 6. GIGLIOLI, G. Malaria Survey of the North Studies on benign terian malaria. 4. On the re- Rupununi Savannah. Georgetown, 1939, pp. 1-14. fractoriness of Negroes to inoculation with P. vivax. (Legislat. Council Paper No. 3/1944) Amer. 1. Hyg. 18:485-489, 1933. 7. GIGLIOLI, G. Malaria, Filariasis and Yellow 2. CLARK, H. C. A comparison of the spleen and Fever in British Guiana: 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. fedical Department, United Fruit Company, Seven- 8. GIGLIOLI, 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, Amerindian problem. In Development Report. 1959. Georgetown, Government of British Guiana, 1948, 9. GIGLIOLI, G., F. J. RUTTEN, and S. RAMJATTAN. "op. 10-22. (Mimeo.) Interruption of malaria transmission by chloro- 4. GIGLIOLI, G. Malarial Nephritis. Epidemiologi- quinised salt in Guyana. Bull. Wld. Hlth. Org. 36: al1 Notes on Malaria, Blackwater Fever, Albuminu- 283-301, 1967. ria and Nephritis in the Interior of British Guiana, 10. RAICHENOW, E. Zur frage der Malaria-resistenz Based on Seven Years Continual Observation. Lon- bei Negren. Behefte Z. Arch. J. Schi#s u. Tropen don, Churchill, 1930. 164 pp. Hyg. 33:58-64, 1929. 5. GIGLIOLI, 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 FOOD AND NUTRITION OF THE MAYA BEFORE THE CONQUEST AND AT THE PRESENT TIME'

Moisés Béhar

". . . and grinding then the yellow and the white ears In this paper I shall try to review our unfortu of corn, the goddess Ixmucané prepared nine beverages, nately' small store of information on the dietar- and from this food came strength and stoutness and with practices and nutritional status of the Mayar it they created the muscle and vigor of man. ... populations before the Spanish conquest of thi. "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 th( A great civilization developed in Middle main changes that have come to pass. It is m America on the basis of corn. The above quota- hope that, in so doing, some general principle tions from the Popol Vuh, the sacred book of applicable not only to this particular group o the ancient Quiché Maya, emphasize the im- American Indians but to others as well wil portance ascribed to this staple grain by the emerge. This seems to me more useful than: Mayan populations. Nevertheless, our present purely descriptive paper on the nutritional proL knowledge of nutrition has demonstrated that lems now afflicting the American Indian groups corn, if not properly supplemented (16), is Dietary practices and nutritional status of th nutritionally very poor as a staple. As in most of pre-Columbian Mayan Indians the cereal grains, its protein concentration is low; in addition, corn proteins have a very low Corn, prepared in many different ways, wa biological value. They are deficient in at least eaten at every meal, every day, as the most im two of the essential amino acids: lysine and portant food. In addition, the Mayas cultivatec tryptophane (5). Furthermore, the deficiency of and 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 egg' ment of pellagra in those whose diet is based were apparently not a regular item in thei on it (10). dietary; they were consumed sporadically an( It must therefore be asked how an important on special occasions, and were obtained mainl civilization like the Mayan could develop with by hunting. Very significant in this regard i such a great dependence on corn as its staple the existence, in a Mayan dialect of the Guate food; and why the present-day descendants of malan highlands, of a special word, tcibex, fc the Mayas have serious nutritional problems, "to eat meat," as distinct from the general wor also because of their dependence on corn. lobex meaning "to eat" (14). Only the group living on the coasts consumed fish. Milk was no 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 indicate

114 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 ents: protein, calcium, and niacin. The defi- would have to consume about 450 g of corn a 'ciency of calcium and niacin was corrected by day, or about 800 g of tortillas, to satisfy his ,he way in which corn was processed. For the protein needs. Not only is this physically im- Spreparation of tortillas and other corn products, possible but, in addition, the child would be the grain was first soaked overnight in lime receiving many more calories than he needs. 'water and then cooked in the same water (the Of course, the Mayas did consume other foods purpose was to remove the husk). The result besides corn. From what we know, the other was a product very rich in calcium, for which important items in the regular daily diet were ,he Mayas had no other adequate dietary beans, fruits and vegetables, and roots and lources. This method also yielded a higher con- tubers. Beans are a more concentrated source of centration of tryptophane, in relation to the 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, but The situation would again be one in which it is now evident that it was indispensable for adults 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 that other animal products tein. As I have said, the main drawback of corn (meat and 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 developed the required processing facilities. protein a day, which satisfies the average protein There was only one solution, and they applied requirement in an adult male weighing 65 it-prolonged lactation. The information now kilograms (17), does contain the needed available indicates, in fact, that Mayan mothers amounts of all essential amino acids (12). This breast-fed their children for at least three years amount of protein is provided by about 370 g and frequently even longer (15, p. 44). of corn, which is not an unreasonable quantity Our present knowledge indicates that breast 'tor a man to consume per day-in fact, 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, it is therefore mothers started to give their children what the possible for adults to satisfy their protein needs rest of the family ate but continued to feed them with this grain as the main staple, if they con- breast milk as an indispensable protein supple- sume enough of it. ment. By the time they were completely weaned, But the situation is 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 for tional requirements without the use of any essential amino acids is also greater. A two-year- special product, such as milk in the case of ")ld child, for instance, would need about 18 g Western civilization. It is also 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 as could be the main reason for the small size of the source, the child would need 36 g of protein, the adults, which is well documented. This

115 reduction in body size, a consequence of en- 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 ancestors. body size resulting in a similar reduction in The changes that have occurred, however, havel' nutrient requirements, the 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 information on Corn, processed in the same way, is still the the health conditions of children in those popu- I main staple. It provides up to 80 per cent of thq lations. Data on infant and early childhood calories and 70 per cent of the proteins in the mortality could be particularly illustrative. The diet of the adult population (9). Beans, vege- information available indicates that, in general, 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-Indian first two years. Later, their skin waxed dark with con- population are better able to com,! tinuous bathing by mothers and many suns; they were groups in the nonetheless bonny and mischievous throughout child- pete. This is even more true of animal products. hood, never ceasing to wander about carrying bows and Wild animals have practically disappeared, and arrows and frolicking among themselves and thus they hunting is no longer an important means of grew until they began to adopt the ways of the youths obtaining precious foods. Domestic animals are and to acquire a more exalted self-concept, and to leave of animal prod- behind the things of childhood. now the only important source ucts, and the market for them is very competi We cannot say the same about the majority of tive. The majority of the Indian families raise preschool children of Mayan descent today; they some animals in their homes, mainly poultry are usually apathetic and very often sick. and pigs. But these are more a source of badly Another important deduction that can be needed cash than of food. They and their prod- made from the information on late weaning ucts are consumed only on very rare occasions; concerns birth spacing. The mothers probably most of the time, they are sold. The meat most had at least three years between one birth and frequently consumed is beef, which has to be the next. To what extent prolonged lactation obtained from the market and therefore is one contributed to this situation we still do not of the luxury items for which cash is needed. know. However, it was definitely very con- Not only have the Indians lost their old, venient, because only in this way could the sources of food in the wild forests, but also the mothers breast-feed their children as required. amount of land available to them for cultivation Accurate information on the nutritional condi- has been drastically reduced. Before the Con- tion of pre-Columbian Mayan adults is also quest, they had all the land they needed to nonexistent, but all known facts indicate that, of although small, they were in general strong and produce their corn. Although the method healthy. We can thus conclude that, in terms of cultivation they used cannot be considered very nutrition, they did not have serious problems efficient today, its relative inadequacy was 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 nological development. They did, however, produce cash crops, which now compete with undergo acute crises, and there is information on food crops for the best lands. the famines they suffered when natural disasters Under present-day circumstances, the Indians (droughts or locust invasions) interfered with are frequently forced to buy even their corn, their corn crops. which they had always produced before. Owing

116 o their poverty, this greater dependence on the at the present time is barely adequate for adults narket to satisfy food needs has caused their and frequently inadequate for children. As a Jiet to deteriorate. Another change in the adult result, environmental stresses, particularly the Jiet is the introduction of coffee, which has heavy burden of frequent infections in early replaced the more nutritious corn and chocolate childhood, 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 severe cases of nutritional deficiencies found ;umption habits, with deleterious effects from particularly in early childhood. Briefly, the nu- the nutritional and even more from the eco- tritional problems now affecting the Guatemalan nomic standpoint. Indians are as follows: 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, the period of weaning cient and inadequate dietary intake and fre- kas been shortened. In pure Guatemalan In- quent infectious episodes, particularly diarrheal dians, prolonged lactation is still essential to processes and the common diseases of childhood 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 and early 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 the lence of severe cases of protein-calorie malnu- Conquest, is being progressively reduced as the trition (kwashiorkor and marasmus) (1), which process of integration into the ladino or Western are usually fatal if not properly treated in a culture advances (11). This situation is very hospital. Even more important from the public dangerous and has caused acute nutritional health standpoint are the chronic, subclinical problems, since the populations adopting earlier forms of malnutrition, which usually go unrec- weaning are not as a rule prepared culturally ognized. 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, cannot afford adults (20). If these observations are confirmed to feed milk to their children. -and studies are now under way to test the The situation is further aggravated by the hypothesis-it is easy to understand the tremen- introduction, by foreign civilizations, of refined dous importance of the problem 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 their participation in the social and economic and deleterious for the child during the post- development of their societies and couritries. ,weaning period. An 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 often used nowadays in gruels result of diets in which the sources of this vita- for small children, instead of the more nutritious min are vegetable products with low biological whole-corn gruels of old. activity consumed in insufficient amounts (8). The Guatemalan Indians of today thus seem This deficiency results in problems as serious as to be in a less favorable balance with their en- keratomalacia, which ends in total blindness 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 vitamin defi-

117 ciencies (8). It is closely interrelated with pro- the mother's breast for as long as possible. They tein deficiency, since the adequate dietary 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 of and very inadequate weaning foods. We must this deficiency are frequently seen. Its possible consider the situation of these groups of primi general effects are not clearly understood. tive Indians as they become incorporated intc Nutritional anemias are also highly prevalent; our societies, with the hope that we will find the deficiencies of iron and folic acid are the means to prevent or minimize the nutritional factors most frequently responsible, but again deterioration 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 pattern and the fortunately it has now been controlled by means nutritional condition of the Mayan populations of salt iodization (7). before the Conquest and at the present time The nutritional problems herein summarized indicates that, although corn constitutes the do not affect the Indian population in Guate- main staple for both population groups, the mala specifically or preferentially. They are asso- pre-Columbian Mayas were probably better ciated with low educational and economic levels adapted to ecological conditions. The availability and poor sanitary conditions. There are non- of game, greater availability of land for cultiva' Indian populations in similar conditions, but tion, the absence of competitive markets, pro- practically all the Indian populations fall into longed lactation of infants, and the lack of this category. Their difficulty is one of proper industrialized products were among the factors integration into and adaptation to new ecologi- that brought about a better diet for the Indians cal and cultural conditions and social structure. at that time. The sociocultural and economic In my opinion, the situation described in relation to the Mayan Indians, and the changes changes that have taken place since the Con- that have come to pass, are applicable to a lesser quest have worsened the diet of the present or greater extent to other Indian groups now Indians and resulted in serious nutritional prob- living in touch with Western civilization in the lems, among which protein deficiency-particu- Latin American countries. On the other hand, larly in small children-is the most important. many Indian groups still living in isolation from This experience should be considered both in Western culture and from the present Latin trying to solve the nutritional problems of this American societies have less serious nutritional and other American Indian groups now living problems; some of them seem to be living under with populations of Western culture and in pre- excellent conditions in this respect. These groups venting similar changes that could occur as the are still very dependent on hunting and fishing still primitive and isolated Indian groups are to satisfy their food needs, and still follow the incorporated into modern Latin American cul- "primitive" practice of raising their children at ture.

REFERENCES

1. AUTRET, M., and M. BÉíHAR. Síndrome poli- in four rural communities in Guatemala. Bull. Wld. carencial infantil (kwashiorkor) and its prevention Hlth. Org. 19:1093-1102, 1958. in Central America. Rome, Italy, Food and Agri- 3. BÉHAR, M., and N. S. SCRIMSHAW. Epidemi- culture Organization of the United Nations, 1954. ology of protein malnutrition. In lago Galdston (FAO Nutritional Studies, 13) (ed)., Human Nutrition Historic and Scientific. 2. BÉHAR, 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. (Institute of Social and Historical

118 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. lation of the amino-acid composition of proteins MITCHELL. The availability to pigs of nicotinic acid with their nutritive value. Nutrition Abstr. Rev. in tortilla baked from maize treated with lime- 16:249-278, 1946-1947. water. Brit. J. Nutrition 13:363-384, 1959. 5. BRESSANI, R., N. S. SCRIMSHAW, M. BÉHAR, and 14. LA FARGE, O. The Religion of a Cuchumatán F. VITERI. Supplementation of cereal proteins with Indian Town. Chicago, University of Chicago Press, amino acids. II. Effect of amino acid supplementa- 1947, p. 37. tion of corn-masa at intermediate levels of protein 15. LANDA, FRAY DIEGO DE. Relación de las cosas intake on the nitrogen retention of young children. de Yucatán (compuesto por el año 1560). 8a. ed. I. Nutrition 66:501-513, 1958. México, Editorial Porrua, 1959, pp. 36, 37. (Biblio- 6. CRAVIOTO, J. Application of newer knowledge teca Porrua, 13) of nutrition on physical and mental growth and 16. ORGANIZACIÓN DE LAS NACIONES UNIDAS PARA development. Am. I. Pub. Health 53:1803-1809, LA AGRICULTURA Y LA ALIMENTACIÓN. El maíz en 1963. la alimentación. Estudio sobre su valor nutritivo. 7. DE LEÓN, J. R., and 0. G. RETANA. La erradi- Roma, FAO, 1954. (Estudios sobre Nutrición, 9) cación 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. GARCíA, Z. FLORES, and M. Y. WHO. Rome, Food and Agriculture Organization LARA. Annual patterns of family and children's of the United Nations, 1965. (FAO Nutrition diet in three Guatemalan Indian communities. Brit. Meeting Report Series, 37; WHO Tech. Report I. Nutrition 18:281-293, 1964. Series, 301) 9. FLORES, M., and E. REH. Estudios de hábitos 18. SCRIMSHAW, N. S., and M. BÉHAR. Malnutri- dietéticos en poblaciones de Guatemala. I. Magda- tion in underdeveloped countries. New England 1. 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. BÉHAR. Protein 10. GOLDSMITH, G. A., J. GIBBENS, W. G. UNGLAUB, malnutrition in young children. Science 133:2039- and O. NEAL MILLER. Studies of niacin requirement 2047, 1961. in man. III. Comparative effects of diets containing 20. TORÚN, B., F. VITERI, F. SARAVIA, and M. lime-treated and untreated corn in the 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. GONZÁLEZ, M. L. SOLIEN DE. Breast-feeding, 21. VITERI, F., J. ALVARADO, D. G. LUTHRINGER, weaning and acculturation. 1. Pediat. 62:577-581, and R. P. WooD, II. Hematological changes in pro- 1963. tein-calorie malnutrition. Vitamins & Hormones. 12. HEGSTED, D. M. Variation in requirements of 26, 1968. In press.

119 IODINE DEFICIENCY WITHOUT GOITER IN ISOLATED YANOMAMA INDIANS: PRELIMINARY NOTE

René Riviere, Dominique Comar, Mentech Colonomos, Jean Desenne, and Marcel Roche 1

Roche et al. (10) showed that, in a region of endemic goiter, subjects without as well as those with goiter had an elevated thyroid radioactive iodine uptake. A similar finding was reported in New Guinea by Choufoer, Van Rhijn, Kas- senaar, and Querido (1). In an isolated primitive Venezuelan Indian community, Roche (9) demonstrated a generally elevated thyroid uptake without goiter; and this was again described in the eastern region of the Congo Republic by Delange, Thilly, and Ermans (3). In the present preliminary note, further quantitative aspects of thyroid metabolism in a Yanomama group from the upper Orinoco region are presented.

Material and methods ° The area chosen for study lies roughly be- 72- 70 tween the Ocamo and Mavaca rivers on the FIGURE 1. Area selected for study of thyroid metabolism Orinoco (Figure 1), in the extreme south of in Yanomama Indians (see page 77 for map of Vene- Venezuela. It is reached by plane to La Esme- zuela) ralda-an old mission site visited by Humboldt either not palpable or barely palpable. A (7) in 1800-and upstream from there by fast Catholic and a Protestant mission were estab- river boat in about four hours. Two expeditions lished at the site, but the Indians still lived in were carried out. wide-open thatch huts, ate their own food, and During a preliminary one in February 1962, went completely naked. thyroid iodine uptake was determined on 41 carried out in Janu- Yanomama Indians living in Bisaasiteri, at the Another expedition was and March 1968 to Bisaasiteri confluence of the Orinoco and the Mavaca. All ary, February, and to the neighboring the subjects studied were adequately nourished, Ocamo River (Tyowei- teri). The living conditions of the Indians had in good health, and with thyroids that were changed, although they remained very primi- 1 Presented by Dr. Roche. tive. They no longer lived in the characteristic

120 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 female-were studied. They water was 3.2 ,ug/liter (average of six samples). differed from those studied on the first expedition. They were in apparently good clinical nutritional state, of short stature (males 1.57 m Discussion and 52 kg, females 1.44 m and 45 kg, on the average) and without any clinical evidence of As Figure 2 shows, the thyroid iodine uptake thyroid dysfunction. In 45 of them the thyroid is higher in all the subjects studied in 1962. In was not palpable; in 10 it was palpable but 1968, although many uptakes are still high, this estimated to be of normal size. In one subject time five individuals have uptakes below 40 per (a male) there was a small diffuse goiter, esti- cent. It is presumed that the lowered uptakes are mated to weigh around 50 g. due to the introduction of foreign food and medications; none of the uptakes Twenty-four-hour thyroid uptake was deter- in the more mined after the oral or intravenous administra- isolated group were low, and the average in that group was higher than in the other two (Table tion of 5 to 30 ,tc of 13'I, in the manner recom- 1). mended by the International Atomic Energy Agency (12). A figure on the uptake of the first The urinary iodine/creatinine ratios (Table group has been published (6). Kinetic studies 1) are low compared to normal values (5). The are being continued after the oral administration total 24-hour iodine excretion can only be indi- of 100 [tc of 131I and will be reported later, as rectly evaluated. The values obtained are ap- will the effect of TSH and of methimazole and proximations, since urinary excretion of iodine other measurements. is not uniform throughout the day and since Radioactivity was measured on the site of the 24-hour urinary creatinine is not an accurately study; the various iodinated fractions of plasma fixed value. As a matter of fact, creatinine excre- were separated on ion exchange resins and mea- tion is related to muscle mass and in these short, sured separately. Measurements of urinary 127I thin Indians is likely to be less than is taken here as normal (4.) In a and creatinine were performed on single urinary group of Makiritare In- samples because of the practical impossibility of obtaining reliable 24-hour samples. 1963 5 1968 The blood, urine, and river drinking water samples were refrigerated around 40 C when indicated and sent by plane to the central labora- * lo tory in Caracas. Stable iodine was measured by activation analysis in the Nuclear Chemistry Department of the Venezuelan Institute for B1TTbi Scientific Research and at the Service Hospitalier Frédéric Joliot in France (2). The iodine/ creatinine ratio was measured to give an idea of the daily urinary iodine excretion. The average Thyroid Uptake (%) 24-hour creatinine excretion was assumed to be FIGURE 2. Frequency distribution of iodinel31 thyroid 1.6 g for men and 1.0 g for women (5, 8). uptake of Yanamama Indians

121 TABLE 1. Iodine metabolism in isolated Venezuelan Indians (average, standard deviation, range, and number of subjects)

24-HOUR PLASMA MAXIMUM 1251 1 2 7 THYROID 1311 IODINE (~g)/ THYROXINE PLASMA UPTAKE CREATININE (g) 1271 URINARY 127I THYROXIN SITE (% OF DOSE) RATIO (g/1 00 cc) (¡Lg/DAY) (%/LITER)

64.3 29.1 4.42 42.6 0.445 -7.4 -5.2 +0.53 ±7.8 +0.107 Bisaasiteri 36.0-81.3 10.0-104.0 2.8-5.7 15.0-166.0 0.194-1.100 (26) (35) (22) (35) (18) 61.1 0.679 -10.3 4-0.221 Tyoweiteri 48.6-70.0 0.207-1.450 (10) (10) 74.1 19.1 5.07 30.5 ±12.4 +±4.0 ±0.84 -6.4 Macorimateri 32 9 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 (11) found a 24-hour iodine excre- though, as is likely in this case, iodine deficiency tion of 21.2 /g/day with a mode between 10 and has been present through many generations. 20 /.g. lodine 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 /ag 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 the in 48 Indians from the Yanomama tribe living observation that these subjects are clinically 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/creatinine ratio, normal urinary 127I 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 be of goiter. It is 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.

RENCES

1. CHOUFOER, J. C., M. VAN RHIJN, A. A. M. 3. DELANGE, F., C. THILLY, and A. M. ERMANS. KASSENAAR, and A. QUERIDO. Endemic goiter in lodine deficiency, a primitive condition in the western New Guinea: iodine metabolism in goitrous development of endemic goiter. 1. Clin. Endocrinol. and nongoitrous subjects. 1. Clin. Endocrinol. and and Metab. 28:114-116, 1968. Metab. 23:1203-1217, 1963. 4. DE VENANZI, F., M. ROCHE, and A. GERARDI. 2. COMAR, D., R. RIVIERE, and C. KELLERSHOHN. Captación de yodo radiactivo (I131) por sujetos L'analyse par activation et ses applications aux eutiroideos de nuestro medio, y algunas considera- sciences biologiques. In Troisieme Congres Interna- ciones sobre aplicación al diagnóstico de las enfer- tional de Biologie de Saclay. Paris, Presses Univer- medades tiroideas. Acta Méd. Venez. 3:114-118, sitaires de France, 1964, pp. 307-336. 1955.

122 5. DWORIN, H. J., j. P. CHANDLER, and W. H. the absence of goiter in isolated Venezuelan Indians. BEIERWALTES. Estimation of fecal and urinary excre- I. Clin. Endocrinol. and Metab. 19:1440-1445, 1959. tion of iodine. 1. Clin. Endocrinol. and Metab. 25: 10. ROCHE, M., F. DE VENANZI, J. VERA, 149-151, 1965. E. COLL, M. SPINETTI-BERTI, J. MÉNDEZ MARTíNEZ, A. GER- 6. GAEDE, K. Algunos estudios recientes sobre el ARDI, and J. FORERO. Endemic goiter in Venezuela, bocio endémico. Acta Científica Venez. Supp. 1:58- 1 3 1 74, 1963. studied with I . I. Clin. Endocrinol. and Metab. 17:99-110, 1957. 7. HUMBOLDT, A. DE. Voyage aux régions équi- noxiales du Nouveau Continent. Paris, Librairie 11. ROCHE, M., H. PERINETTI, and A. BARBEITO. Grecque-Latine-Allemande, 1816. Urinary excretion of stable iodine in a small group 8. JOLIN, T., and F. ESCOBAR DEL REY. Evaluation of isolated Venezuelan Indians. I. Clin. Endocrinol. of iodine/creatinine ratios of casual samples as 21:1009-1012, 1961. indices of daily urinary iodine output during field 12. The Calibration and Standardization of Thy- studies. 1. Clin. Endocrinol. and Metab. 25:540-542, roid Radioiodine Uptake Measurements. Vienna, 1965. International Atomic Energy Agency, November 9. ROCHE, M. Elevated thyroidal pI31 uptake in 28-30, 1960.

123 STUDY OF ENDEMIC GOITER IN THE AMERICAN INDIAN'

José Barzelatto and Edmundo Covarrubias2

The existence of endemic goiter in America mented examples are those of the United States can be traced back to pre-Columbian times (5, (22), Mexico (40), Guatemala (39), Colombia 43), although the opposite view can be sustained (6), and Argentina (33). Iodine prophylaxis (14, 15). Endemic goiter areas have been programs are being increased in number and demonstrated during the last decades in practi- effectiveness in America and will soon influence cally every country in the Americas. From the goiter epidemiology, at least among Indians review published by Kelly and Snedden in 1960 more closely integrated with the surrounding (21), it can be estimated that at least 20 per society. cent of the population is affected by goiter. The idea that endemic goiter reflects only the From epidemiological surveys comprising in adaptation of man to iodine deficiency can no some cases huge samples, it can be inferred that longer be sustained. As in other parts of the all races are affected. Unfortunately, there is no world, it is contradicted in America by such precision about racial distribution in most of examples 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- and the persisting development of endemic cluded in variable proportions. Those Indians goiter in children born in areas where effective who live in primitive 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 (11, 23, 32, 34). An example in demic goiter areas in America, as judged by our continent is the piñon or nut of the Arauca- urinary excretion, has been found in practically ria 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, 41). This is not surprising, Contrasting with the situation in sporadic since except for the nitrate deposits in northern goiter, genetic mechanisms have not been well Chile (20) iodine is not abundant in the surface documented in the pathogenesis of endemic of this hemisphere as it is in the rest of the goiter. Enzymatic defects have not been demon- world. Wherever the diet has been supple- strated in its etiology. A concentration of en- mented with iodine, the incidence of endemic demic goiter cases within families has been goiter has been greatly reduced; well-docu- reported in America (16, 25) and elsewhere (7, 27). A positive association of nodular goiter 1 Work supported by Comisión de Ayuda a la Investi- with the distribution of nontasters of phenyl- gación Científica, Faculty of Medicine, University of Chile, Contract 64/4, and Comisión Nacional de Investi- thiocarbamide (PTC) has been reported in a gación Científica y Tecnológica, Project 41. general mixed population in Brazil (28). How- 2 Presented by Dr. Barzelatto. ever, the search for this association among

124 American Indians is rendered difficult by the Endemic goiter in Pedregoso small proportion of nontasters usually found In accordance with the foregoing line of 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 rTewenche 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 pbysiological characteristics of goiter may these ideas. vary in a single individual during his lifetime; Pedregoso is a geographically isolated Andean other levels of variation are to be defined among valley at an altitude of 1,310 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 live there, by relatively primitive ability make it a hard task to compare results agriculture and some cattle raising. Piñion is a of different authors, especially because there is standard item in their diet. The available still no general agreement on definitions. Some genetic data show that these Indians are less methodological limitations should also be borne mixed than other groups of the Araucanian in mind, such as 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 population in 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 retrosternal. comprising 81 per cent of the total population (the remaining 19 per cent, mainly young The impact of endemic goiter in public health people, were not examined in our house-to- is another matter of debate. Severe endemic house survey because for one reason or another goiter is associated with endemic cretinism, they were away from home). The distribution deaf-mutism and neurological defects, but the according to sex and age is shown in Table 1, 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 in with retrosternal extensions). As in most other individuals within an endemic goiter area. In studies, 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 life are 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- exceptional and only one cretin and no deaf- demic goiter. They presumably have very differ- mutes were found. ent 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 is are independent effects. Nodularity increases probably unique in each population; hence with age in both sexes, appearing earlier among studies in depth of endemic goiter in certain women (Table 2). Large goiters are more fre- communities may be scientifically rewarding, quently nodular than small ones with advance besides providing the basis for rational pro- in age, but some individuals have small nodular phylaxis programs. goiters in the first decades. It is obvious from

125 TABLE 1. Goiter in Pedregoso

MEN WOMEN

EXAMINED GOITROUS EXAMINED GOITROUS % AGE % % % % % (YEARS) N GOlTROUS N NODULAR LARGE N GOITROUS N 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 50 and over 29 76.9 22 68.2 63.6 28 96.4 27 77.8 81.5 Total 228 57.9 132 29.5 26.5 248 75.0 186 37.1 36.6

these data that if the influence of other etiologi- 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 should The data should therefore be analyzed both be considered separately. A number of facts by sex and by age group. In each of these cate- reinforce this conclusion: (1) The greater sever- gories, five subgroups have to be studied: with- ity of goiter among women, as judged here by out goiter and with small diffuse, large diffuse, greater frequency, larger size, and more nodu- small nodular, and large nodular goiters. Since larity, and also by the earlier appearance of these the size of the population and the distribution of characteristics, is due to poorly understood 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 life it is only important to working in nearby towns or farms, while compare subjects with and without goiter, ignor- women and children tend to stay on the reser- ing their sex. The second decade is a transitional vation. (3) Since most of the goiters become group, in which large and nodular goiters are nodular with age, nodularity among younger still infrequent and cannot be considered apart. individuals may have a different meaning from Among the adults there are relatively adequate that seen in most older subjects. (4) Estimates numbers of all groups except that of women of size among older individuals are liable to without goiter. Finally, among older individuals greater error, because of retrosternal extension, the meaning of these groups becomes less pre-

TABLE 2. Nodularity and goiter size in Pedregoso

MEN WOMEN

SMALL GOITERS LARGE GOITERS SMALL GOITERS LARGE GOIT ERS AGE

(YEARS) N g NODULAR N %ENODULAR N g NODULAR NE O NO[ )ULAR

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 6 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 cise, except for small diffuse goiters, which are those whose parents were born in different less subject to errors in classification. reservations. Other comparisons show no signifi- Let us first consider the effect of pregnancy cant differences. To sum up, it can be said that upon goiter variability in Pedregoso. In each inbred-people in Pedregoso develop goiters at a decade, women with either large or nodular later age and that their goiters are somewhat goiters have had a greater average number of less prone to develop nodules. pregnancies (Table 3)-which would suggest PTC-tasting ability was determined, with the that gestation increases goiter size and nodu- Harris and Kalmus technique, in 255 persons in larity if this interpretation were not challenged Pedregoso. Since only 4.3 per cent were non- by the fact that the same tendencies are ob- tasters, this characteristic could not be properly served among men. Men and women under 30 related to goiter. Nevertheless, a comparison years of age were not included in Table 3, made among tasters showed that both men and although they showed these same tendencies. women with nodular goiters in the third and The differences are significant only for men fourth decades of life had significantly lower with large as against those with small goiters in mean threshold values than those with diffuse the older group ("t"3,,)=2.337; p<0.05). It goiters (9). should be added that men without goiter aver- Height was also correlated with goiter (Table age a slightly higher number of descendants 5), and it was shown that taller subjects of both then goitrous men in these two age groups. sexes had larger goiters, in the third and fourth Inbreeding also seems to affect goiter in decade. An analysis of variance combining all Pedregoso (Table 4). The prevalence of goiter these groups showed that this relation is signifi- among children under 10 years whose parents cant at the .05 level (F(l,lo),=4.04). Nongoi- are consanguineous is significantly lower than trous men of the same age are in general taller among children of parents who were born in than goitrous ones, but this difference is not Pedregoso but in whom consanguinity was not significant even when the comparison is made detected (X(1) 2(yates)= 4.275; p<0.05) or chil- only with males with small goiters ("t"(5o)= dren of parents born on different reservations 1.857). (X(1)2(Yates)=4.275; p<0.05). The opposite Nutritional surveys made in Pedregoso re- trend seen among persons over 20 years of age vealed two relevant facts: first, the existence of is not significant. Inbred adults show a consist- iodine deficiency as judged by an average daily ent and significant tendency to have fewer nodu- urinary excretion of 33.2 micrograms (2); and 2 2 6 4 0 2 second, the fact that all year round the people lar goiters (X(1) (Yates)=6. ; p<0. 5) than consumed piñion (1), which has been demon- in rats (42). Goiter is TABLE 3. Fertility and goiter in Pedregoso (total off- strated to be goitrogenic spring, abortions excluded) apparently influenced by these two factors. Subjects with large or nodular goiters have 30-39 YEARS 40 YEARS OR OVER lower mean urinary iodide excretions than those with small or diffuse goiters (Table 6). These GOITER N X Se N X Se differences are better demonstrated by the per- Women centage of subjects in each group excreting 50 Diffuse 11 4.73 0.406 7 7.43 0.922 micrograms or more. With this approach, sta- Nodular 15 5.73 0.547 31 8.29 0.605 Small 18 4.89 0.427 9 7.33 1.414 tistical significance at the .05 level is reached Large 8 6.25 0.620 29 8.38 0.531 when large goiters are 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 9 5.00 0.408 13 7.00 1.104 direct relation between mean piñion ingestion Large 6 5.83 0.792 20 10.20 0.841 and thyroid size (Table 7), nongoitrous subjects

127 TABLE 4. Inbreeding and goiter in Pedregoso (men and women)

EXAMINED GOITROUS

AGE GROUP INBREEDING N %O GOITROUS N % NODULAR % LARGE

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

a Parents born in Pedregoso; consanguinity detected. b Parents born in Pedregoso, consanguinity not detected. e Parents born on different reservations. showing the lowest consumption. Since the comparable and slightly-above-normal figures values observed differ widely, it is preferable to for net iodine intake by the thyroid and periph- study in this case the proportion of subjects eral consumption of thyroid hormones; (2) a eating greater amounts, say 100 or more grams considerable "iodine leak," since the total out- daily; again the same trend is seen, the difference put of iodine by the thyroid is five to ten times being significant when subjects with large thy- the amount incorporated into thyroid hor- roids are compared with the rest (Xi)2(Yates)= mones; and (3) the abnormal presence in the 4.061; p<0.025). Diffuse and nodular goiters do blood of increased amounts of endogenously not differ by this approach. labeled triiodothyronine (T3), iodinated poly- Kinetic studies of iodine metabolism per- peptides (NBEI), and iodotyrosines. These formed in Pedregoso (3, 4) are summarized in facts, plus increased radioiodine uptake and Figure 1, the most striking findings being (1) PBI' 2 5 levels, express an accelerated turnover of

TABLE 5. Height (centimeters) and endemic goiter in Pedregoso

GOITROUS

NONGOITROUS DIFFUSE NODULAR SMALL LARGE AGE

(YEARS) N X Se N se N X se N X R Se N X Se

Men 20-29 14 162.1 1.85 17 159.8 1.22 6 160.0 2.81 18 159.4 1.41 5 161.5 1.00 30-39 9 164.2 2.41 9 160.8 1.18 8 163.1 1.47 11 160.7 1.02 6 164.2 1.67 40 and over 10 159.5 1.10 11 159.8 1.37 20 159.0 1.48 13 159.0 1.04 18 159.4 1.15 Women 20-29 2 150.0 2.50 32 150.2 0.90 10 150.5 1.86 23 148.4 1.07 19 152.5 1.01 30-39 - - 15 150.8 1.54 17 149.0 1.12 22 149.3 1.02 10 151.0 1.07 40 and over 2 150.0 2.50 9 151.9 1.76 30 149.8 0.86 9 150.3 1.88 30 150.3 0.85

128 TABLE 6. Urinary iodide and goiter in Pedregoso TABLE 7. Piñon ingestion and goiter in Pedregoso (micrograms per day) (grams per day)

50 ,g oR 100 GOR 2 N X S MORE (%) N X s2 MORE (%)

Nongoitrous 12 36.6 388.99 25.0 Nongoitrous 22 46.0 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 inefli- shows that nodular goiters have significantly ciency in handling iodine in terms of the per- higher values for this parameter, as is also true 125 centage used for the synthesis of thyroid hor- for NBEI . In a previous publication (4) we mones. Some of these findings have been found have shown that nodular goiters have the widest 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 largest figures 125 How should radioiodine studies be correlated observed. This finding may explain why PBI to goiter variation? The PBI125 reflects the speed values have the greatest variance among nodular of the thyroid in incorporating radioiodine into goiter, small pools determining the high values circulating iodinated compounds, and Table 8 observed. In support of this view, the thyroidal

Thyroid iodine pool = 11 mg.

I- < 97,6Ag/doy I + TYROSINE -* MIT-DIT ) MIT-DIT

- NBEI /

T3- T4 I I 102 )ug/day 1 1 1 T 3 - T4 I- +TYROSINE

Thyroid output = 512 - 833 - 974 jg/day

FIGURE 1. Iodine metabolism in Pedregoso

129 TABLE 8. PB1125 and NBEII2 5 in Pedregoso (o%/dose/ p<0.01), while there is no significant difference liter) between subjects with small and large goiters. 25 Nongoitrous persons have labeled PBI 1 NBEI 125 iodotyrosines in a smaller proportion than goitrous ones, 2 N X S2 N X S particularly in comparison to those with nodular Nongoitrous 10 0.17 0.0135 7 0.012 0.04176 goiters. Subjects with labeled iodotyrosines have significantly higher PBI'25 values ("t"(4 6)= Goitrous 0 5 Diffuse 30 0.18 0.0204 19 0.012 0.04823 2.31; p<0. ); this is not true for those with Nodular 34 0.50 0.1874 29 0.028 0.038736 labeled triiodothyronine. The demonstration of Small 30 0.31 0.1147 23 0.022 0.037575 iodotyrosines can be directly ascribed to small Large 34 0.39 0.1529 25 0.022 0.035058 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 1.97 mg+0.327, while five a p < 0.001. others in whom they were not demonstrated had b p < 0.010. an average value of 6.22 mg_1.757 ("t"( 4)= 2.37; p <0.05 ). iodine pool among 21 subjects in Pedregoso, calculated by the Riggs isotopic equilibrium Discussion method (4), shows a significant inverse rela- Multiple causes are involved in the etiology tionship with the PBI125 values observed in and pathogenesis of endemic goiter, many of these subjects (r= -4.72; p<0.02 5). A similar which still require proper identification and correlation has been observed in euthyroid pa- evaluation. Iodine deficiency is usually the pre- tients in Scotland (45). Furthermore, small dominant etiological factor; other environmental iodine pools could also explain high NBEI1 25 influences are probably also present and, where values, since this parameter shows a significant 25 there is adequate iodine, would sometimes positive correlation with PBI1 (r=0.50; p< suffice to explain endemic goiter. Natural goitro- 0.001). gens are the best-known example. Genetic fac- Endogenously labeled triiodothyronine is chro- tors have not been shown to influence the preva- matographically present in the same proportion lence of endemic goiter, although the possibility among all goitrous subjects and absent in the of their playing some role cannot be discarded. nongoitrous (Table 9), although the difference The participation of all these factors should 5 9 is not significant (X(1)2(yates)=2. 4 ). On the 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 a significantly higher percentage are seldom made, presumably because of the of the Indians with nodular goiters than of difficulties in definition and methodology that those with diffuse goiters (Xo()2(yates) =5.711; have been mentioned. It is upon variation that genetic factors have been demonstrated, as ex- TABLE 9. Endogenously labeled triiodothyronine and iodotyrosines in Pedregoso emplified by the association of PTC-tasting ability with nodular goiter. Weak environmental T IODOTYROSINES 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 Goitrous In Pedregoso we have looked into goiter varia- Diffuse 11 36.4 28 10.7 tion with three different elements in mind: Nodular 25 36.0 34 41.2 presence, size, and nodularity of 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 data a multi- larity is related to the two clearly defined genetic variate analysis through which the proper value aspects studied so far-inbreeding and PTC- could be attached both to main factors and to tasting ability-and to some radioiodine parame- the many kinds of interactions present. This ters. Inbred people develop fewer nodular goi- might not be possible, however, for two main ters; Indians with nodular goiters have lower reasons: the small size of our sample, especially thresholds for tasting PTC; and radioiodine since the subsamples in which parameters were 2 5 25 studies show high PBI1 and NBEI' values recorded did not always comprise the same indi- and a greater proportion of endogenously la- viduals, and the fact that the categories of goiter beled iodotyrosines in subjects with nodular contain different types of error in a nonrandom goiters. All these radioiodine findings can be way and have different meanings. For example, linked to the predominance of small thyroidal relationships with age are difficult to study be- iodine pools among those with nodular goiters. cause small diffuse goiters at an early age may Large goiters, on the other hand, are associated develop into large and/or nodular goiters. Our with the two best-defined environmental factors interpretation of the Pedregoso data requires us studied by us: the subjects eat less iodine and to extend these studies to other similar Pewenche more piñon. Size is also positively associated communities, but even then comparisons with with two complex factors-height and fertility. Presence, unlike the other two elements, has entirely different situations using similar types been connected significantly only with inbreed- of analysis would be necessary to determine the ing and during the first decade. It is noticeable bearing of such findings on the general problem here that early appearance of goiter, like nodu- of endemic goiter. Is endemic goiter the adapta- larity, is negatively influenced by inbreeding. tion of man to iodine deficiency? Our data 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. The fact be sought mainly through interaction types of that Indians with large goiters are taller and effects. have more children might be in line with such In view of the complexity and number of an interpretation, even if we are unable to in- factors to be considered acting together, it would terpret the mechanisms involved.

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131 15. GREENWALD, 1. Texas Rep. Biol. Med. 15:874, 32. PELTOLA, P. In C. Cassano and M. Andreoli 1957. (eds.), Current Topics in Thyroid Research. New 16. GREENWALD, 1. 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 GROOT, M. PAREDES, and 34. PODOVA, J., and P. LANGER. Naturally Occur- W. PEÑAFIEL. Rev. Ecuator. Med. Cienc. Biol. 5: ring Goitrogens and Thyroid Function. Bratislava, 15, 1967. Slovak Academy of Sciences, 1964. 18. FOLLIS, R. H., JR. Bol. Ofic. Sanit. Panamer. 35. ROCHE, M. 1. Clin. Endocr. Metab. 19:1440, 60:28, 1966. 1959. 19. FOLLIS, R. H. Am. 1. Clin. Nutr. 14:253, 1964. 36. ROCHE, M., F. DE VENANZI, J. VERA, E. COLL, 20. lodine Facts, I (217). London, lodine Educa- M. SPINETTI-BERTI, J. MENDEZ-FARTINES, A. GERARD, tional Bureau, 1940-1946. and J. FORERO. 1. Clin. Endocr. 17:99, 1957. 21. KELLY, F. C., and W. W. SNEDDEN. Bull. Wld. 37. ROCHE, M., H. PERINETTI, and A. BARBEITO. Hlth. Org. 44:27, 1960. 1. Clin. Endocr. 21:1009, 1961. KIMBALL, 22. O. P. Arch. Int. Med. 107:290, 1961. 38. ROTHHAMMER, F., R. BLANCO, E. COVARRU- 23. KRusIus, F. E., and P. PELTOLA. Acta Endocr. BIAS, and M. DIXON. Ztschr. f Morph. v Anthrop. u 53:342, 1966. In press. 24. LOBO, L. C. G., FRIDMAN, J. D. ROSENTHAL, 39. SCRIMSHAW, N. S. Public Health Rep. 75:731, R. ULYSSEA, and S. FRANCO. 1. Clin. Endocr. 22: 1960. 1182, 1962. 40. STACPOOLE, H. H. Bull. Wld. Hlth. Org. 9: 25. LOBO, L. C. G., A. QUELCE-SALGADO, D. ROSEN- 283, 1953. THAL, A. FREIRE-MAIA, and M. A. REBELLO. VI 41. STANBURY, J. B., G. L. BROWNELL, D. S. RIGGS, Pan American Congress of Endocrinology. Excerpta Medica, Internat. Congress Series 99, 1965. H. PERINETTI, J. ITOIZ, and E. B. DEL CASTILLO. Endemic Goiter: The Adaptation of Man to Iodine 26. MAISTERRENA, J. A., E. TOVAR, A. CANCIO, Deficiency. Cambridge, Massachusetts, Harvard Uni- and O. SERRANO. 1. Clin. Endocr. 24:166, 1964. versity Press, 1954. 27..MALAMOS, B., K. MIRAS, P. KOSTAMIS, J. 42. TÉLLEZ, M., A. GIANETTI, E. COVARRUBIAS, MANTZOS, A. C. KRALIOS, G. RIGOPOULOS, N. ZERE- FOS, and D. A. KOUTRAS. In C. Cassano and M. and J. BARZELATTO. 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 published. 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., and R. ETCHEVERRY. Nature 197: 44. WAHNER, H. W., E. GAYTAN, and P. CORREA. 187, 1963. 1. Clin. Endocr. 26:279, 1966. 30. NAGEL, R., R. ETCHEVERRY, C. GUZMÁN, A. 45. WAYNE, E. J., D. A. KOUTRAS, and W. D. HILLE, J. BARZELATTO, and E. COVARRUBIAS. Rev. ALEXANDER. Clinical Aspects of lodine Metabolism. Med. Chile 90:616, 1962. Philadelphia, F. A. Davis, 1964, p. 48. 31. PARRA-JIMÉNEZ, N., P. RODRíGUEZ-GARCíA, M. 46. WEINSTEIN, E. D., J. V. NEEL, and F. M. ROCHE, and K. GAEDE. J. Clin. Endocr. 22:754, 1962. SALZANO. Amer. I. Hum. Genet. 19:532, 1967.

132 DISCUSSION

J. D. Niswander

The speakers today have each considered one diabetes mellitus. The rank order for the hos- special disease problem of the Indian. I should pitals changes very little if, instead of the total like to discuss three of these in relation to one number of discharges, the total beneficiary popu- another. 1 will discuss diabetes mellitus, gall- lation of each hospital is used as the denomi- bladder disease, and nutrition in the context 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 reported an unusually tive frequency comparisons, although one cer- high frequency of diabetes among several North tainly would not want to attribute a great deal American tribes-with the studies being con- of precision to them. ducted among the Pima, reported by Dr. Miller, The map shows the location of the various being the foremost example. There are, how- Public Health Service Hospitals throughout the ever, very few data on the question of just how United States and their grouping into seven prevalent diabetes is throughout the various administrative areas. The eighth area (Portland) American Indian groups or whether it is more delivers service through contract with local fa- or less uniquely confined to particular tribes. cilities and not directly through Public Health Certainly such a hypothesis as that put forward Service Hospitals. by Dr. Neel-that diabetes may represent the The mean frequency for the 46 Indian hos- effects of changing dietary patterns on genomes pitals is 2.2 per cent. This compares with figures previously adapted to a "feast or famine" situa- of 1 to 1.5 per cent for similar data for several tion-would suggest that a high frequency of surveys of general non-Indian hospitals through- diabetes might be expected among populations out the United States representing over 26,000,- who have been both recently and rapidly trans- 000 hospital admissions. formed from this primitive pattern to 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 that I 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 ideal, is the records of the U.S. Public of 2.2 per cent is very wide, with roughly a Health Service, Division of Indian Health Hos- twentyfold difference between the lowest and pitals. Accordingly, I have obtained tabulations the highest. The table also shows the relative of discharge diagnoses from these hospitals for frequency of admission for gallbladder disease, the four-year period 1963-1967. This represents which very closely parallels that for diabetes. In approximately 250,000 discharges. I have tabu- fact, the rank correlation is slightly higher than lated the proportion of discharges from each .5 and extremely significant. There is, however, hospital in which the primary diagnosis was a considerably narrower range of variability than

133 .S.Aol Id.c R.ml$ S-.lcL Adi F-. - OAhAn~ 1. H. A.. C for diabetes. It can be seen that there is an various Alaskan Indians-and 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, by kan natives and the Navajo are probably the hospital, is highly significant. least acculturated of all the Indians remaining Specifically in regard to diabetes there are in the United States. several points 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 significantly greater proportion of of these groups, independent of environmental the patient population than in non-Indian hos- factors, cannot be excluded. Likewise, it could pitals. Most of the major linguistic groups are be argued that birth weight and diabetes are represented in the high and intermediate dia- related by virtue of both being the 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 me, 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 that To speculate a bit more than is perhaps war- these early agriculturists would be the ones most ranted by the data, I would suggest that virtually susceptible to acculturation and hence would all 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-diabetes group is notable for includ- tors associated with present-day reservation life. ing all of Alaska-the Aleuts, Eskimos, and In this regard, it is unfortunate that there are

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

# DIABETES * GALLBLADDER * M EAN MAJOR (% OF ALL (% OF ALL BIRTH HOSPITALS TRIBES ADMISSIONS) ADMISSIONS WEIGHT -1

Window Rock Area (6 hospitals-New Mexico, Navajo Arizona)

Belcourt, North Dakota Chippewa Mescalero, New Mexico; White River, Arizona Apache 0-1 1.9 6.68 Keams Canyon, Arizona Navajo, Hopi

Alaska (7 hospitals) Aleut, Eskimos "Alaskan Indians"

Montana (3 hospitals) Blackfoot, Crow Cheyenne, Gros Ventres Assiniboine, Cree Cass Lake, Minnesota; Red 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, Papago Shurz, Nevada; Owyhee, Nevada Paiute, Shoshone Winterhaven, California; Parker, Arizona Cocopa, Mohave, Chemehuevi

* Based on 257,320 discharge diagnoses, 1963-1967. 1IBased on 27,288 births.

135 no very adequate data available on relative of increased mortality and morbidity figures. obesity between tribes or on the increase in Difficult related questions, not yet answered, con- obesity that may have occurred in recent years. cern the proportion of genes potentially capable There is, however, one study that reports ap- of provoking 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. over a twenty-year period. The reservation All the specific health problems mentioned in Navajo, however, are still considered relatively this session are characterized by a complex lean in body build, without the great tendency causality involving multiple genetic, environ- toward obesity that is seen in some of the other mental, and cultural factors. The genetics of a tribes. It would be of considerable interest to disease like diabetes is a complicated subject, in follow Navajos who have now moved to the which no precision as to the part played by genes major urban centers and compare 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 who of disorders.1 A similar situation exists for have remained on reservations. endemic goiter. In the case of malaria, in which My purpose in this discussion has been pri- increased resistence is found among individuals marily to point up the need not only for detailed heterozygous for hemoglobin S, other still un- studies of special diseases in specific groups but known genetic factors, probably polygenic, may also for data giving comprehensive coverage 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 environmental causes. the interactions between genotype and changing Thus, the variability of all these diseases is 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, and no geneti- and between various human populations. cism or environmentalism will be of real value If my speculations prove to be correct, diabetes to an over-all comprehension of the problems.3 may become an increasingly important medical Among the specific questions raised to-day, I 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- relationship suggested for Pewenche Indians ning for dealing with the health problems of between nodular goiter and lower phenylthio- emerging primitive groups. carbamide thresholds in the group of "tasters" of this substance. Similar results may appear Acknowledgment from the studies of diabetes and gallbladder I am grateful to Mr. Abraham Ranofsky, Program disease among the Pima Indians, in relation to Anal\ysis and Statistics Branch, Division of Inclian Health, blood groups. Whether or not such associations for furnishing the special tabulations on diabetes and galibla lder disease. are real is to be found in the future, but they may constitute starting points for the analysis of polygenic systems in man. Another interesting Edmundo Covarrubias question here concerns how the epidemiology of The presentations we have heard this afternoon make it clear that the American Indians 'Rimoin, D. L. Genetics of diabetes mellitus. Diabetes 16:346-351, 1967. received from the conquisatadors a rather heavy 2 Allison, A. C. Genetic factors in resistance to malaria. burden of evils: bio'ogical agents of disease and Ann. N.Y. Acad. Sci. 91:710-729, 1961. the pressure of a different culture, stresses to 3 Thoday, J. M. Geneticism and environmentalism. In J. E. Mead (ed.), Biological Aspects of Social problems. which they had to become adapted at 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 of and that if 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 to be learned, I of diagnosis due to cultural differences. One believe, from different groups of people and effort in this direction was recently made in different traditional systems. I think that we in Chile: a new structural theory of psychoses led the so-called Western world have a considerable to transcultural definitions that made it possible body of dogma in our own infant feeding that to distinguish considerable 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. Béhar pointed out, in most of pre-conquis- Finally, I feel tempted to generalize to other tador South America and also in Polynesia 5 systems the words of Book concerning the animal milk was not traditionally drunk. Yet etiology of mental illness: ". .. my general people there reared children and fed them per- medical and biological background together with fectly adequately without it. There are messages my fragmentary knowledge of psychiatric re- for us from these different groups. search work has led to the tentative conclusion Another important aspect of Dr. Behar's paper that any unitary explanation of the etiology is the fact that disruption of a culture can very even of what is now considered a special type frequently have its most sensitive and devas- of mental illness, or a clinical entity, will not tating effect on the age group that is most pre- contribute to the advancement of psychiatric cariously poised physiologically. That, of course, research." is the young child-the weanling, who is in a To sum up, biomedical research among the process, if you like, of biocultural transition. American Indians is apt to disclose new facts When we consider the Indian communities in that may be of value both for specific public the 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 Genera! Discussion were found to have a pattern of infant feeding well adapted to their community-breast feed- Jelliffe: I should like to congratulate Dr. Béhar ing, bone marrow, pre-chewed meat, and so on his excellent paper. It seems to me that, late forth. Five years later another survey was car- as the hour is, it would be wrong to let this go ried out, and the pattern of infant feeding and by without emphasizing its importance. As those childhood malnutrition had changed completely. of us here who are working in the field of child- In the first survey there was very little protein- hood malnutrition will agree, the lessons to be learned from his paper are of great importance. calorie malnutrition. In the second it was very First, I think that we all have to realize that common. all groups that have been living in a certain What practical messages are there to be gained from Dr. Béhar's paper? I think there are these: 4Mufioz, L., J. Marconi, J. Horwitz, and P. Naveil- We should realize that as well as the rapid lán. Cross-cultural definitions applied to the study of spread of infections due to viruses, such as functional psychoses in Chilean Mapuches. Brit. 1. Psy- chiat. 112:1205-1215, 1966. measles, or to bacteria, such as tuberculosis, the Boo;k, J. A. Genetical etiology in mental illness. In spread of inappropriate non-adapted ideas con- Causes of Mental Disorders: A Review oj Epidemio- cerning infant feeding can have a very consid- logical Knowledge. New York, Milbank Memorial Fund, 1961. erable effect on local child health and on the

137 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 groups-in too easily in the dietary habits of American northeastern India, for example. But it is so Indian groups, careful thought should be given localized as to suggest a racial factor in the to who 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 Waterlow: 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 h4 I think, therefore, that in dealing with the Amerindian. American Indian groups that have been dis- Giglioli: 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 Indians. These are people who are Furthermore, any imports of foods into these at our mercy both morally and physically. The communities through, for instance, 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. This 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 I believe, 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. This 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 measles restraint in his investigation to be sure that his or tuberculosis. activity is in no way harmful to the populations Waterlow: May I ask Dr. Giglioli one ques- he is studying. tion? Is it 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 whether others? we are not dealing with the well-known modula- Giglioli: My paper touches on this question. tion effects that derepressors and repressors Not that I have any competence to discuss it. might have on the transcription and translation I mean, it is a moot question whether malaria phenomena. It is now, I think, reasonably well pre-dated the Conquest or was introduced. I documented that the influence of a variety of think the consensus now is that it 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 Amerindian 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 for ages. are now being explained in molecular biological We have no data to go by. Consider the Medi- terms of messenger RNA and all that goes with terranean populations, which were exposed to it. This must mean that there is an interplay by malaria for centuries. Certainly there was no a 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 elicit a qualitative changes. great pattern of enzymatic changes that are not As I listened to many of these presentations, due to an instantaneous transition of the genes. diabetes in Indians and so on, I thought it quite We know this. Just a simple high carbohydrate conceivable that this could be a quantitative intake as against a low, high nitrogen intake variation of one or another enzyme pattern that as against low, will cause changes of level of would lead to a metabolic anomaly of the kind I high orders of magnitude. These can 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 question still remains 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 before we get too Moderator: A discussion of this would take us involved in genetic markers as an absolute rather far afield. I am not sure whether you are known relationship, which is clearly the case in using the term modulation in the sense used by many diseases that have been defined in these the bacterial geneticist or whether you are re- terms, we give some thought to the possibility ferring to the kind of nature-nurture inter- that in many of these adaptive phenomena we actions for which we have come to have great are dealing with something quite different. respect in 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 the en- up to our discussion today. This is where the vironmental difference is 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 Moderator: The final item on the agenda is quite a bit. On the other hand, perhaps we labeled "Commentary and Summarization." 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, and cer- of a good meeting is the extent to which old tainly one opportunity is in transitional popula- ideas are rescrutinized and questioned, and tions. For instance, if it can be documented that from beginning to end of this session we have the polymorphisms, which are a readout for been questioning some of our old ideas, begin- proteins, alter in frequency more rapidly in a ning with the matter of the origins of the population in cultural transition than can pos- American Indians and ending with details of sibly be ascribed to genetic selection, this would disease susceptibility and resistance. The session be hard evidence that the readout is different. has been very profitable, and it is my privilege That is the kind of evidence that I think it to thank PAHO for making this gathering would take to satisfy most of us. possible, both financially and in terms of facili- Cohen: Not to prolong this, it is very clear ties.

139 (1 Appendix

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. Schull

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, it is 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 scat- an unusual challenge to those interested in tered; it seemed that a compilation of those recent developments in genetic taxonomy (see studies which deal with the major genetic poly- 1, 2), but attempts to apply these developments morphisms would prove useful in assisting in- may be vitiated by substantial racial admixture. vestigators both in planning and facilitating Accordingly, 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 1 to 11 probably essentially no admixture, (2) non- 1 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. tribution to the gene pool probably greater than These are all systems exhibiting marked poly- 5 per cent. morphism. We have not included systems, such As every student of the problem knows, the as the glucose-6-phosphate dehydrogenase or classification of certain samples into one of these hemoglobin system, that are polymorphic in three categories is difficult, and we are sure not some populations but essentially nonvariant in only that there are grounds for issue with some the Indian. Nor have we included some systems of our decisions, but also that some of these that while probably polymorphic are still inade- decisions are incorrect 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 results of the genetic studies themselves, and, 1Work supported in part by U.S. Atomic Energy Commission Contract AT(11-1)-1952. where given, the author's statement.

141 The results of typings for the ABO system tribal organization has long since disappeared, have been thought to provide especially critical these names may be confusing and occasionally material for decisions regarding admixture. We meaningless. Thus there are two samples of accept the working hypothesis that IB and IA2 Mayans so called by the authors, an unsatisfac- are not found among American Indians and tory name since several other samples are tabu- that IA1 does not occur in South America. How- lated of Mayan-speaking Indians-Chol, Kek- ever, we note that in the few South American chi, Mam, Pocomam, Quiché, and others. Five samples where IR and I4 do occur, the ratio of samples are designated Aymara, two Quechua, the frequency of the former to that of the latter and one Chibchan, each of which is a large tends to be higher than in western Europe and linguistic stock including several distinct cultures Africa, the presumed origin of these genes. The and several millions of persons. Three linguistic possibility must be recognized that IB was pres- stocks of simpler culture are similarly employed ent in South America in pre-Colombian times as names of samples-Arawak, Carib, and (see also 6), perhaps introduced by trans-Pacific Guaraní. From four to ten samples are desig- contacts. nated by tribal names that fall under each of Additional genetic grounds for decisions re- these stocks. garding sample purity are the presence of the The estimates of gene frequencies in the following antigens, wherever tested for: Kell, tables that follow were computed on an IBM Lewis (a), Berrian (Be), Henshaw (He), 1130 Model, using programs written in the De- Lutheran (Lua), Miltenberger (Mia), Sutter partment of Human Genetics. The ABO pro- (Jsa), V of the Rh system, Verweyst (Vw) of gram has been published (4); the other pro- the MN system, and Wright (Wr). All of these grams are available on request. The "goodness are thought to be absent in unmixed Indians, of fit" of the observed phenotype frequencies and we have not included tabulations for any with those expected under Hardy-Weinberg of them. References establishing the virtual or equilibrium and with the calculated gene fre- complete absence of these traits in samples of quencies has been tested by chi square. The latter unmixed Indians are listed in Table 12. We are not given, but the corresponding probabili- have felt, on the other hand, and contrary to ties when significant are indicated by one 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 frequencies 0.01>P>0.001; C-0.001>P. The validity of may be of significance. applying such x2 tests to small samples in which In our opinion, many types of critical com- the mean coefficient of relationship is probably parisons are impossible when there is more 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 incorporated letters and followed by a brief indication of into Tables 1 to 11, with samples of the first locality, such as that of a state or province, class designated by asterisks. The references to River (R), Mountain (M), Lake (L), or region samples with more than 5 per cent admixture (N for north, and so on). Exceptions are made are given in Table 13, which lists the samples for French Guiana and for Guyana and Suri- in alphabetic order. nam (formerly British and Dutch Guiana, re- The names of tribes and groups are those spectively), for which no locality is added, since given by authors, to which an arabic numeral their areas are small. has been added in cases 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, where pooled data of the "tribe." The grounds for

142 recognizing subtribes are seldom clear; they a 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 cases, we have retained cent non-Indian ancestry. Such data are not the author's designation but listed his data under tabulated here, but the references are listed the main tribal unit, with a cross-reference apart (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 under ity, 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 been published on

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150 243. SANTIANA, A. Los indios del Ecuador y sus Javanese, Djuka Negroes and Oyama and Carib In- características serológicas. Terapía 10:13-18, 1952. dians. Nature 185:324-325, 1960. 244. SCARO, J. L. Investigación del factor Diego 258. SUÁREZ MORALES, O., B. BRODER, and A. -en aborígines de la Quebrada de Humahuaca. Rev. LOAYZA. Antígenos de grupos sanguíneos en indios Soc. Argent. Biol. 34:71-74, 1958. chipayas de Bolivia. Anal. Acad. Nac. Cien. quad. 2, 245. SCOTT, E. M., I. W. DUNCAN, V. EKSTRAND, Cien. Nat. (La Paz) pub. 4, pp. 1-9, 1964. and R. C. WRIGHT. Frequency of polymorphic types 259. SUÁREZ MORALES, O., E. LINARES I., and of red cell enzymes and serum factors in Alaskan J. GUTIÉRREZ B. Grupos sanguíneos en poblaciones Indians and Eskimos. Amer. 1. Hum. Genet. 18: del altiplano andino con bocio endémico. Anal. 408-411, 1966. Acad. Nac. Cien. quad. 2, Cien. Nat. (La Paz) f 246. SHIM, B. S., and A. G. BEARN. The distribu- pub. 5, pp. 1-10, 1965. tion of haptoglobin subtypes in various populations, 260. SUTTON, H. E., G. A. MATSON, A. R. ROBIN- including subtype patterns in some nonhuman pri- SON, and A. W. KoUCKY. Distribution of haptoglo- mates. Amer. J. Hum. Genet. 16:477-483, 1964. bin, transferrin and hemoglobin types among In- 247. SHREFFLER, D. C., and A. G. STEINBERG. dians of southern Mexico and Guatemala. Amer. Serum protein groups and the SC trait of saliva in 1. 1 Hum. Genet. 12:338-347, the Simóes Lopes and Sao Marcos Xavantes. Amer. 1960. 1. Hum. Genet. 19:514-523, 1967. 261. SUTTON, H. E., J. V. NEEL, F. B. LIVING- STONE, G. BINSON, P. KUNSTADTER, SILVA, DA, see DA SILVA. and L. E. TROM- SILVA, R. M., see MARTíNEZ SILVA, R. BLEY. The frequencies of haptoglobins in five popu- 248. SILVER, R. T., J. M. HABER, and A. KELLNER. lations. Ann. Hum. Genet. 23:175-183, 1959. Evidence for a new allele in the Kidd blood group 262. TEJADA, C., M. SANCHEZ, M. A. GUZMAN, E. system in Indians of northern Mato Grosso, Brazil. BREGNI, and N. S. SCRIMSHAW. Distribution of blood Nature 186:481, 1960. antigens among Guatemalan Indians. Hum. Biol. 249. SILVER, R. T., J. M. HABER, and A. KELLNER. 33:319-334, 1961. Blood studies of jungle Indians of the upper Xingu 263. TEJADA, C., M. SANCHEZ, M. GUZMAN, N. S. River, Mato Grosso, Brazil. Bibl. Haemat. 13:167- SCRIMSHAW, and E. BREGNI. Blood groups in Mayan 169, 1962. population. Abstract. Rec. Genet. Soc. Amer. 28:102, 250. SNYDER, L. H. Human blood groups: their 1959. inheritance and racial significance. Amer. 1. Phys. 264. THOMAS, J. W., M. A. STUKEY, H. S. ROBIN- Anthropol. 9:233-263, 1926. SON, J. P. GOFTON, D. O. ANDERSON, and J. N. BELL. 251. SOCARRAS DEL Ríio, J. A. Contribución al Blood groups of the Haida Indians. Amer. 1. Phys. estudio de los grupos sanguíneos en el Perú. Rev. Anthropol. 22:189-192, 1964. Cienc. (Lima) 32, 1930. Cited in Sacchetti, 1953. 265. URIZAR, R. Grupos sanguíneos de autóctonos 252. SOCARRAS DEL Río, J. A. Contribución al del Chaco Paraguayo. Amer. Indígena 2:49-50, 1942. estudio de los grupos sanguíneos en Colombia. Univ. 266. VELA, C. Los grupos sanguíneos en Quito. Nac. Col. (Bogotá), 1930. Cited in Sachetti, 1953. Quito, Imp. Univ. Central, 41, 1933. 253. SORIANO-LLERAS, A., and R. MARTíNEZ SILVA. 267. VELLARD, J. Contribution á l'étude des in- Estudios immunohematológicos entre los indios diens Uru ou Kot'suns. Trav. Étud. And. 1:191-209, lloreos. Rev. Fac. Med. (Bogotá) 28 (7-9):101-106, 1949. Abs. HLAS 17:33. , 1960. VIANNA MARTINS. See MARTINS. 254. STEFFAN, P., and S. WELLISCH. Die geogra- phische Verteilung der blutgruppen. Z. Rassen- 268. WALKER, M., F. H. ALLEN, and M. T. NEW- physiol. 5:180-5, 1932; 6:28-36, 1933; 7:176-185, MAN. Blood groups of Cakchiquel Indians from 1935; 8:38-47, 1936. Sumpango, Guatemala. Amer. J. Phys. Anthropol. 25:239-242, 1966. 255. STEINBERG, A. G., M. S. CORDOVA, and R. Lls- KER. Studies on several genetic hematological traits 269. WEISS, P. Estudio sobre los lamistas; su grupo e, of Mexicans. XV. The Gm allotypes of some Indian sanguíneo; algunas pruebas psico-técnicas. Rev. Mus. tribes. Amer. 1. Hum. Genet. 19:747-756. Nac. (Lima) 18:19-41, 1948. Abs. HLAS 15:48. 256. STEINBERG, A. G., R. STAUFFER, B. S. BLUM- 270. WIENER, A. S., J. ZEPEDA, E. B. SONN, and BERG, and H. FUDENBERG. Gm phenotypes and geno- H. R. POLIVKA. Individual blood differences in Mexi- types in U.S. whites and Negroes; in American In- can Indians, with special reference to the Rh types dians and Eskimos; in Africans; and in Microne- and Hr factor. 1. Exp. Med. 81:559-571, 1945. sians. Amer. 1. Hum. Genet. 13:205-213, 1961. 271. WITKOP, C., and J. GAISER. Genetic blood ·257. STEINBERG, A. G., R. STAUFFER, and H. FUD- studies. Bethesda, Maryland, NIH, Interdepart. Com- a ENBERG. Distribution of Gm and Gm-like among tee. Nutrit. Nat Defense, 1960.

151 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 several areas or sub- British Honduras BRH tribes, in which case "P" appears in column 14 preceded w French Guiana FRG by the serial number in column 13. (Subsamples are Guyana GUY presented separately, also, whenever available.) "P" Surinam SUR stands for "pooled."

Abbreviations of nations and colonies and Other abbreviations other abbreviations North N East E Nations South; Saint; Argentina ARG Sierra S Bolivia BOL West W Brazil BRZ Mountain (s) M(TS) Canada CAN Another; Others ANO Chile CHL River R Colombia COL Lake L Costa Rica COS Delta D Ecuador ECD Island(s) I(S) Guatemala GUA Pooled data, Honduras HON from several Nicaragua NIC samples P

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180 TABLE 12. Data on genes that are probably absent in Indians who have no Caucasian or Negro admixture. Tabulation restricted to samples in which non-Indian ancestry is believed to be less than 5 per cent, as listed in Table 1. Samples with relatively high frequencies that may be due to technical problems are starred (*).

Lutheran

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

Tribe or Group Area Reference Sample Size Lu(a+) Aymara Bolivia: Yungas 169 120 1 Cuna Panama: San Blas Is. 178 388 2 Diegueño U.S.A.: California 202 58 2 Kalapalo (2) (*) Brazil: Mato Grosso 201 73 12 639 * Among six samples tested with both anti-Lua and anti-Lub, no case of either Lu(a+) nor of Lu(b-) was revealed (References 33, 176, 177 and 178).

Kell

Among. 40 samples tested with anti-K serum alone, 28 contained no positives, and may be considered to consist wholly of homozygous kk cases. They are reported in seventeen references: 42, 64, 74, 81, 90, 114, 132, 134, 145, 171, 175, 195, 223, 231, 262, 263, and 268. The 12 exceptions are:

Tribe or Group Area Re/erence Sample Size K+ Athabascan U.S.A.: Alaska 3 206 1 · Aweikoma Brazil: Sta. Catarina 233 122 3 Carajá Brazil: Sta. Isabel 108 35 7 Chol Mexico: Chiapas 64 137 4 Cara Mexico: Nayarit 64 96 6 Diegueño U.S.A.: California 202 56 6 Huastec Mexico: Vera Cruz 64 75 2 Huichol Mexico: Jalisco 64 70 2 Kalapalo (2) (*) Brazil: Mato Grosso 201 73 17 Maztec (1) Mexico: Oaxaca 64 136 1 Y 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 contained no case of Kell positive and may be considered to consist of homozygous kk individuals only. They are 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 and 262. The 4 exceptions are:

9 Tribe or Group Area Re/erence Sample Size K+ . Cherokee U.S.A.: No. Carolina 209 78 1 Chippewa (*) U.S.A.: Minnesota 159 161 24 Goajiro Venezuela: W. Zulia 128 119 1 Navajo (4) U.S.A.: Arizona 63 237 4 595 30

All of the 30 K-positivc cases were heterozygous Kk except two of the four Navajos, who were KK.

Wright

Among 41 samples tested with anti-Wra serum, not one contained a positively reacting case. References: 63, 91, 164, 167, 168, 169, 171, 173, 175, 176, 177, 178, 209, 231, and 233.

181 TABLE 12-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. The other 39 samples are reported in references 164, 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 Cobán, Guatemala (ref. 173), one individual was Vw-+. The other references are 91, 164, 167, 168, 169, 171, 175, 176, 177, 178, and 209.

"V" (in Rh-hr system)

Among 28 samples tested with anti-V serum, only two samples contained positive cases. Among 12 Kekchis tested at Cobán, Guatemala (ref. 173), two were V+, and among 45 Totonacs (sample 2) at Vera Cruz, Mexico (ref. 170), one was V+. The other samples are referred to in 28, 167, 170, 171, 175, 176, 177, 178, and 209.

Berriens

Among 18 samples tested with anti-Bea serum not one positive case was found. References are: 167, 171, 173, 176, 177, 178, and 209.

Sutter

Among 15 samples tested with anti-Jsn serum, only one contained a positive case-one individual in a sample of 119 Goajiros, sample no. 3, in western Zulia State in Venezuela (ref. 128). Other references are: 28, 126, 167, 171, 173, 176, 177, and 209.

182 TABLE 13. Tribes and groups estimated to have more than 5 per cent of non-Indian ancestry. Some tribes list-d here will be found also among one or another of tables 1-11. Such duplication results from two or more samples having been drawn from the same tribe, whether by the same or by different workers, one or more samples indicating less than 5 per cent non-Indian ancestry and one or more indicating more than 5 per cent non-Indian admixture.

TRIBE OR GROUP COUNTRY REFERENCES TRIBE OR GROUP COUNTRY REFERENCES

Alukuyana Surinam 109 Chorotega Nicaragua 165, 175 Andido Argentina 205 Chorotí 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, 196,211,237 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 Aymará Bolivia; Peru 223; 225 Emerillon French Guiana 114

Beaver U.S.A.; Flathead 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 Guaraní Brazil 216 Boruca Costa Rica 177 Guatuso Costa Rica 81 Botocudo Brazil 153 Guayquerí Venezuela 134, 135 Cáchama Venezuela 118, 119 Haida Canada 102,150,264 ' Caddo U.S.A. 99 Hopi U.S.A. 42,250 Caingang Brazil 76,77 Huichol Mexico 192 Canela Brazil 22,65,146 Hupa U.S.A. 104 Carajá Brazil 106,107 Ica Colombia 22, 134 Caramanta Colombia 11 Itonama Bolivia 169 Carib Surinam 257 Itza Maya Mexico 170,260 Carrier Canada 98 Catawba U.S.A. 210 Jicaque Honduras 165, 174 Catio Colombia 12 Kansas U.S.A. 112 Catopaxi Ecuador 242 Kariri Brazil 207 Ceará 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 217 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 Chipaya Bolivia 258 Lamistas Peru 269 Chipewyan U.S.A. 96,98, 143 Lenca Honduras 165, I174 Chippewa U.S.A. 52,56, 159 Llorcos Colombia 253 Chiriguano Argentina 181,254 Choctaw U.S.A. 57,112,250 Makah U.S.A. 102 Chol Guatemala; Mapuche Chile 74, 101, 184, 188, 189, Mexico 64, 170, 171,260, 262 196, 211, 238,239 Chontol Mexico 64 Mataco Argentina 7, 182,254

183 TABLE 13-Continued

TRIBE OR GROUP 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 Salish U.S.A.; Canada 62, 87, 89, Mixe Mexico 64, 165 102, 141 Mixtec Mexico 64,221 Sapelo Canada 219 Mojave U.S.A. 42 Sarcee Canada 50 Montagnais Canada 33,219 Sekani Canada 98 Moré Bolivia 169 Senaca U.S.A. 70 Muckleshoot Canada 102 Seri and Yaqui Mexico 192,221 Muskhogean U.S.A. 112,250 Shoshone U.S.A. 250 1 Sibundoy Colombia 198 ,i 221 Nahua Mexico Sioux U.S.A. 154, 157,250 Naskapi Canada 33,183,219 Slave Canada 87, 97, 98, 156 Navajo U.S.A. 4,6,40,42, 63, Stoney Canada 50, 97, 98, 155 ' 191,203 Subtiaba Nicaragua 165,175 80 Nez Perce U.S.A. Swinomish U.S.A.; Canada 102,103 Nootka Canada 102 Tacana Bolivia 169 187 Oajana Surinam Tarahumara Mexico 187,221 Okanagon U.S.A.; Tarasco Mexico 24,221 Canada 102,103 Tariana Brazil 31 157 Omaha U.S.A. Tewa (Pueblo) U.S.A. 4 211 Ona Chile Tinneh U.S.A. 206 Otomí Mexico 187,226, 229 Tlaxcalteca Mexico 227 Oyama Surinam 257 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 Pataxó Brazil 208 Trumai Brazil 249 Paya Honduras 165, 174 Tsimshian Canada 102,217 Pawnee U.S.A. 99 Tulalip Canada 102 Pehuelche Chile 189,254 Tungurahua Ecuador 242 Pehuenche Chile 74,254 Tuscarora U.S.A. 185 Penobscot U.S.A. 3 Uru Peru 267 v 155 Piegan Canada Ute U.S.A. 161, 163 Pijáo Colombia 213 99 Pilagá Argentina 182,199,254 Wichita U.S.A. 157 Pima U.S.A. 42, 100 Winnebago U.S.A. 104 Ponca U.S.A. 250 Wintun U.S.A. Pueblo New Mexico 4,5,6,39,40 Yagan Chile 75 Puruhá Ecuador 242 Yakima U.S.A.; Canada 102, 103 . Yamana Chile 74 Quechua Ecuador; 3,28,93, 167, 168, Yaqui and Seri Mexico 192,221 Peru 214,223,244 Yurok U.S.A. 104 Quillacinga Colombia 198 Quilleute U.S.A.; Canada 102 Zapotec Mexico 64, 170, 187, 260

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