INT J TUBERC LUNG DIS 4(5):395400 2000 IUATLD

UNRESOLVED ISSUES

The origins and precolonial epidemiology of tuberculosis in the Americas: can we gure them out?
T. M. Daniel
Center for International Health, Case Western Reserve University School of Medicine, Cleveland, Ohio, USA SUMMARY

Paleologic evidence of tuberculosis in the precolonial Americas is reviewed to cast light on its origins and subsequent epidemiology. The genus Mycobacterium is an ancient one, and M. tuberculosis may have differentiated 20 400 to 15 300 years ago. The Americas were peopled by migrants from Asia in two major migrations, one occurring more than 20 000 years ago and the other 12 000 to 11 000 years ago. Tuberculosis reached the Americas with these migrants, persisting at a low level of

endemnicity in small, dispersed population groups. Beginning about 1500 years ago, an epidemic of tuberculosis began, probably in the Andean region of South America. It did not reach or subsided in time to leave highly susceptible indigenous American populations at the time of European colonization. K E Y W O R D S : tuberculosis; history of tuberculosis; precolonial America

THAT MYCOBACTERIAL INFECTION occurred in the Americas in the precolonial epoch has now been rmly established by the identication of mycobacterial DNA in a pulmonary lesion typical of tuberculosis in a 1000-year-old Peruvian mummy by Salo and associates.1 This demonstration would seem to have put an end to the discussion and speculation on whether tuberculosis existed in indigenous native Americans prior to European colonization, a subject that had been the source of considerable controversy, as summarized by the skeptical reviews of Morse2,3 and of Paulsen.4 That the matter is of some importance for modern public health planning has recently been emphasized by Stead and his coworkers, who also pointed out that the technique used by Salo et al. did not permit distinction between infection with Mycobacterium tuberculosis and M. bovis.5 Among issues that remain moot are how tuberculosis rst reached the Americas, whether precolonial American tuberculosis was human or bovine, how widespread tuberculosis was among early indigenous Americans, whether tuberculosis was ever epidemic in the Americas in precolonial times, and whether there was sufcient tuberculosis in the prehistoric Americas to have inuenced herd immunity or lack thereof among native American people. I have recently reviewed the early history of tuberculosis in Africa.6 For Africa, there are accounts of tuberculosis by early missionaries and explorers, many of whom had medical backgrounds. Although some

of the indigenous people had developed hieroglyphics, which were used on stelae, no useful medical accounts have been found among them. The early explorers and settlers of the Americas included virtually no medical observers, and there is a paucity of relevant written material from these early explorers upon which to draw. The Spanish priest and chronicler, Fr. Diego Durn, recorded that the 50 to 60 slaves sacriced at the funeral of the Mexica ruler Axaycatl in 1479 included many hunchbacks,7 and we may suppose that some of them had Potts disease. On the other hand, there is a trove of American archeologic material. From this material, one can make a number of inferences and tentative conclusions, although it is unlikely that all observers will agree with whatever conclusions are drawn. The review presented here should stimulate further healthy scientic debate on this subject.

THE ORIGINS OF THE TUBERCLE BACILLUS

Most of the members of the genus Mycobacterium are water or soil organisms, and it is reasonable to suppose that the genus had its origins in such an environment. Distinct species have emerged over evolutionary time, and have been subject to selection pressures leading to differentiation. Hayman considered the habitat and geographic distribution of M. ulcerans.8 He concluded that this organism must have become established on the primitive continent of Gond-

Correspondence to: Thomas M Daniel, MD, Professor Emeritus, Center for International Health, Case Western Reserve University School of Medicine Room T-505, 10900 Euclid Avenue, Cleveland, OH 44106-4978, USA. Tel: (216) 3686321. Fax: (216) 368-8664. e-mail: tmd5@po.cwru.edu Article submitted 20 May 1999. Final version accepted 21 December 1999.

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wanaland approximately 150 million years ago. If his conclusions are correct, then the genus may predate the origin of primates, including Homo sapiens, and may have differentiated to include at least one modern species at that ancient time. Kapur, Whittam and Musser noted that M. tuberculosis has an unusual lack of nucleotide diversity and a much lower frequency of mutation than other bacteria.9 They estimated that the species originated 20 400 to 15 300 years ago. It has been asserted that M. tuberculosis evolved from M. bovis, making this evolutionary step in association with the domestication of cattle by humans.5,1012 In considering this hypothesis, it is rst important to note that these two species are both members of the M. tuberculosis complex and are so closely related genetically that they might well be considered a single species.13,14 Historically, they have been distinguished by only a few biochemical differences and by greatly different host ranges. M. tuberculosis is pathogenic for humans, other primates, and guinea pigs; M. bovis is pathogenic for many animals, although its historic host was almost certainly cattle, other species having acquired it from bovine sources. It has been argued that M. bovis is a more primitive species than M. tuberculosis because it carries only one copy of the transposon designated IS6110. However, single copy isolates of M. tuberculosis have been found in several locations in Asia by Fomukong and colleagues.15 They believe that the IS6110 genetic element is an ancient one present in both species and antedating the differentiation of these two species. The broader host range of M. bovis has also been used to argue that it is the more primitive species.12

communities are the Clovis, New Mexico, site in the American southwest, dated to about 11 500 years ago, and the Monte Verde site in central Chile, dated to about 12 500 years ago.25 An earlier site at Pedra Furada in northeastern Brazil dated to 14 300 years ago is not accepted by all authorities.18 In fact, more primitive habitation probably existed at the Monte Verde site as early as 33 000 years ago. Urbanization at these and other sites almost certainly developed independently. It is likely that the people of Monte Verde and similar sites were descendants of the earliest migrants, and one must also conclude that huntergatherer groups of humans were widely dispersed throughout the Americas at very early times.

PREHISTORIC TUBERCULOSIS IN THE AMERICAS

How and when did tuberculosis reach the Americas, and where did it come from? The earliest evidence for tuberculosis in the Americas comes from the studies of Peruvian mummies carried out by Allison, Arriaza, Salo, and their colleagues.1,26,27 Their work rmly establishes the presence of tuberculosis in the arid deserts of Peru and northern Chile, where mummication was widely practiced, during the rst millennium of the common era. In this region, substantial communities with inhabitants numbered in the thousands or tens of thousands also developed at that time, and this aggregation of people may have facilitated the spread of tuberculosis. The Table presents eleven precolonial instances of skeletal remains for which the evidence of tuberculosis is convincing, either including typical tuberculous spondylitis or identication of mycobacteria by acidfast staining or polymerase chain reaction DNA amplication. These cases begin in about the year 500, and continue until the beginning of the colonial era. If these cases are representative of the actual epidemiology, they suggest that tuberculosis may have moved northwards from South America, but one must be cautious in drawing such a conclusion because only in the arid regions of Chile and Peru did mummies survive. Arriaza and coworkers reviewed skeletal remains from Andean sites dating from 4000 years ago to about 500 years ago, at the inception of the colonial era.27 They found tuberculosis to be present only during the rst millennium of the common era, and then it was present in remains from only two sites, designated Maitas Chiribaya and Cabuza, at prevalences of 2.4 and 2.3 per cent. Morse has argued that the prevalence of putative bony tuberculosis observed in paleopathologic specimens is smaller than one should expect to nd if the disease was common in prehistoric America.2 Morses calculations were based upon data from native Americans at a time of enormous tuberculosis prevalence. He estimated that 7 per cent of persons with tubercu-

THE PEOPLING OF THE AMERICAS

All authorities agree that humans reached the Americas across a land bridge from Siberia to Alaska during epochs when the Bering Strait, which now separates the two continents, was above the level of the sea. Archeologic, linguistic, and genetic evidence support this hypothesis. Most authorities now believe that two major migrations took place, one about 25 000 to 20 000 years agoperhaps, in the view of some, beginning as early as 35 000 to 30 000 years ago and one about 12 000 to 11 000 years ago.1622 Between those two times, the Beringa land bridge was covered with glacial ice; after about 10 000 years ago it was submerged. The people who made this journey probably originated in Central Asia, and at least some of them probably migrated down the Pacic coast in boats.23,24 In more recent times, a third group moved across the Bering Strait to populate North American circumpolar regions. These early migrants were wandering hunter-gatherers, and they did not live in permanent sites, nor did they domesticate animals. The earliest established

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Table

Certain and probable precolonial tuberculosis in paleontological specimens in the Americas

Location Peru Chile Peru Tennessee, USA New Mexico, USA Tennessee, USA Peru Peru Illinois, USA Ohio Ontario, Canada 500 1040 700 2001000 8281130 10271617 Early colonial or late precolonial 10001300 8001050 1275 1490 Date

Investigator, reference Arriaza et al27 Case AZ71 T194 Case SRI Allison et al26 Morse2 Case 15 Case 10 Case 12 Case 4 Salo1 Buikstra and Cook28 Widmer and Perzigian29,30 Pfeiffer31

losis would have bone disease. Ortner and Putschar, using data from nineteenth century Europe, estimated that about 3 per cent of persons dying of tuberculosis would have bone disease.32 El-Najjar found osseous lesions in 8 per cent of skeletons of tuberculous individuals from the Hamann-Todd collection assembled in Cleveland, Ohio, in the early twentieth century.33 In these reports, classical spondylitis, the condition most easily identied as tuberculous and used as a criterion for inclusion in the Table ranged from 3 per cent33 to 14 per cent of bone disease.32 If we assume that about 6 per cent of precolonial indigenous Americans dying with tuberculosis had bone disease, then approximately 40 per cent of persons had some form of tuberculosis at death in Maitas Chiribaya and Cabuza, the two sites noted by Arriaza and his colleagues.27 While this calculation must be considered only approximate at best, it is sufcient to demonstrate that the archeologic evidence of tuberculosis in the precolonial Americas supports a concept of substantial epidemic disease at some locations at some times. Potts disease is identiable in early indigenous American art with varying degrees of certainty.34 There are many causes of dorsal kyphosis, but the dowagers hump of osteoporosis was probably not common among short-lived, prehistoric people. No meaningful inferences about tuberculosis prevalence can be made from art, but a few conclusions can be drawn. Figures depicting probable tuberculosis of the spine have been found in North America, South America, Central America, and the Caribbean. This implies that tuberculosis was widely dispersed, an implication that cannot be made from skeletal remains because climate does not favor the preservation of human remains in many locations. In a limited review of gures at the Museo Popol-Vuh in Guatemala, we noted possible gibbosities in about 10 per cent of gures from two sites, but none in gures from a third.35 Finally, artistic representation of Potts disease generally comes from cultures that ourished and often developed large centers of population during the sec-

ond half of the rst millennium of the common era or later, in concordance with paleopathologic ndings. There is nearly universal agreement that tuberculosis arrived in the Americas along with early people who crossed the Beringa land bridge. Tuberculosis was widely dispersed in Europe in neolithic times, but there is little rm evidence of tuberculosis in Asia at a time that antedates the early migration of people to the Americas. From India and China there are literary allusions to tuberculosis, but no archeologic specimens. It is perhaps most rational to assume that tuberculosis was not common among the early migrants or perhaps that it aficted only a few groups. At the same time, in order to understand the nding of tuberculosis in Chilean and Peruvian mummies, one is led to assume that tuberculosis came to the Americas with some of the earlier rather than later migrants. The earliest people of the Americas almost certainly lived in small, dispersed population groups. One can perhaps draw certain inferences about them from indigenous tribes now living in the remote areas of the upper Amazon River basin of Brazil. These people have little contact with one another and with modern Brazil, and they live in villages of about 300 persons; larger communities usually divide.36 Tuberculin reactor rates in these isolated populations range from 0 to 80 per cent.35,36 McGrath used a mathematical model to estimate that in prehistoric cultures a social network of between 180 and 400 persons was required for coexistence (endemnicity) of M. tuberculosis and humans; with smaller populations either the pathogen or the host did not survive.37 The parameters used to model the populations she studied may not be generally applicable to all early American populations, but the general concept that tuberculosis can exist in small, isolated groups is pertinent. Thus, it seems likely that early prehistoric people, whose huntergatherer groups were necessarily small, supported tuberculosis in their populations only occasionally, and that tuberculosis was a geographically dispersed disease. Only when larger, settled communities devel-

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oped did tuberculosis become epidemic. In the Americas, that appears to have happened about 1500 years ago.

M. TUBERCULOSIS OR M. BOVIS
There is a wealth of experience from the clinic and the autopsy table to indicate that M. bovis has a predilection for causing tuberculosis of the bones and lymph nodes in humans. This has led some to conclude that all of the bone tuberculosis observed in paleologic specimens, including that from the Americas, is bovine in origin. Hare, for example, asserts that all tuberculosis prior to about 4000 years ago was due to M. bovis,10 and Manchester accepts this view.38 More recently, Haas and Haas,12 Bates and Stead,11 and Stead39 all accept the view that early human tuberculosis was due to M. bovis. This line of thinking offers further support for the thought that M. tuberculosis originated by mutation from M. bovis. Cattle were domesticated in the eastern Mediterranean basin about 7000 to 9000 years ago, and perhaps even earlier in Africa.38,40 If M. tuberculosis has a bovine origin, it is important to realize that bovine tuberculosis is rarely transmitted by the aerial route and does not readily spread from human to human, so that epidemic spread could only have occurred after the new species had originated. This implies that the disease would have been maintained in animals until the time it crossed host species lines and differentiated into M. tuberculosis. The early nomads who crossed the Beringa land bridge were hunter-gatherers, and did not bring domestic animals with them. At none of the early American sites is there evidence of domestication of animals. Archeologic evidence from Argentina suggests that the rst animals domesticated, about 9000 years ago, were dogs.16 By the time of the arrival of the rst Spanish conquistadores, the Quechuans of the Andean altiplano had domesticated llamas and guinea pigs, called by the Spaniards conejos (rabbits) de los indios, and cattle were rst introduced into the Americas by the Spaniards. Guinea pigs are not hosts for M. bovis, and although bovine tuberculosis has been found in modern llamas, they are not a natural host for this organism. In fact, llamas are camelids, and were separated from other camels at the time of the breakup of Gondwanaland, approximately 150 million years ago. Wild bison hunted by early North Americans might have served as hosts for bovine tuberculosis, but one has to ask how they might have become infected. Thus, there appears to have been no logical primary host for M. bovis at the time when tuberculosis rst appeared among indigenous Americans. If one accepts, as I believe one must, the argument that the earliest tuberculosis in the Americas was caused by M. tuberculosis, then one has to doubt the widely accepted hypothesis that tuberculosis spread

from early domesticated cattle to humans. Unless identical mutations occurred independently in the earths two hemispheres, there is no reasonable way for M. tuberculosis to have existed on the eastern end of the Beringa land bridge unless it also existed on the western end, and the Bering Strait became covered with arctic sea water before the rst cattle were domesticated in Asia Minor.

SUSCEPTIBILITY TO TUBERCULOSIS OF NATIVE AMERICANS

It is abundantly clear that native Americans in historical times have become highly susceptible to tuberculosis. This fact was dramatically demonstrated by the course of epidemic tuberculosis among Eskimos.41 Similarly, tuberculosis found susceptible hosts among the indigenous people elsewhere in North America.4,42,43 In fact, it is apparent that most indigenous Americans were immunologically nave and probably genetically susceptible to tuberculosis when they met the often phthisical Europeans who colonized their lands. Grigg argues convincingly that tuberculosis epidemics have a duration of about 300 to 400 years, the epidemic ending with the development of herd immunity.44 He bases his thesis on data from Europe. A similar time period for an epidemic wave of tuberculosis, beginning in the Andean region of South America about 1500 years ago, would leave many generations for susceptibility to redevelop among native Americans. This epidemic probably did not reach some remote people, including the Brazilian Yanomami43 and the Eskimos;41 these populations were nave and extremely susceptible.

CONCLUSIONS
The information reviewed here allows one to conclude that tuberculosis reached the Americas with its early immigrants from Asia at least 10 000 years ago, possibly earlier than 20 000 years ago. It then must have persisted in some small, dispersed social groups for thousands of years. Tuberculosis prevalence appears to have risen to epidemic proportions about 1500 years ago, perhaps beginning in the Andean region of South America and spreading slowly northward. This epidemic appears to have subsided well before the arrival of European colonialists, leaving a highly susceptible indigenous population to be ravaged by the reintroduction of tuberculosis. There is no evidence to support the belief that tuberculosis in the prehistoric Americas was due to M. bovis, and considerable reason to doubt this hypothesis. Acknowledgements
The author wishes to thank Janet W McGrath for her helpful comments and review of this manuscript.

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References
1 Salo W, Aufderheide A C, Buikstra J, Holcomb T A. Identication of Mycobacterium tuberculosis DNA in a pre-Columbian Peruvian mummy. Proc Natl Acad Sci USA 1994; 91: 2091 2094. 2 Morse D. Prehistoric tuberculosis in America. Am Rev Respir Dis 1961; 83: 489504. 3 Morse D. Tuberculosis. In: Brothwell D, Sandison A T, eds. Diseases in antiquity. A survey of the diseases, injuries and surgery of early populations. Springeld, IL: Charles C Thomas, 1967: pp 249271. 4 Paulsen H J. Tuberculosis in the Native American: indigenous or introduced? Rev Infect Dis 1987; 9: 11801186. 5 Stead W W, Eisenach K D, Cave M D, et al. When did Mycobacterium tuberculosis infection rst occur in the new world? An important question with public health implications. Am J Respir Crit Care Med 1995; 151: 12671268. 6 Daniel T M. The early history of tuberculosis in central East Africa: insights from the clinical records of the rst twenty years of Mengo Hospital and review of the relevant literature. Int J Tuberc Lung Dis 1998; 2: 784790. 7 Thomas H. Conquest: Montezuma, Corts, and the fall of old Mexico. New York: Simon and Schuster, 1993: pp 26. 8 Hayman J. Mycobacterium ulcerans: an infection from Jurassic time? Lancet 1984; 2: 10151016. 9 Kapur V, Whittam T S, Musser J M. Is Mycobacterium tuberculosis 15 000 years old? J Infect Dis 1994; 170: 1348 1349. 10 Hare R. The antiquity of diseases caused by bacteria and viruses. A review of the problem from a bacteriologists point of view. In: Brothwell D, Sandison A T, eds. Diseases in antiquity. A survey of the diseases, injuries and surgery of early populations. Springeld, IL: Charles C Thomas, 1967: 115131. 11 Bates J H, Stead W W. The history of tuberculosis as a global epidemic. Med Clin N Am 1993; 77: 12051217. 12 Haas F, Haas S S. The origins of Mycobacterium tuberculosis and the notion of its contagiousness. In: Rom W N, Garay S M, eds. Tuberculosis. Boston, MA: Little, Brown and Company, 1996: pp 319. 13 Wayne L G. Mycobacterial speciation. In: Kubica G P, Wayne L G, eds. The mycobacteria. A sourcebook. Part A. New York, NY: Marcel Dekker Inc, 1984: pp 2565. 14 Eisenach K D, Crawford J T, Bates J H. Genetic relatedness among strains of the Mycobacterium tuberculosis complex. Analysis of restriction fragment heterogeneity using cloned DNA probes. Am Rev Respir Dis 1986; 133: 10651068. 15 Fomukong N G, Tang T H, Al-Maamary S, et al. Insertion sequence typing of Mycobacterium tuberculosis: characterization of a widespread subtype with a single copy of IS6110. Tubercle Lung Dis 1994; 75: 435440. 16 Fiedel S J. Prehistory of the Americas. 2nd ed. Cambridge: Cambridge University Press, 1992. 17 Weiss K M. American origins. Proc Nat Acad Sci USA 1994; 91: 833835. 18 Meltzer D J. Clocking the rst Americans. Annu Rev Anthropol 1995; 24: 2145. 19 Gibbons A. The peopling of the Americas. Science 1996; 274: 3133. 20 Wallace D C. Mitochondrial DNA in aging and disease. Scientic American August 1997: 4047. 21 Gibbons A. Mother tongues trace steps of earliest Americans. Science 1998; 279: 13061307. 22 Hurtago de Mendoza D, Braginski R. Y chromosomes point to native American Adam. Science 1999; 283: 14391440. 23 Sandweiss D H, McInnis H, Burger R L, et al. Quebrada Jag24

25 26

27

28

29 30

31

32

33

34 35

36 37

38 39 40 41 42 43

44

uay: early South American maritime adaptations. Science 1998; 281: 18301832. Keefer D K, deFrance S D, Moseley M E, Richardson J B III, Satterlee D R, Day-Lewis A. Early maritime economy and El Nio events at Quebrada Tacahuay, Peru. Science 1998; 281: 18331835. Grayson D K. Conrming antiquity in the Americas. Science 1998; 282: 14251426. Allison M J, Mendoza D, Pezzia A. Documentation of a case of tuberculosis in pre-Columbian America. Am Rev Respir Dis 1973; 107: 985991. Arriaza B T, Salo W, Aufderheide A C, Holcomb T A. PreColumbian tuberculosis in northern Chile: molecular and skeletal evidence. Am J Physical Anthropol 1995; 98: 3745. Buikstra J E, Cook D C. Pre-Columbian tuberculosis in westcentral Illinois: prehistoric disease in biocultural perspective. In: Buikstra J E, ed. Prehistoric tuberculosis in the Americas. Evanston, IL: Northwestern University Archeological Program, 1981: pp 115139. Perzigian A J, Widmer L. Evidence for tuberculosis in a prehistoric population. JAMA 1979; 241: 26432646. Widmer L, Perzigian A J. The ecology and etiology of skeletal lesions in late prehistoric populations from eastern North America. In: Buikstra J E, ed. Prehistoric tuberculosis in the Americas. Evanston, IL: Northwestern University Archeological Program, 1981: pp 99113. Pfeiffer S. Paleopathology in an Iroquoian ossuary, with special reference to tuberculosis. Am J Physical Anthropol 1984; 65: 18811889. Ortner D J, Putschar W G J. Identication of pathological conditions in human skeletal remains. Washington, DC: Smithsonian Institution Press, 1981. El-Najjar M Y. Skeletal changes in tuberculosis: the HamannTodd collection. In: Buikstra J E, ed. Prehistoric tuberculosis in the Americas. Evanston, IL: Northwestern University Archeological Program, 1981: pp 8597. Ponce Sangins C. Tenupa y Ekaku. La Paz, Bolivia: Los Amigos del Libro, 1969. Daniel T M. An immunochemists view of the epidemiology of tuberculosis. In: Buikstra J E, ed. Prehistoric tuberculosis in the Americas. Evanston, IL: Northwestern University Archeologic Program, 1981: pp 3548. Black F L. Infectious diseases in primitive societies. Science 1975; 187: 515518. McGrath J W. Social networks of disease spread in the lower Illinois valley: a simulation approach. Am J Physical Anthropol 1988; 77: 483496. Manchester K. Tuberculosis and leprosy in antiquity: an interpretation. Med Hist 1984; 28: 162173. Stead W W. Tuberculosis in Africa. Int J Tuberc Lung Dis 1998; 2: 791792. Kaiser J. Were cattle domesticated in Africa? Science 1996; 272: 1105. Grzybowski S, Styblo K, Dorken E. Tuberculosis in Eskimos. Tubercle 1976; 57 (Suppl): S1S58. Holmberg S D. The rise of tuberculosis in America before 1820. Am Rev Respir Dis 1990; 142: 12281232. Sousa A O, Salem J I, Lee F K, et al. An epidemic of tuberculosis with a high rate of tuberculin anergy among a population previously unexposed to tuberculosis, the Yanomami Indians of the Brazilian Amazon. Proc Natl Acad Sci USA 1997; 94: 1322713232. Grigg E R N. The arcana of tuberculosis. With a brief epidemiologic history of the disease in the USA. Amer Rev Respir Dis 1958; 78: 151172.

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RSUM

Nous avons pass en revue les preuves palontologiques de lexistence de la tuberculose dans les Amriques prcolombiennes afin dclairer ses origines et les consquences pidmiologiques qui en ont rsult. Le genre Mycobacterium existe de trs longue date et M. tuberculosis peut stre diffrenci il y a 20 400 15 300 ans. Les Amriques se peuplrent dimmigrants provenant dAsie au cours de deux migrations majeures : la premire qui est survenue il y a plus de 20 000 ans et lautre il y a 12 000 11 000 ans. La tuberculose est arrive aux

Amriques avec ces immigrants et elle a persist un faible niveau dendmicit dans de petits groupes de population disperss. Cest depuis 1500 ans environ que lpidmie de tuberculose a commenc probablement dans la rgion andine de lAmrique du Sud. Elle na pas progress ou sest rduite temps pour laisser en place des populations indignes amricaines fortement prdisposes la maladie au moment de la colonisation par les Europens.

RESUMEN

Se revisa la evidencia paleontolgica de la tuberculosis en la Amrica precolonial para aportar luz sobre sus orgenes y ulteriores consecuencias epidemiolgicas. El gnero Mycobacterium es muy antiguo y M. tuberculosis puede haberse diferenciado entre 20 400 y 15 300 aos atrs. Las Amricas estaban pobladas por inmigrantes de Asia, que llegaron en dos grandes contingentes, uno hace ms de 20 000 aos y el otro, entre 12 000 y 11 000 aos atrs. La tuberculosis lleg a las

Amricas con estas migraciones y persisti con un bajo nivel de endemicidad en grupos pequeos y dispersos de la poblacin. Una epidemia de tuberculosis comenz hace alrededor de 1500 aos, probablemente en la regin andina de Sud Amrica. No persisti o se redujo a tiempo para dejar una poblacin americana indgena altamente sensible cuando se produjo la colonizacin europea.