advances in the paleopathology of andean south...

Journal of World Prehistory, Vol. 11, No. 2, 1997

Advances in the Paleopathology of Andean South America

John W. Verano I

Over the past 100 years, paleopathology has grown from the occasional description of the odd and curious to what is today a dynamic and multidisciplinwy field. South American skeletal and mummified remains played a key role in stimulating the development of the discipline in the late nineteenth and early twentieth centuries. The past 20 years have witnessed a renewed interest in the paleopathology of prehistoric Andean populations. Recent studies are characterized by growing sophistication in both research methods and theory, and significant advances have been made in understanding diet, disease, and general health patterns in the prehistoric Andean world.

KEY WORDS: paleopathology; skeletal biology; Andean area; mummies.

INTRODUCTION

Paleopathology is broadly defined as the study of disease in ancient populations. It is a field that traces its roots to various academic disciplines, including anthropology, pathology, and the history of medicine (Angel, 1981; Ubelaker, 1982). South American skeletal and mummified remains, primarily from Peru, Chile, and Bolivia, played a key role in stimulating the development of the field in the late nineteenth and early twentieth centuries (Mufiiz and McGee, 1897; Bandelier, 1904; Hrdli~ka, 1914; Moodie, 1923). Over the past 20 years there has been renewed interest in the paleopathology of prehistoric Andean populations, along with growing sophistication in research methods and theory. This article provides a brief historical background of the field and highlights some recent advances.

1Department of Anthropology, Tulane University, 1021 Audubon Street, New Orleans, Louisiana 70118.

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Fig. 1. Geographic extent of the Inca Empire at its apogee in the early sixteenth century (shaded area).

Andean South America

Andean South America constitutes a unified culture area that forms a natural unit of anthropological study (Moseley, 1992; Lechtman, 1996). Defined as that portion of South America which formed the Inca Empire at its period of greatest expansion, early sixteenth-century Andean South America stretched approximately 5000 km north-south, from the present day Ecuador-Colombian border in the north to the Maule River in central Chile to the south (Fig. 1). East-west it extended from the Pacific Coast

Paleopathology of Andean South America 239

to the eastern slopes of the Andean Cordillera (roughly 300-600 km in width). The Inca Empire itself was quite short-lived, existing for less than 100 years before being conquered by the Spanish in 1532 (Morris and Von Hagen, 1993). However, much of its cultural and administrative foundations were rooted in a long sequence of complex societies, several of which had, like the Inca, periodically united large areas of the Andean highlands and coast (Isbell and McEwan, 1991; Burger, 1992; Kolata, 1993),

Andean South America is littered with ancient ruins, abandoned agricultural terraces, and irrigation canals which provide evidence of dense and long-term human occupation of the region. The vast majority of these sites predates the Inca Empire, and some of the largest Andean cities were already abandoned or in decline by the fifteenth-century expansive phase of the Inca Empire (Rowe, 1963; Isbell, 1988). The earliest dates for human occupation of the Andes are still the subject of debate (eg., Lynch, 1990; Dillehay and Meltzer, 1991; Meltzer, 1993), but it is clear that the region records a long sequence of human adaptation to a uniquely diverse set of environments defined by the backbone of the Andes mountains.

Historical Background

Paleopathology in the Andean Area traces its roots back to the mid nineteenth century, with early collections of skeletal material made by explorers and anthropological expeditions (Squier, 1877; Dorsey, 1898; Bandelier, 1904; Hrdli~ka, 1914; Eaton, 1916). The development of paleopathology in South America followed a similar trend witnessed by the field in general (Buikztra and Cook, 1980; Ubelaker, 1982), from a focus on the single unusual speci- men to a paleoepidemiologieal approach, examining disease from a population perspective. While the identification of specific diseases in skeletal and mummified remains continues to be an important activity, researchers are increasingly interested in broader questions of human adaptation across space and time, the health implications of major shifts in settlement pattern and diet, and events such as European contact (Allison, 1984; Benfer, 1984, 1990; Verano and Ubelaker, 1992). New analytical techniques have joined traditional visual and radiographic methods to provide a substantial increase in the kinds of questions paleopathologists ask,

Data Sources

Information on health and disease in prehistoric Andean populations is available from various sources. Eyewitness accounts and secondary de-

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scriptions imbedded within the Spanish chronicles provide information on health problems and medical treatment in the Inca Empire (Rowe, 1947; Lastres, 1951). Ethnographic data from modern-day traditional healers may also provide insight into pre-Columbian health and disease (Bastien, 1987; Joralemon, 1993; Cabieses, 1993). Artistic depictions of individuals with missing limbs or other physical deformities are well-known from some pre- Columbian cultures such as the Moche of northern coastal Peru (Donnan, 1978; Weiss, 1958, 1961, 1984; Urteaga-Ballon, 1991). However, the most direct source of information on prehistoric health consists of skeletal and mummified human remains. These remains constitute one of Andean South America's richest archaeological resources.

DOCUMENTING SKELETAL PATHOLOGY

Pathological conditions commonly observed in skeletal remains include trauma, developmental and nutritional disorders, and various infectious, de- generative, and neoplastic diseases that can affect bone (Steinbock, 1987; Ortner and Putschar, 1981). While acute trauma and cultural practices such as cranial deformation are relatively easy to identify, diagnosis of specific diseases is more difficult. Only a relatively small number of diseases affect the skeleton, and some affect bone in similar ways (Buikstra, 1977; Ortner, 1991, 1992). Diseases that affect the skeleton are most commonly long-term ailments such as arthritis, chronic infections, and certain dietary deficiencies. Most illnesses that kill quickly, particularly acute infectious diseases such as influenza and pneumonia, do not leave their signature on the skeleton. Paleopathologists studying skeletal populations are therefore limited to a relatively narrow field of inquiry, although the development of new molecular techniques in recent years holds new promise for the diagnosis of disease in ancient skeletons.

Preservational conditions on the desert coast of Peru and Chile (and some highland microclimates) afford unique opportunities for paleopathological research. In addition to well-preserved human skeletons, mummified remains and coprolites have been recovered from numerous archaeological sites in South America, some dating back as early as 9000 years B.P. (Mufioz et al., 1993). The extremely dry western coast of Peru and Chile, in particular, is an exceptional preservational environment, and spontaneous mummification of bodies is common. Over the past 20 years, studies of these mummified remains have documented the presence of not only chronic infectious diseases such as tuberculosis and blastomycosis, but also acute infectious diseases such as bronchopneumonia and lobar pneumonia. Mummy dissections and eoprolite studies have provided additional

Paleopathology of Andean South America 2,41

insight into diet and intestinal parasite infestation in precontact populations of the Americas.

The past decade has seen renewed intensity in the study of South American mummies, brought about in part by the development of new laboratory techniques for paleodietary analysis and for the extraction and amplification of DNA from bone and mummified tissue (P/i/ibo, 1985; He,~-r~ann and Hummel, 1994; Lassen et al., 1994; Salo et aL, 1994). Re- search questions have evolved as well, reflecting the application of paleode- mographic and epidemiological models to the identification of disease in the archaeological record.

Disease and Community Health

Studies of disease and general health conditions in pre-Columbian An- dean populations fall into two broad categories: case reports and population studies. Case reports, which document specific pathologic conditions in single specimens have been important in establishing the presence of particular diseases in pre-Columbian New World populations (Allison et al., 1973, 1974, 1979; Kelley and Lytle, 1995). Population studies examine broader patterns of morbidity across space and time. Some of these focus on a single disease, such as tuberculosis or Carri6n's disease, and use demographic or epidemiological models to explain the observed frequencies and characteristics of skeletal and soft tissue lesions (Schultz, 1968; Buikstra and Williams, 1991; Arriaza et aL, 1995). Others look at general indicators of community health such as childhood growth, anemia, nonspecific bone infection, and dental disease (Elzay et al., 1977; Allison, 1984; Ubelaker, 1984, 1996; Benfer, 1990; Kelley et aL, 1991; Guill6n, 1992). All three kinds of studies have made important contributions to the understanding of patterns of health and disease in pre-Columbian South America (Verano and Ubelaker, 1991; Verano, 1992; Merbs, 1992).

New Analytical Techniques

In recent decades new analytical tools have been developed for reconstructing ancient diet, evaluating environmental health risks such as heavy metal poisoning, and identifying the ingestion of medicinal plants such as coca and tobacco. Elemental and stable isotope analyses of ancient human tissue have been used to reconstruct the paleodiet in various prehistoric population samples from Peru and Chile. The results of such studies have provided information on the relative importance of marine and terrestrial

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foods in the diet (Benfer, 1990; Sandness, 1992, 1995; Aufderheide et al., 1993; Verano, 1997b), dietary adaptations of highland immigrants to coastal environments (Aufderheide et aL, 1994; Aufderheide, 1996), the relative contribution of particular cultigens, such as maize, to the diet (Burger and van der Merwe, 1990; Sandness, 1992, 1995), and the possible population affiliation of sacrificial victims (Verano and DeNiro, 1993). In conjunction with the analysis of coprolites (Callen and Cameron, 1960; Weir and Bonavia, 1985; Holden, 1991; Holden and Nufiez, 1993; Reinhard and Bry- ant, 1992), such studies have made significant advances in reconstructing dietary patterns in prehistoric South America.

Chemical analysis of mummy hair has provided important information on the consumption of medicinal plants as well. Cartmell and colleagues (1991; Aufderheide, 1996) have identified cocaine and cocaine metabolites (indicating the chewing of coca leaves) in the hair of mummies from coastal Chile. Positive results first appear in a single mummy dated to approximately 3000 B.P. and become common in mummies from the later phases of the Alto Ramirez culture (2000-1650 B.P.). These results correlate well with archaeological evidence of increased cultural contact and exchange with the highlands (the intermediate source of coca) during this time period. Cartmell et al. (1995) also have found nicotine and its metabolite cotinine in prehistoric hair samples from Chile and Peru, confirming archaeological and ethnohistoric evidence of tobacco use by native South American populations (Wilbert, 1987). There appears to be potential for identifying metabolites of hallucinogenic plant extracts as well, which would open another avenue for ethnobotanical research. Finally, chemical analysis of mummified tissues has also been applied to evaluate suspected environmental hazards to health, such as chronic arsenic poisoning from local water sources (Allison and Gerszten, 1996).

TRAUMA

The general category of trauma includes acute injuries such as bone fractures and joint dislocation, as well as bony changes resulting from chronic stressors such as cranial deformation. Surgery, dental modification, and tattooing are also generally classified as forms of trauma (Ortner and Putschar, 1981; Merbs, 1989).

Early collections of Peruvian skeletal material made by Tello, Hrdli~ka, and others contain dramatic examples of skull fractures that document the damage potential of traditional Andean weapons such as clubs and slings (Tello, 1913; Tyson and Alcauskas, 1980; Merbs, 1989). Some of these collections also contain examples of trephination, a proce-


Fig. 2. Incomplete trephination associated with skull fracture: Cinco Cerros, central Peruvian highlands. (National Museum of Natural History, Smith-

sonian Institution Catalog Number 293780.)

dure in which a portion of the skull vault was cut, drilled, or scraped away. Many of these trephinations are associated with skull fracture (Fig. 2), an observation noted by early workers such as Tello and Hrdli~ka and confirmed by later researchers (Stewart, 1957; Allison and Pezzia, 1976; Ver- ano and Williams, 1992; Verano, 1997a). My own research with J. M.

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Fig. 3. Cranium with four healed trephinations: Late Horizon, Cuzco area. (Museo Arqueol6gico de la Universidad San Antonio Abaci, Cuzco, Catalog Number 2580.)

Williams of Hahnemann University on a sample of 621 trephined skulls from coastal and highland Andean sites found a frequent association between skull fracture and trephination. Trephinations associated with visible skull fracture ranged from a low of 6.8% in crania from Paracas on the south coast of Peru, to 11.8% in crania from the southern highlands of Peru, to a high of 26.2% in crania from the central Peruvian highlands. Actual frequencies were probably higher, since the trephination procedure itself probably removed the evidence of fracture in some cases (Verano, 1997a). Head injury and its complications were probably the primary mo- tivation for the practice in prehistoric South America, as is the case in modern times among traditional surgeons in Kenya (Margetts, 1967; Fur- nas et al., 1985). Survival rates for prehistoric Andean trephinations are impressive, particularly in later samples from Inca sites (Fig. 3). Evidence of long-term healing (extensive remodeling of the margins of trephinations) was present in 32.7% of trephinations at Paracas, 53.8% in the central highland sample, and 70.9% in the Inca sample from the southern highlands of Peru (Verano, 1997a).

Trephination is known to have been practiced in various parts of Peru and Bolivia for over 2000 years, from around 2500 B.P. until the midsix- teenth century (Verano and Williams, 1992; Verano, 1997a). Curiously, al-


Fig. 4. Nasea trophy head. (Museo National de Antropologfa, Arqueologfa y Historia, Lima, Catalog Number AF:7047.)

though there is abundant osteological evidence that trephinations were performed by the Inca, there is no mention of the practice in the Spanish chronicles (Lastres, 1951; Lastres and Cabieses, 1960). The tradition may have been lost following the Spanish conquest, although Adolf Bandelier (1904) claimed that trephination was still being practiced by traditional healers in Bolivia in the late nineteenth century.

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Warfare, Trophy Taking, and Human Sacrifice

I have recently published a review (Verano, 1995b) of the ethnohistoric and archaeological evidence for human sacrifice, secondary offerings of human remains, and the collection and curation of human body parts among prehistoric Andean societies. Human bodies and body parts were manipu- lated and curated in complex ways by Andean peoples. Human sacrifice was practiced by the Inca and by various cultures before them, and the offering of human remains in ceremonial contexts is a long Andean tradition (Scho- binger, 1991; Reinhard, 1992; Rowe, 1995). Early Spanish chroniclers reported that the Inca sacrificed war prisoners and collected a variety of body parts as war trophies (Rowe, 1946, p. 279; Hemming, 1970, p. 54), and this practice also appears to have considerable time depth in the Andes.

Mummified "trophy heads" are well-known from the Paracas and Nasca cultures of the south coast of Peru (Fig. 4). The diagnostic features of these heads include a hole in the forehead area for the attachment of a suspensory cord and damage to the base of the skull through which the brain was removed. More than 100 of these heads exist in museum collections, and additional examples are known from private collections (Baray- bar, 1987; Verano, 1995). There has been some debate about whether these heads were the product of warfare or human sacrifice (Tello, 1918; Baray- bar, 1987; Proulx, 1989), and one researcher has even suggested that they might represent ancestor worship (Guillrn, 1992). From my own examination of 84 Nasca trophy heads, I have concluded they are most likely war trophies, based on my assessments of the age and sex of the individuals (primarily adolescent and adult males), the way in which the heads were prepared, and the contexts in which trophy heads appear in Paracas and Nasca art (Verano, 1995; also Proulx, 1989; Silverman, 1993).

Interpreting the specific context and meaning of human offerings is difficult for pre-Inca societies, for which ethnohistoric data are sparse or absent, although artistic depictions of warfare, prisoner taking, and sacrifice can provide important interpretive clues in some cases (Donnan and Castillo, 1992; Alva and Donnan, 1993). For example, I used both ethnohistoric accounts and iconography, in conjunction with osteological analysis, to interpret a mass burial of sacrificial victims (Fig. 5) found in 1984 at the site of Pacatnamu in the Jequetepeque River Valley (Verano, 1986, 1994). More recently, the first skeletal evidence of large-scale prisoner sacrifice by the Moche of northern Peru was found at the Pyramid of the Moon in the Moche River Valley (Bourget et al., 1996; Verano, 1997c). To date, more than 50 skeletons have been excavated from two sacrificial contexts at the site. The demographic characteristics of the victims and patterns of trauma observed on their skeletal remains correspond closely to depic-


Fig. 5. A mass burial of 14 individuals at Paeatnamu, during excavation.

tions of prisoner sacrifice in Moche art (Alva and Donnan, 1993; Verano, 1997e). The discovery at the Pyramid of the Moon is important in providing the first well-documented archaeological evidence of human sacrifice among the Moche.

Cultural Modification of the Body

Common ways in which the body can be culturally modified include cranial deformation, dental modification, tattooing, and scarification. All were practiced in prehistoric Andean South America, although some cus- toms were more widespread than others. "l~ttooing (Fig. 6) is known on the central and north coast of Peru from at least 1500 B.P. onward, as documented by mummified remains from various sites (Ubbelohde-Doering, 1967; Vreeland and Cockburn, 1980; Allison et al., 1981b; Verano, 1995a). Individuals with tattoos or scarification are also represented in Moche art (Donnan, 1978). Moche artists also depicted what appears to be intentional mutilation of the nose and lips of certain individuals. This has generally been interpreted as representing punitive mutilation (Donnan, 1978; Ur- teaga-Ballon 1991), although some examples more likely depict congenital malformations such as cleft palate or damage caused by infectious diseases

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Fig. 6. Right arm of a young adult female from the Late Period occupation of Pacamamu (ca. 900-600 B.R) with tattooing (H28 Burial 5).

such as leishmaniasis (see Leishmaniasis and Chagas' Disease, below). In- dividuals with missing limbs were also depicted by Moche artists. Urteaga- Ballon (1991) has reported examining 65 ceramic vessels of individuals with missing limbs and argues that they represent surgical amputations. Ander- son and Verano (1996) have recently reported what may be the first osteological evidence of a Moche amputation from the monumental center of El Brujo in the Chicama Valley. It is the skeleton of an adult male missing all bones of the feet. The distal ends of the left and right tibiae and fibulae show extensive bony outgrowths and nonfunctional joint surfaces suggesting loss of the feet followed by healing (Fig. 7).

Cranial Deformation

Cranial deformation, both unintentional and intentional, was by far the most widely distributed form of body modification among prehistoric Andean populations. Unintentional flattening of the back of the skull as a

Fig. 7. (a) Moche ceramic vessel of an individual with missing feet, holding what appears to be a prosthesis. Height, 21 cm. [Courtesy of the American Museum of Natural History (Catalog Number IM919); photo by the author.] (b) Distal ends of the tibiae and fibulae of an adult male Moche burial from Huaca Cao Viejo, El Brujo, with extensive bony outgrowths covering the distal articular surface~

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Fig. 8. (a) Cradleboard found at a Late Intermediate Period site (ca. 900-500 B. P.) in the Viru Valley. Dimensions: 68 x 26 crn. (Museo Arqueol6gico de la Universidad Nacional de Trujillo, Peru.) (b) Doll placed on the cradleboard to demonstrate how an infant's head was fixed to the board by cloth bands.


Cranial Deformation

Cranial deformation, both unintentional and intentional, was by far the most widely distributed form of body modification among prehistoric Andean populations. Unintentional flattening of the back of the skull as a result of cradleboarding of infants is common in cranial samples from the central and north coast of Peru, and actual examples of cradleboards have been recovered archaeologically (Fig. 8) (Verano, 1994, 1997b). Moreover, intentional shaping of the head was a widespread practice in the Andes at the time of Spanish contact and is frequently noted in early colonial sources (reviews by GuiU6n, 1992; Hoshower et aL, 1995). Since the late nineteenth century, various schemes have been proposed to classify the varieties of cranial deformation found in the Andean Area. The best-known of these is the general descriptive system of Imbelloni (1933). Other workers have attempted to develop more fine-grained classificatory schemes linking cultural association and deformation type and, where possible, deforming ap- paratus (Weiss, 1961, 1972; Tello and Xesspe, 1979; Allison et aL, 1981a; Munizaga, 1987; Gerszten, 1993). These studies document great diversity in deformation techniques. At the same time they highlight some of the problems involved in trying to classify the end result of a complex interplay between a specific deformation device and a growing infant cranium. Clas- sificatory schemes utilizing idealized skull shapes do not adequately reflect within-sample variability resulting from variation in the placement of the deforming device, the amount of pressure applied, and other idiosyncratic factors. Hoshower et aL (1995, p. 146), in a recent study of cranial deformation in southern Peru, caution against simple assumptions about the re- lationship between deformation type and cultural identity, since head shape "may signal group membership at any one of a number of levels: regional, community, and/or lineage." While cranial deformation was clearly an important social marker in prehistoric Andean South America, there is still work to be done to define better its cultural and temporal dynamics.

INFECTIOUS DISEASE

Various bacterial, protozoal, and parasitic infections plagued prehistoric populations of Andean South America. The documentation of these diseases is important not only for the perspective it provides on prehistoric health in the Andes, but also for understanding the geographic distribution of particular diseases prior to extended contact between Old and New World populations (Verano and Ubelaker, 1992; Merbs, 1992; Larsen and Milner, 1994). Historically, the presence or absence of diseases such as tu-

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berculosis and syphilis or of parasites such as hookworm in the New World has been a subject of considerable debate. Recent advances in paleopathology have resolved some, but not all, of these issues.

Tuberculosis

There has been a long debate over whether tuberculosis was present in the New World in pre-Columbian times (Buikstra, 1981). The number of reported skeletal lesions suggestive of pre-Columbian tuberculosis has grown rapidly in recent decades, but confident diagnosis of many of these cases has remained elusive. Since the early 1970s, however, South American mummified remains have provided convincing evidence that tuberculosis was indeed a pre-Columbian health problem.

In 1973, Allison et al. published the first diagnosis of pre-Columbian tuberculosis based on both skeletal and soft tissue evidence. The case involved the mummy of a child from a Nasca (ca. 1300 B.P.) cemetery in southern coastal Peru. The mummy showed evidence of chronic bone and soft tissue disease suggestive of tuberculosis, but more importantly, Allison and colleagues were able to extract and identify acid-fast bacilli morphologically similar to Mycobacterium tuberculosis. Since this pioneering study, many additional cases of probable tuberculosis have been identified from skeletal and mummified remains from southern Peru and northern Chile (Allison et al., 1981c; Buikstra and Williams, 1991; Arriaza et aL, 1995). While differentiating bone lesions produced by tuberculosis from those of other mycotic diseases is challenging (Buikstra, 1977, 1981; Ortner and Put- schar, 1981), recent developments in DNA recovery and amplification by the polymerase chain reaction (PCR) have resulted in a major advance in the identification of tuberculosis in ancient remains. Salo et al. (1994) recently reported the extraction of DNA characteristic of M. tuberculosis from a pre-Columbian mummy from southern Peru. Similar identifications of M. tuberculosis and M. leprae DNA in skeletal and mummified remains have been reported by other researchers, supporting the utility of PCR amplification of ancient DNA in the identification of pathogenic organisms (Spigelman and Lemma, 1993; Lombardi, 1994; Raft et at , 1994). It now appears unequivocal that tuberculosis was present in the Americas before European contact. Some questions for future research include the original source (or sources) of the disease and the amount of genetic variation among ancient and modem strains of M. tuberculosis.

Treponemal Disease

Since the late nineteenth century there also has been debate over whether syphilis was present in prehistoric South America (Stewart, 1950a).


A number of examples of probable treponemal disease have been reported from Peru (Williams, 1932; Weiss, 1984; Zink et al., 1994; Tyson and AI- causkas, 1980), but convincing evidence of venereal or congenital syphilis has been elusive. A nonvenereal treponemal disease appears to have been common in some prehistoric populations of northern Chile. Most cases con- sist of individuals with pronounced tibial inflammation, although one individual showed cranial involvement as well (Allison et al., 1982; Standen et al., 1984; Arriaza, 1995).

Rothschild and Rothschild (1995) have recently presented criteria for distinguishing between the different treponemal syndromes that affect bone (yaws, bejel, venereal syphilis). From their examination of prehistoric skeletons from the site of San Lorenzo in Ecuador, they conclude that lesions symptomatic of venereal syphilis are present (Rothschild and Rothschild, 1996). If these results are confirmed, this would indicate that both venereal and nonvenereal forms of treponemal disease Were present in South Amer- ica prior to European contact.

Carri6n's Disease

Carri6n's disease, also known as bartonellosis, is caused by the bacte- rium Bartonella bacilliformis. It is a disease transmitted by sandflies that is endemic to certain highland valleys of modern-day Peru, Ecuador, and Co- lombia. Carri6n's disease manifests i~elf in two distinct forms--an acute form known as Oroya fever, which infects red blood cells and has a high mortality rate, and a relatively mild second stage, characterized by wart-like skin lesions, known as verruga (Schultz, 1968; Recavarren and Lumbreras, 1972).

In 1974, Allison and colleague s (1974a) identified an example of the verruga phase of Carri6n's disease in a pre-Columbian mummy dating to the Middle Horizon (ca. 1400-1100 B.P.) from the south coast of Peru. Their diagnosis was made on the basis of skin lesions containing clumps of bacilli morphologically similar to Bartonella bacilhformis. Interestingly, this was a mummy buried on the coast, in an area where Carri6n's disease presumably was not endemic. The mummy was from a cemetery showing highland (Huari) influence, and the individual may have been an immigrant from the highlands. No additional cases of Carri6n's disease have been reported in mummies subsequent to Allison and co-workers' (1974a) publi- cation, although examples of ceramic vessels depicting individuals with verruga-like skin lesions are known from the Moche and Chancay cultures of central and northern coastal Peru (Moodie, 1923, p. 488; Schultz, 1968; Weiss, 1984, Fig. 11).

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Leishmaniasis and Chagas' Disease

Protozoal diseases known to have been endemic to South America include leishmaniasis and Chagas' disease. Leishmaniasis (uta), an in- sectborne disease found in portions of Peru and Brazil today (Omran, 1961), appears to have afflicted prehistoric Andean populations as well. Most of the evidence is indirect and based on artistic representations of individuals with facial lesions typical of the mucocutaneous form of the disease (Weiss, 1961; Urteaga-Ballon, 1991), although a skull from Chavifia in the central highlands of Peru presents bony changes suggestive of the disease (Merbs, 1992). Reinhard has also reported a mummy from the Chiribaya Culture of southern Peru that shows facial lesions suggestive of leishmaniasis (Reinhard, 1995).

Chagas' disease is caused by infection with the protozoan parasite Try- panosoma cruzi. The most common route of infection is through contact with the feces of reduviid bugs of the subfamily Triatominae, although other forms of transmission are known (Rothhammer et al., 1985). The most common species of reduviid bug found in Andean South America is Triatoma infestans, which frequently occupies the roofs and walls of houses in rural areas. After malaria, Chagas' disease is the most common vectorborne disease in present-day Latin America (Schofield et al., cited by Rothhammer et al., 1995). Recent studies of South American mummies suggest that the disease has a long history of adaptation to human populations in the Andes. Rothhammer et al. (1985) reported nine possible cases of Chagas' disease in mummies from the Quebrada de Tarapacfi in northern Chile dating from between approximately 2450 B.E and 1400 B.P. Their diagnosis was based on the presence of "megavisceral syndromes"--enlargement of the colon, heart, or esophagus, which are typical manifestations of infection with T. cruzi. More recently, Fornaciari and colleagues (1992) have reported a study of an Inca period mummy from Cuzco that showed similar megavisceral syndromes. Ultrastructural and immunohistochemical study of the mummy indicated infection with T. cruzi.

Parasites

Studies of mummies and coprolites have demonstrated that some parasites previously thought to have been Old World introductions in fact plagued pre-Columbian New World populations. These include at least one species of hookworm, Ancylostoma duodenale, and the whipworm Trichuris trichiura (Allison et al., 1974b). Research has also documented the presence of roundworms (Ascaris lumbricoides), marine tapeworm (Diphyllobothrium


pacificum), pinworm (Enterobius vermicularis), hairworm (Strongyloides spp), Trichinella spiralis, Echinococcus granulosis, and the ectoparasite Pediculus humanus (louse). Also likely present were Entamoeba spp. and Moniliformis clarki, a thorny-headed worm (Patrucco et aL, 1983; Home, 1985; Reinhard, 1990, 1992; Confalonieri et al., 1991). Coprolite samples from the central coast of Peru document the presence of tapeworm and roundworm in cultural strata dating to approximately 4800 B.P. and pinworm at approximately 4300 B.P. While some parasites show great antiquity, most of the evidence has come from later agricultural populations living in sedentary villages.

Parasitism may, in part, provide a key to explaining regional variation in the frequency of porotic hyperostosis and cribra orbitalia in prehistoric Ecuador and Peru. Porotic hyperostosis and cribra orbitalia, porous lesions found on the external table of the skull vault and orbital roofs, are generally considered to represent a physiological response to iron deficiency anemia, which can be caused and exacerbated by a variety of factors, including dietary deficiencies, intestinal parasites, infectious disease, and their synergistic effects (Goodman et al., 1984; Kent, 1986; Walker, 1986; Stuart-Macadam, 1987). No evidence has been found to suggest that hereditary anemias such as sickle cell and thalassemia were present in the Americas in pre-Colum- bian times, so these can be excluded as possible causes (Ortner and Put- schar, 1981, p. 259; Merbs, 1992).

In his studies of skeletal samples from prehistoric Ecuador, Ubelaker (1992) found a significantly higher frequency of porotic hyperostosis in coastal populations. He hypothesized that parasitism, due to hookworm and possibly other parasites which require warm and damp conditions to propa- gate, may have been a significant contributing factor, citing similar geographic patterning of hookworm infestation in modern Ecuador. Ale~ Hrdli~ka (1914, pp. 57-59) and Pedro Weiss (1961, pp. 125-127) found similar differences in the frequency of porotic hyperostosis in prehistoric Peruvian crania from coastal and highland sites. While recognizing that parasitism is probably not the only factor that might explain these differences, Ubelaker (1992, 1996) believes that it played a significant role in the patterning of porotic hyperostosis observed in prehistoric population samples from Peru and Ecuador.

NEOPLASTIC DISEASE

Evidence of cancer is relatively rare in the paleopathological record (Steinbock, 1976; Ortner and Putsehar, 1981). This rarity has stimulated considerable debate over whether cancers are indeed relatively rare or

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whether short life expectancy, misdiagnosis, or other factors might explain the small number of cases reported (Zimmerman, 1977; Micozzi, 1991a, b; Roberts and Manchester, 1995).

Soft tissue tumors are particularly rare in the paleopathology litera- ture. In autopsies performed on over 1000 mummies from Chile and Peru, Gerszten and Allison found only two examples: a subcutaneous lipoma on the right side of the chest of an adolescent male and a rhabdomyosarcoma on the right cheek of a male child approximately 12 to 18 months old (Gerszten and Allison, 1991). Tumors affecting bone, whether primary or metastatic in origin, are more widespread, probably due to the greater availability of skeletal material relative to mummified soft tissues. Never- theless, the number of cases reported in South American skeletal remains is relatively small (MacCurdy, 1923; Moodie, 1926; Steinbock, 1976; Tyson and Alcauskas, 1980; Ortner and Putschar, 1981; Baraybar and Shimada, 1993; Verano, 1997b).

POPULATION STUDIES

Temporal Trends in Human Health

Rather than focusing on specific diseases, some researchers have attempted to identify general trends in human health through time. Most focus on the effect of subsistence and settlement pattern changes on community health, as measured by the frequencies of skeletal and dental stress indicators such as Harris lines, enamel hypoplasias, periostitis, porotic hyperostosis, dental caries, and other indicators (Goodman et al., 1984). The results of these studies reveal some general patterns in health through time, thus providing important baseline data for future comparative studies.

Robert Benfer and associates have examined the early transition from migratory hunting and gathering to sedentary village life and incipient agriculture on the central coast of Peru at the site of La Paloma, which dates between 8000 and 4500 B.P. (Benfer, 1984, 1990; Quilter, 1989). Based on the age distribution of burials in different occupational levels and the relative frequency of skeletal stress indicators, Benfer concluded that although the earliest inhabitants of La Paloma were severely stressed, there was evidence of decreased infant mortality, greater adult life expectancy, and some indications of improved nutritional status with the adaptation to sedentism and incipient food production.

Robert Malina (1988a, b) has published a study of the skeletal remains of 67 individuals from Late Preceramic tombs (ca. 3400-3800 B.P.) at the site of La Galgada in the northern highlands of Peru. Malina compared

Palcopathology of Ande~m South America 257

mortality profiles at La Galgada with those of the later population at La Paloma and a found lower infant mortality and a slightly greater life expectancy at La Galgada. An interesting finding in the La Galgada sample was the relatively high incidence of fractures. Malina found that 4 of 45, or 9%, of adults presented at least one healed fracture. Most of the fractures were of the wrist (Colles' fracture), ankle, or collarbone and are probably the result of accidents rather than violence, with osteoporosis being a possible contributing factor in some cases (compression fractures of verte- brae). Although few comparative data are available from other highland sites, the risk of accidental fracture was probably substantially higher for highland populations than it was for coastal inhabitants.

Marvin Allison (1984) has published an overview of temporal trends in paleopathology based on a study of 16 populations from coastal Peru and Chile ranging in time from approximately 4000 B.P. to the early colonial period. Allison found no evidence of a general improvement in health with the adoption of sedentism and intensification of agricultural production. On the contrary, he concludes that sedentary village life was generally det- rimental to health, due to crowding and sanitation problems, and that the increasing social stratification seen at some later sites resulted in health benefits only for the elite minority. The results of some of my own work on the north coast of Peru are consistent with the latter observation. At the site of Paeatnamu, I found that skeletal indicators of anemia (porotic hyperostosis and cribra orbitalia) were much more common in lower-status cemeteries than in higher-status cemeteries during the Late Intermediate Period (ca. 900-600 B.P.) occupation. Samples from earlier Moche cemeteries at the site showed fewer indications of anemia and did not present significant between-cemetery differences. Based on these differences, I hypothesized that there was an increase in social stratification during the later occupation of Pacatnamu that resulted in deteriorating health conditions for the nonelite population (Verano, 1992).

In studies of human skeletal samples from various time periods in precontact Ecuador, Douglas Ubelaker found a general increase in skeletal and dental stress indicators through time. Ubelaker found that while early coastal populations had low levels of infectious disease, anemia, dental caries, and various measures of nutritional stress, samples of human remains from more recent times show regular temporal increases in the frequencies of these problems. Such trends are particularly notable on the coast and correlate with increased sedentism and a less varied diet (Ubelaker, 1984, 1992, 1996).

While Benfer's study of La Paloma suggests that there was some improvement in general health and life expectancy during an initial adaptation to sedentism and food production on the central coast of Peru, surveys of

258 Verano

long-term temporal trends in skeletal pathology by Allison and Ubelaker point to a general deterioration in health conditions with increasing sedentism, agricultural intensification, and population growth. Studies of mummified human remains from coastal Peruvian and Chilean sites indicate that both chronic infectious diseases, such as tuberculosis, and acute pulmonary infections (Allison, 1984) were significant factors in morbidity and mortality in prehistoric Andean populations. Coprolite studies have documented the presence of various intestinal parasites in Andean populations as early as 5000 B.E, and some data suggest that parasite infestation may have been an important contributing factor in elevated frequencies of porotic hyperostosis seen in later prehistoric coastal Ecuadorian and Peruvian populations.

Studies in paleopathology indicate that prehistoric South American populations faced significant health challenges in their adaptation to sedentism, agricultural production, and population growth. With increasing population size and density came inevitable problems of sanitation, parasitism, and an increase in infectious disease. Agricultural intensification and social stratification led to a less varied diet for many, exacerbating the effects of infectious disease and parasitism. It should be emphasized, however, that such problems were not unique to populations of Andean South America. Similar challenges were faced by human populations worldwide in the transition from mobile foragers to sedentary agriculturists (Larsen, 1995).

The Biological Effects of European Contact

Native populations of Andean South America experienced major demographic decline following European contact in the sixteenth century. Warfare and social disruption were contributors, but the most important factor was a series of epidemics of European diseases that swept through Andean cities in the sixteenth and seventeenth centuries (Dobyns, 1963; Cook, 1981, 1992; Verano, 1992).

To date, historic documents are the primary source of information on changing health conditions following European contact because few sys- tematic studies of human skeletal remains from the early colonial period have been conducted. An exception is recent research by Douglas Ubelaker (1994, 1995a,b) on skeletal samples from historic churches in Quito, Ecua- dor. Ubelaker's research provides important comparative data on skeletal and dental health indicators before and after European contact, and dem- onstrates the potential for further studies of changing patterns of morbidity and mortality following contact.


CURRENT CHALLENGES AND FUTURE PERSPECTIVES

Over the past 100 years, paleopathology has grown from the occasional description of the odd and curious to what is today a dynamic and multidisciplinary field. This field has seen substantial advances in both research methods and theoretical perspective, although challenges still re- main. Diagnosis of specific disease conditions in archaeological remains is not always simple since bone can respond to disease in only a limited number of ways, and different diseases may produce similar lesions. More- over, diseases and their hosts evolve through time in response to changing environmental conditions and cultural practices, and this can be expected to result in variable expression of diseases in past populations. One of the most basic challenges, however, continues to be the need for accurate description and communication of paleopathological observations. Several recent publications address this issue and provide recommendations for recording observations on skeletal lesions (Ortner, 1991, 1992; Buikstra and Ubelaker 1994).

Paleodietary reconstruction from trace elements and stable isotopes also faces methodological challenges. Trace element studies have to take into account the potential for significant diagenetic alteration of bone and soft tissue following burial (Aufderheide, 1989), and detecting diagenesis has proven to be more difficult than previously suspected (Sandford, 1993; Edward and Benfer, 1993), While diagenetic change does not provide as great a challenge to stable isotopic analysis, early models for estimating the relative proportion of dietary components based on stable isotopic ra- tios have been shown to be oversimplified (SiUen et al., 1989; Ambrose, 1993). Despite methodological challenges, however, paleodietary analysis continues to be a dynamic and growing research area.

One of the important achievements of Andean paleopathology has been the development of a population perspective on the health and demography of prehistoric Andean peoples. In recent years a growing number of carefully excavated human skeletal samples has become available for study, making it possible to examine in greater detail patterns of disease and demography across space and time. Ongoing studies of mummified remains and coprolites promise to provide additional insight into the history of specific infectious and parasitic diseases. Andean South America is an unusually rich field for research into the health challenges of sedentism, changes in subsistence pattern, and population growth. Upcoming decades should see substantial progress in our understanding of the unique adaptations of prehistoric Andean populations to these challenges.

260 Verano

ACKNOWLEDGMENTS

I am grateful to Jane Buikstra and Douglas Ubelaker for critical read- ings of a preliminary draft of this paper and to several anonymous reviewers for helpful comments. Financial support for aspects of research reported in this article came from a Tulane University Committee on Research Sum- mer Fellowship, Fulbright Lectureships (1989, 1996) from the Council for the International Exchange of Scholars, and travel funding from the Re- search Opportunities Fund of the National Museum of Natural History, Smithsonian Institution. I am grateful to the following museums and archaeological projects for permission to study and photograph their collections: in Peru, the Museo Arqueol6gico de la Universidad National de Trujillo; the Museo National de Antropologia, Arqueologla y Historia, Lima; the Museo Arqueol6gico de la Universidad San Antonio Abad, Cuzco; and the Proyecto Arqueol6gico Complejo El Brujo; and in the United States, the National Museum of Natural History, Smithsonian In- stitution, and the American Museum of Natural History.

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