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2008 Parasite Pathoecology of Salmon Pueblo and Other Chacoan Great Houses: The eH althiest and Wormiest Ancestral Puebloans Karl Reinhard University of Nebraska-Lincoln, [email protected]

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Reinhard, Karl, "Parasite Pathoecology of Salmon Pueblo and Other Chacoan Great Houses: The eH althiest and Wormiest Ancestral Puebloans" (2008). Karl Reinhard Papers/Publications. 25. http://digitalcommons.unl.edu/natresreinhard/25

This Article is brought to you for free and open access by the Natural Resources, School of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Karl Reinhard Papers/Publications by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Published in: Chaco’s Northern Prodigies: Salmon, Aztec, and the Ascendancy of the Middle San Juan Region after AD 1100, edited by Paul F. Reed (Salt Lake City: The University of Utah Press, 2008), pp. 86-95 & 389-427. Copyright © 2008 The University of Utah Press. 5

Parasite Pathoecology of Salmon Pueblo and Other Chacoan Great Houses

The Healthiest and Wormiest Ancestral Puebloans

KARL J. REINHARD

ARCHAEOPARASITOLOGY AND affected parasitism at Antelope House and Salmon PATHOECOLOGY IN THE SOUTHWEST Ruin is another application ofpathoecology. Two fields of paleopathological investigation orig­ Ancestral Pueblo communities have long been inated in the Southwest. Archaeoparasitology is the focus of archaeoparasitology. Samuels (196S) the study of ancient parasite infection (Reinhard developed the methods for helminth (parasitic 1990, 1992b). It includes comparisons between worm) egg recovery with coprolites from Mesa time periods of single societies as well as compari­ Verde. Subsequently, Stiger (1977) provided the sons of parasitism between different, contempora­ first intersite comparison analysis for sites on Mesa neous cultures. For example, Fry (1980) compared Verde. Fry and his colleagues conducted the first re­ Fremont and Anasazi parasitism, and also Archaic gional comparisons ofparasitism, focusing on Can­ hunter-gatherer and ancestral Pueblo parasit­ yon de Chelly and Glen Canyon (Fry 1977; Fry and ism. All of these studies fall into the definition of Hal1197S, 1986). Fry (1977) also presented the first archaeoparasitology. cross-cultural analysis ofArchaic, ancestral Pueblo, By contrast, pathoecology is the reconstruction and Fremont sites, and pioneered the comparison ofrelationships among behavior, environment, and of parasitism among populations practicing differ­ disease organisms in the development of illness ent subsistence strategies. Building on this previous (Martinson et al. 2003; ReinhardandBuikstra2003; work, I have analyzed the diversity of helminths Reinhardet al. 2003; Santoro et al. 2003).1his field that parasitized ancestral Pueblo peoples (Rein­ developed from the need for fine-grained analysis hard 1985a, 1985b, 198sc, 1990; Reinhard et al. ofprehistoric ecological and behavioral conditions 1987). By 1985, archaeoparsitologists had identi­ to assess factors that affected disease. Pathoecologi­ fied eight species of helminth that infected ances­ cal interpretation depends on archaeological infor­ tral Puebloans (Figure S.I). mation regarding parasitism, community size, trade Aidan Cockburn's insight into the origins of patterns, water sourceS, subsistence practices, envi­ disease influenced the development of pathoecol­ ronment, medicinal use, and many other topics. Al­ ogy in the archaeoparasitology of ancestral Pueblo though the term is new, pathoecology developed sites. Cockburn (1967, 1971) argued that the evolu­ over several decades. I view EI-Naijar et al.'s (1976) tion of infectious diseases followed human evo­ study of ancestral Pueblo anemia as the first patho­ lution and the development of human cultures. ecology study. Perhaps the most advanced exam­ Inspired by Cockburn, Reinhard (198Sa) compared ple ofpathoecology is Stodder and Martin's (1992) the parasitic state of Colorado Plateau Archaic multifactorial perspective on ancestral Pueblo dis­ peoples to ancestral Pueblo an sites. He verified ease. My study (Reinhard 1996) of the factors that Cockburn's hypothesis that occasional infections

86 Parasite Pathoecology

~stercoralis n=3(0.6%) Strongylate worm I n=7(1.4%)

FIGURE 5.1. Diagram showing the wide spectrum of parasites that infected ancestral Puebloans. in hunter-gatherers became major health hazards vermicularis) prevalence in coprolites (Reinhard in agricultural populations. Reinhard (1988) pre­ 1988). The pinworm was chosen as an indicator of sented anumberofpathoecological findings regard­ general infectious disease because it is transferred ing the development ofparasitic disease in ancestral by person-to-person and by environmental con­ Puebloans relative to earli~r hunter-gatherers. Para­ tamination (Figure 5.3). sitism was limited in hunter-gatherers due to small Over millions of years of mutual evolution band size, band mobility, diffuse regional popula­ with hominids and modern humans, pinworms tions, and the presence of natural anti-helminthics have evolved multiple routes of infection, includ­ (worm poisons) in hunter-gatherer diets. Hunter­ ing anal-oral, hand-to-hand, and airborne routes. gatherer parasitism was promoted by the consump­ Pinworms are exceptionally remarkable among tion of uncooked meat and insects. Parasitism human parasites because the female worm wrig­ was promoted in ancestral Puebloan communi­ gles out of the anus of her host at night to scatter ties by contaminated water sources, concentrated her eggs. Once outside of the intestine, she dis­ populations, a more sedentary lifestyle, crowded perses eggs by two different mechanisms. Two (apartment-style) living conditions, establishment types of eggs are produced in two parts of the pin­ oflarge latrines, activities centered on water (agri­ worm uterus: light and heavy. Heavy eggs are laid culture), and activities that expanded wetlands (in­ on the perianal folds with an irritant excretion. eluding irrigation ofall types). The resulting itching (pruritis) and nocturnal host By the 1990S, Reinhard (1992a) had identi­ scratching transfers the infective eggs to the host fied wide variation in parasitism among ancestral fingers. Other eggs are distributed by aerosol when Pueblo villages (Figure 5.2). At some settlements, the female's desiccated body bursts, which releases parasitism was controlled, but others were over­ thousands of light eggs into the air. Ultimately, whelmed by their pathogens. This topic was ex­ these light eggs contaminate the environment, plored with a comparison of pinworm (Enterobius settling on food, in water, and throughout the KARL J. REINHARD

-----I I I , L- I I Colorado i Utah , 1- __

, 21%•

. ,• New

I Mexico

FIGURE 5.2. Map showing variation in percentages of pinworm parasitism among ancestral Pueblo villages.

Eggs are transferred by direct contact between humans and by t­ Inhalation ot eggs in air currents. ""

Noctural egg-laying results in retrointection as some eggs hatch and larvae enter the same host used by the mother worm.

FIGURE 5.3. Diagram showing modes of pinworm transmission to human hosts. habitation. How long these eggs remain infective perinanal region, and the larvae wriggle back into depends on warmth and humidity. In general, even the host. Hand-to-hand transfer of the eggs occurs in arid environments, human habitations have an when humans interact upon waking. Autoinfec­ elevated humidity. Thus, several infection routes tion occurs when humans eat food contaminated result from the pinworms' nocturnal excursions. with the eggs from their own hands. Airborne in­ Retroinfection occurs when the eggs hatch on the fection occurs when humans inhale the eggs, or

88 Parasite Pathoecology

30

Q) u c Canyon de Chelly Q) • -co Chaco Canyon 20 • Q)> a.... E .. Inscription House 0 10 • C Salmon Ruin -a..• • at 01---~~---P--.-~~-.--~--~~---t 40 50 70 80 90 0/0 Porotic Hyperostosis Prevalence

FIGURE 5.4. Graph comparing pinworm parasitism with porotic hyperostosis prevalence for several southwestern locales. The chart shows that the prevalence ofpinworm parasitism covaried with porotic hyperostosis prevalence at ancestral Pueblo sites where both coprolites and skdetons were studied (Reinhard 1992.a).

when the air dissemination of eggs results in the sance, but reflects a serious health risk, particu­ contamination offood and water. Ofcourse, other larly when one considers that other pathogens are pathogens follow the same hand-to-hand, hand­ spread by the same means. to-mouth, and aerosol routes as pinworm infec­ Reinhard (1992a) showed that the prevalence of tion. Therefore, high rates of pinworm prevalence pinworm parasitism covaried with porotic hyper­ suggest high rates of infection by other pathogens ostosis prevalence at ancestral Pueblo sites where that are passed through the same modes of infec­ both coprolite and skeletons were studied (Figure tion (see Figure 5.1). 5.4). Porotic hyperostosis is an indicator of gen­ Some ancestral Pueblo communities were ex­ eral skeletal pathology that has been used to assess tremely parasitized. In fact, some sites have the maternal-infant health. The fact that these indica­ highest levels of pinworm infection recorded for tors of disease had a positive, statistically signifi­ ancient or modern peoples. In a modern clini­ cant correlation underscores the use of pinworms cal setting, only 5 percent of feces from pinworm­ as a general gauge of ancestral Pueblo disease state infected people are positive for pinworm eggs. The (Reinhard 1992a). percentages of coprolites positive for pinworm Pinworms are not very pathogenic but are a from several sites far exceed this level. For example, good proxy for the infectious disease environment ~9 percent of the coprolites from Antelope House, (Reinhard 1996). Rates of pinworm infection at 19 percent of those from Inscription House, and ancestral Pueblo and Fremont culture sites were ex­ 21 percent of those from Chaco Canyon sites were plored by Hugot and his colleagues (1999). They positive for pinworm eggs. This indicates that pin­ found that sites in rockshelters without walled worm parasitism was unavoidable, and that in all villages (some Glen Canyon sites) had the lowest probability people had heavy infections. In such levels ofparasitism. Such sites had pinworm preva­ populations, pinworm infection is not just a nui- lence comparable to hunter-gatherers. Next, village KARL J. REINHARD sites outside of rockshelters had intermediate levels persal capability and the ability to find hosts when of parasitism. Finally, walled villages built within needed. Also, they must possess attributes enabling rockshelters had the highest prevalence of pin­ them to survive in the external environment. Fea­ worm (see Figure S.4). tures of the host have less effect on survival and re­ If we look at data from the Chacoan realm, we production of these parasites. find that Chacoan great houses are anomalous: Nidicolous parasites live in the host's immedi­ they include both the wormiest and healthiest ate environment: in beds, walls, granaries, caves, sites. Salmon Pueblo has among the lowest prev­ rockshelters, and under floors. Fleas, mites, bed­ alence values (7 percent). In contrast, Pueblo Bo­ bugs, triatomiid bugs, and the diseases transmitted nito and Pueblo Alto in Chaco Canyon are among by these bugs are examples of nidicolous parasites. the highest (21 percent). Clearly, the pathoecology They depend upon the host not only for food but of great houses was defined by factors other than for creation of their habitat. size. Puebloans living in great houses adapted their Permanent parasites live on or in the host ex­ use of the structures in ways that either promoted cept when dispersing between hosts. These in­ or limited parasitism. The remainder of this chap­ clude most protozoa, roundworms, flukes, and ter explores the factors that could have limited tapeworms. They are completely dependent upon parasitism at Salmon relative to other great house their host for food and all other environmental communities. requirements.

CHACOAN GREAT HOUSES FACTORS OUTSIDE GREAT AS NIDI FOR INFECTION HOUSE ENVIRONMENTS Realizing that for parasitic disease to occur, all fac­ Ttater Source, Giardiasis, tors related to the survival and reproduction of the and Amoebic Dysentery parasite must be present, Pavlovsky (1966) com­ Water sources in desert environments are foci for bined ecological factors into a predictive tool for in­ human activity and can therefore become nidi. As fection. These can include vectors, reservoir hosts, long as they are plentiful and flowing, and popu­ humans, and favorable external environments. He lations are not too concentrated around them, defined a nidus as that portion of a natural geo­ water sources are not necessarily a pathoecological graphic landscape that contains a community con­ factor in the spread of parasitism. However, when sisting of a pathogen, vectors, reservoir hosts, and water sources are few in number and stagnant, and recipient hosts, and that possesses an environment when populations aggregate around them, these in which the pathogen can circulate. He further sources often becomes contaminated, providing a found that pathogens possessed nidallty: the ten­ nidus and becoming significant pathoecological dency of an infectious agent to occur in distinct problems. nidi, such as being associated with particular geo­ Giardia lamblia has been found in ancestral graphic, climatic, or ecological conditions. Thus, a Pueblo coprolites (Gon'ralves et al. 2002). This par­ nidus is a focus of infection. For humans, a nidus asite is not highly pathogenic in most adults. In fact; can be as confined as a single room accessed by ro­ most infected people show no symptoms. How­ dents carrying plague-infected fleas, or as large as ever, when G. lamblia becomes established in stag­ an entire community and its agricultural use-area, nant water sources, it becomes a problem. It is most as is the case for the transmission of hookworms. perilous to pregnant women and their babies. Dis­ Various types of parasites circulate in nidi. ease in mothers and children is due to poor mater­ Temporary parasites, which live in the external en­ nal nutrition caused by malabsorption, resulting in vironment and come to the host only to feed, in­ intrauterine growth retardation. G. lamblia causes clude mosquitoes, chiggers, ticks, and leeches. In malabsorption when the intestinal villi become these species, every individual must have good dis- blunted and the function of intestinal mucous di- Parasite Pathoecology minishes (Carden ~d MacLeod 1988). Clinical the large bowel or ileum. Amoebas can cause nodu­ symptoms include cramps, watery diarrhea, nausea, lar granuloma formation, colitis, and diarrhea. The vomiting, and sometimes fever. Among pregnant disease can become systemic and eventually an ul­ women who exhibit symptoms, G. lamblia causes cerative disease of the large intestine, liver, lung, malabsorption and dehydration at a period when brain, or other organs. Amoebiasis can be symp­ there is a need for accentuated nutritional require­ tomatic or even fatal during pregnancy (Abioye ments. Such women fall into a negative nutritional 1973; Lee 1929; Lewis and Antia 1969; Rivera 1972). balance, as demonstrated by Carden and MacLeod Deaths that occur are due to a rapid onset of pro­ (1988), who summarized the effects of G. lamblia fuse diarrhea with dehydration and severe ane­ on the fetus and newborn. With protracted ma­ mia. Premature delivery results from colitis, diar­ ternal infections, normal fetal development is im­ rhea, dehydration, ketosis, or shock (MacLeod and peded. With asymptomatic maternal infections, Carden 1988). Weigel et al. (1996) found that high low birth weight and infant anemia are common E. histolytica load in asymptomatic infections was (De Morais and Suzuki 1997). Generally, infants associated with decreased maternal serum hemo­ become infected after three months of age. Islam globin and hematocrit levels, and iron-deficiency et al. (1988) found that some immunity is conveyed anemia. Among women who had severe prob­ from, mother to infant, but this immunity is not ef­ lems (spontaneous abortion, stillbirth, low-birth­ fective in infected infants. Immunity to G. lamblia weight babies), there was a fourfold increase in the increases with age (Shettyet al. 1992). Thus, the pa­ prevalence of amoebiasis relative to normal births thology caused by G. lamblia is most significant in (Czeizel et al. 1966). In infected but asymptomatic infants and toddlers (Hjelt et al. 1992). mothers, Weigel et al: (1996) found increased indi­ The epidemiology of giardiasis is well known cators of diminished intrauterine growth. Despite (Taus et al. 1998; Hjelt et al. 1992; Harter et al. immunity conveyed by antibodies passed through 1982). Sullivan et al. (199 I) showed that giardiasis is the placenta and in milk, infants can become in­ highly prevalent in children with chronic diarrhea fected. When this happens, infants exhibit fever and malnutrition, and that giardiasis does not re­ with severe watery, sometimes bloody, diarrhea. spond to standard therapeutic measures. Children Colitis, appendicitis, intestinal ruptqre, and peri­ who have low iron or vitamin BI2levels have more tonitis result in a high mortality among infected in­ severe giardiasis symptoms (Awasthi and Pande fants (MacLeod and Carden 1988). 1997; Olivares et al. 2002). Subadults in the age The Pueblo III occupation ofAntelope House, range of9 months to II years are most susceptible to at Canyon de Chelly in Arizona, is the best­ infection, though infections can occur at 3 months documented case of an ancestral Pueblo village of age. In developing nations, 91 percent of infants that suffered declining health due to water source of infected mothers become infected by 6 months nidi. Morris (1986) describes the pathoecological of age. Of infected infants, 86 percent have diar­ conditions that led to water contamination. To­ rhea. Infected people tend to live in dwellings with wards the end of the occupation, drought affected dirt floors, simple latrines, groundwater drinking the region. As more distant water sources dried up, sources, and close contact with dogs. These aspects the population ofAntelope House and Canyon de of life were common at ancestral Pueblo villages Chelly burgeoned. The increased population and (Reinhard 1996). In addition, person-to-person decreased water resulted in contamination. Gon­ transmission of G. lamblia is common (Birkhead ~alves et al. (2003) found both E. histolytica and and Vogt 1989; Black et al. 1977( Keystone et al. G.lamblia in Antelope House coprolites. El-Naijar 1978). The parasite Entamoeba histolytica also af­ (1986; EI-Naijar et al. 1976) found increased skel­ flicted ancient Pueblo groups (Gon~alves et al. etal evidence ofsystemic disease during the Pueblo 2002). Relative to G.lamblia, E. histolytica causes III occupation of Canyon de Chelly relative to more dramatic pathology, creating ulcerations in other time periods. Thus, there is a relationship

91 KARL J. REINHARD between environmental stress, increased parasit­ cate human hosts and burrow through the skin. ism, and skeletal indicators of morbidity in mother Also,A. duodenale can achieve transmammary and and infants. transplacental infection. Thus, fetuses and infants For Chaco an great houses, coprolites from can be infected without ever coming in contact Salmon Ruin were tested for G. lamblia with nega­ with contaminated soil. tive results (Wilson et al. 2006). The absence ofgi­ Hookworm causes specific problems in preg­ ardiasis at Salmon is logical given the presence of a nancy. One of the most common causes of death flowing water source (the San Juan River) within in labor in the developing world is cardiac failure 200 m of the community. Given these conditions, from severe anemia attributed to hookworm infec­ there was little chance for contamination. No copr­ tion (Cintron Villaronga 1967). As many as 90 per­ olites from Chacoan great houses have been tested cent of pregnant women are infected in endemic for E. histolytica. areas (Ananthakrishnan et al. 1997; Navitskyetal. 1998). Crompton and Whitehead (1993) present Irrigation, Hygiene, and Hookworm calculations comparing effects of hookworms on Hookworm has been found in coprolites from An­ a nonpregnant woman versus a pregnant woman. telope House and Pueblo Bonito, but at no other The model predicts that hookworms more rapidly ancestral Pueblo. Hookworm is the greatest para­ deplete stored iron, with a rapid effect on red cell sitic threat to the mother, fetus, and infant. Iron­ density per milliliter of blood in pregnant women. deficiency anemia resulting from intestinal blood Macleod (1988) verified this model from the clin­ loss is the major consequence of hookworm infec­ ical perspective. Each worm consumes 0.27 mlof tion (Variyam and Banwell 1982; Ali et al. 1990). blood per day, and only 20 weeks after initial in~ Treatment for this type of anemia is administra­ fection, hypochromic, macrocytic anemia can tion of iron supplements. According to Gilman develop. The minor symptoms of infection are (1982), development of hookworm-induced iron­ indistinguishable from complaints of pregnancy deficiency anemia is dependent on the intensity of (epigastric pain, heartburn, and so on). However, infection, the species of hookworm, and the ability with moderate infections there is low-grade fever, of the host to resist infection and to maintain ad­ fatigue dyspnea, heart palpitations, flow murmurs, equate stores of iron. loss of blood is caused by di­ and anemia. In heavy infections, constipation or rect ingestion ofred blood cells and by tissue trauma diarrhea, jaundice, emaciation, cardiac failure, or produced by worm attachment and feeding. pre-eclampsia occur. If a woman survives labor, she The species that causes the more serious pa­ cannot recover as easily from post-partum hemor­ thology and that has been identified in ancient rhage, which can contribute to maternal death. New World remains is Ancylostoma duodenale Hookworms also have a negative impact on fe­ (Allison et al. 1974). This is a fascinating, human­ tuses and infants (MacLeod 1988). Abortion, still­ specific parasite that has evolved several infection birth, and premature labor are associated with modes and adaptations. Perhaps the most remark­ severe hookworm infection. Women infected with able aspect of A. duodenale is its hypobiotic abil­ hookworm give birth to low-birth-weight infants ity. Hypobiosis occurs when a parasite suspends it (a 2 percent hematocrit drop in the mother corre­ development in host tissues in a way that prevents lates to a 100 gram decline in birth weight). Because a strong immunologic response. A. duodenale can of transplacental migration, infants are infected at go into hypobiosis in winter and come out of hy­ birth. Severe and sometimes fatal hemorrhage oc­ pobiosis in summer. This is a significant adapta­ curs in infants less than four months ofage. Chaud­ tion because the females can lay their eggs in the hary and Jayaswal (1984) first described an anemic season that is optimal for larval survival. The larvae infant resulting from transplacental migration. In hatch within a few days, exit the feces, and develop a survey of hundreds of transplacental-infected in­ through three larval stages as free-living soil nema­ fants in China, Yu et al. (1995) defined the symp­ todes. Subsequently, as third-stage larvae, they 10- toms of transplacental infection, which include

92 Parasite Pathoecology

bloody s~ools, melena, anorexia, listlessness, and village location that aggravated pinworm infec­ edema. A. duodenale was the species implicated in tion. Although pinworm prevalence was highest these types of infection. Transplacental migration in walled villages built within rockshelters, it is is not rare. Nwosu (1981) documented that 10 per­ important to note that no thoroughly studied an­ cent of 316 Nigerian newborns (four to five weeks cestral Pueblo site has been found to be pinworm old) were infected with A. duodenale. Transmam­ free. Related cultures also were infected. The earli­ mary infections from mother to infant also occur, est Basketmaker II coprolites from Bighorn Cave with similar health results (MacLeod 1988). Stud­ (Grand Gulch, Utah) have a prevalence of 25 per­ ies of many groups from around the world link cent. Later, the diffuse populations of ancestral hookworm disease, especially from A. duodenale, Pueblo and Fremont in the Glen Canyon area were to severe iron deficiency and anemia in children infected. Even the Sinagua inhabitants of Elden (Albonico 1998; Stoltzfus et al. 1998). Pueblo were infected (Hevly et al. 1979). The in­ Hookworm infection is dependent on mois­ fections resulted from air humidified by human ac­ ture, shade, and warmth. The Colorado Plateau is tivity and contaminated with floating eggs within normally too dry to promote infection, and in his­ confined spaces. toric times hookworm was unknown. Thus, the Although pinworm infection tends to he as­ discovery in Anasazi sites of hookworm eggs and ymptomatic, a high prevalence of heavy infections another parasite with a similar infection mode, can result in severe pathology, including secondary Strongyloides stercoralis, was surprising (Reinhard bacterial infections in juveniles. However, to my 1985C; Reinhard et al. 1987). Clearly, ancestral mind the real relevance ofpinworm relates to other . Puebloans created microenvironment nidi where diseases that are also airborne transferred. For the parasite larvae could hatch and mature in moist, ancestral Pueblos, tuberculosis was the other air­ warm, and shaded soil. Puebloans also spent time borne disease. Among the most poignant epidemi­ in these nidi, where they spread eggs and became ological descriptions of the tuberculosis threat to infected by larvae. It is very likely that irrigated Pueblos, is applicable to pinworms as well, is pro­ fields were hookworm and S. stercoralis nidi. vided by Fink (1985), who examined details ofAna­ Studies of hand and foot washing in Bengal sazi life such as communal living, lack ofknowledge show that the larvae can be washed offeasily within ofgerm theory, and cramped living conditions that a few minutes ofcoming into contact with the skin. promoted infectious diseases. The infection occurred in defecation grounds, and The San Juan period at Salmon Pueblo is enig­ washing was prescribed by religious rules (Nawa­ matic in the context of pinworm prevalence at linski et al. 1978). We do not know if ancestral other sites. Only 7 percent of 112 coprolites stud­ Puebloans had similar rules, but it is very likely that ied contained pinworm eggs-approximately one­ hookworms could penetrate the skin of Puebloan fifth the prevalence recorded for other Pueblo III farmers as they worked in irrigated fields. If the sites, including Pueblo Alto and Pueblo Bonito. division oflabor resulted in men working more in A probable explanation for this relates to Paul irrigated fields, it may be that they were more often Reed's (2006c) finding that the Sanjuan residents infected than women. of Salmon used primarily the second-floor rooms for human activities. Air conditions in the lowest INTERNAL GREAT HOUSE FACTORS rooms, and those closest to the windowless rear Apartments, Plazas, Kivas, wall, would have been more likely to increase pin­ and Second-Floor Living worm infection. Such rooms would have had the Pat Horne (1985) attributed the remarkable pin­ most stagnant and humid air, promoting airborne worm prevalence among ancestral Pueblos to infection with pathogens. Use ofhearths in second­ crowded, apartment-style living conditions. As floor rooms would have produced a much less hu­ noted above, Hugot et al. (1999) elaborated this mid environment, and any rooms opening towards theme by detailing the aspects of architecture and the large Salmon plaza would have been healthier

93 KARL J. REINHARD with ventilation from the dry, relatively breezy air the scolex. As the they progress downward along outside the pueblo. the length of the tapeworm, their ovaries and tes­ Subterranean rooms such as kivas would have tes mature, fertilization occurs, and eggs mature. provided the primary vector for transmitting air­ When a proglottid is filled with mature eggs, it is borne disease. In kivas, the air would have been said to be gravid. In some tapeworms, such as those humid, and the air flow around the ventilator would that infect humans who eat undercooked fish, have been sufficient to transmit particles around the the eggs are laid through gravid proglottid geni­ room, but not to remove infectious particles from tal pores. In other species, such as those that infect the structure. This would explain, in part, why Bas­ humans who eat undercooked beef, entire gravid ketmaker groups, who lived in pithouses with litde proglottids break off of the tapeworm. These pro­ or no circulation, developed a high prevalence of glottids are pardy motile and squirm their way out infection. After Basketmaker times, however, kivas of the host body. would have been the most likely subterranean nidi Hymenolepis lays eggs through genital pores, of pinworm dissemination. One way to test this and these eggs are infective when they pass into the hypothesis is through analysis and comparison of environment. Although they have been found in sediment samples from kiva, living room, and mill­ Canyon de Chelly coprolites, they have not been ing room floors. found in Chacoan great houses. By far the healthiest place to work and live at Salmon was outside, in the plaza and on rooftops. CONCLUSION Sunlight would have desiccated and radiated patho­ The Chacoan great houses provided many poten­ gens, thereby reducing the number of infectious tial nidi for temporary, permanent, and nidiculous airborne contaminants, and the clean air moving parasites. Great house inhabitants created or elim­ across the plaza would have provided people with inated nidi through different activities and prac­ alternate, healthy air. Humidity also was undoubt­ tices. Although there is was no way to completely edly much lower in these open spaces compared to eliminate the transmission of permanent parasites, confinedrooms and kivas. some aspects of life at Salmon retiuced the preva­ lence of pinworm relative to other great houses. Sanitation and Hymenolepidid A lower population density, among other factors, Tapeworm Injection would have accomplished this. With regard to tapeworms, there' are two main The absence of fecal-borne parasitism indi­ types of hosts. The definitive host is the animal cates that nidi of fecal exposure were eliminated at in which tapeworms accomplish sexual reproduc­ Salmon Ruin through the use of specific rooms as tion, whereas intermediate hosts are infected with latrines, an effective way of stopping the spread of nonsexual stages. Usually, tapeworm infections in parasites such as Giardia laMia. humans occur through ingestion of infected inter­ Nidi external to Salmon Ruin where hook­ mediate hosts. worm and S. stercoralis transmission could have The most common tapeworm (Hymenolepis taken place did not exist. This was probably due nana) found in ancestral Puebloan coprolites took to a different type of irrigation (perhaps using the a different infectious pathway. This tiny tapeworm free-flowing San Juan River) and gardening relative evolved the ability to use an intestinal villus as its to that of Pueblo Alto and Pueblo Bonito, where intermediate host. The larvae emerge and become hookworms did infect humans. adults in the intestinal lumen, thus using humans as In the future, more extensive analysis of ances­ both their definitive and intermediate hosts. tral Pueblo coprolites should be conducted using a Tapeworms have two methods of laying eggs. variety ofresearch methods. Some sites, such as An­ The tapeworm's anterior end, the scolex, attaches telope House, are currendy the focus of molecular, to the intestinal wall. Proglottids are the sexually immunological, and microscopic analysis. Other reproducing tapeworm organs that develop from sites, such as those in Glen Canyon, were studied

94 Parasite Pathoecology only through microscopy and could yield beneficial fore, archaeological excavations should include soil data with newer approaches. Once a range ofmeth­ samples from architectural remains to search for ods has been applied to a larger sample of sites, we the presence of insect exoskeletons. Also, analysis will be able construct a more complete picture of of trash sediments for all types of parasites must ancestral Pueblo parasite pathoecology. be developed in order to obtain parasitological In addition, parasitological methods must be data from sites that lack coprolites. Once these ap­ developed for analysis of remains other than copr­ proaches are developed, then a true archaeology of olites. Many nidiculous pathogens such as bed bugs parasitic disease will emerge. and kissing bugs live in walls and roofing. There-

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