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February 2008

Pathoecology and the Future of Coprolite Studies in Bioarchaeology

Karl J. Reinhard University of Nebraska at Lincoln, [email protected]

Vaughn M. Bryant Jr. Texas A&M University

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Reinhard, Karl J. and Bryant, Vaughn M. Jr., "Pathoecology and the Future of Coprolite Studies in Bioarchaeology" (2008). Papers in Natural Resources. 43. https://digitalcommons.unl.edu/natrespapers/43

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Pathoecology and the Future of Coprolite Studies in Bioarchaeology

Karl J. Reinhard University of Nebraska, Lincoln

Vaughn M. Bryant, Jr. Texas A&M University

COPROLITE STUDIES IN (1963) and Martin and Sharrock (1964), but BIOARCHAEOLOGY especially in the seventies and eighties, coprolites were the focus of many interdisciplinary research Human coprolites currently provide an theses and dissertations (Aasen 1984; Bryant 1969; expanding array of information about the diet, health, Clary 1983; Edwards 1990; Fry 1969; Hall 1969, and ecology of prehistoric people in the Southwest, 1972; Kelso 1971, 1976; Napton 1970; Reinhard but for many years coprolites were not recognized or 1985a, 1988a; Sobolik 1988, 1991; Stiger 1977; preserved, or they were not considered important and Stock 1983; Van Ness 1986; Williams-Dean 1978). thus were not saved (Bryant and Dean 2006). With A large number of articles, chapters, and the expansion of archaeological field work during the monographs were published as a result of this early last half of the twentieth century archaeologists have work (Bryant 1974a, 1974b, 1974c; Bryant and increasingly explored the “complete” potentials of Williams-Dean 1975; Callen and Martin 1969; Clary sites, including the collection and analysis of 1981,1984,1987; Cowan 1967; Cummings 1994; Fry geomorphologic, botanical, and faunal data. In some 1977, 1980, 1985; Fry and Hall 1975; Fry and Moore ideal habitats (e.g., very dry or frozen) this includes 1969; Heizer and Napton 1969, 1970; Hevly et al. exploring the scientific potential of human coprolite 1979; Hogan 1980; Holloway 1983; Kelso 1970; studies. This is not easy to do: very few coprolites Lindsay 1980, 1983; Martin and Sharrock 1964; have what might be considered a “characteristic Moore et al. 1969; Napton 1969; Reinhard 1988b, shape and size.” In our experience, the majority of 1992a; Reinhard and Clary 1986; Reinhard et al. coprolites are usually fragmented, flattened by age, 1985, 1987, 1988; Roust 1967; Scott 1979; Steele or in many cases are preserved as amorphous masses 1969; Winter and Wylie 1974). As summarized by of various sizes similar in shape to “paddies” left Reinhard and Bryant (1992) these works explored behind by cattle. These flat, amorphous human the application of many fields to coprolite analysis coprolites are especially common in sites used by including archaeopalynology, archaeobotany, foragers with diets very high in plant fiber. archaeoparasitology, zooarchaeology, biochemistry, Coprolites and coprolite fragments are sometimes starch analysis, and phytolith analysis. (Phytoliths collected in situ during archaeological excavations, are microscopic mineral deposits produced by plants but most often they are found during screening, when within their cells. Phytoliths are extremely durable dirt is being separated from artifacts. If and their morphologies are frequently specific to unrecognized, coprolites may be crushed into dust, family, genus, and even species.) along with clods of dirt, and their contents lost. Each research project offered new In the American Southwest the arid climate, methodological innovations and several salient protected sites, and dry rock shelters provide some of works came from this period. Williams-Dean’s our best areas in North America for the preservation dissertation (1978) was a milestone in combining of human coprolites and the long record of biological studies of modern feces with the contents of Archaic history they help to reveal. Starting in the early Period coprolites. She was also the first researcher to 1960s with the pioneering efforts of Eric Callen present a holistic dietary reconstruction including

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archaeopalynology, archaeobotany, and factors such as crowding, sanitation, hygiene, and zooarchaeology from the same coprolite series. Aasen trade. They also include biotic factors such as (1984) pioneered the application of pollen presence of pathogens, disease reservoirs, and concentration and anthracology to Basketmaker intermediate hosts. Finally, physical factors such as coprolites. Fry and Moore (1969) and Moore and climate and soil conditions can be studied. Englert (1969) demonstrated that parasite eggs could be Pathoecology began to emerge in the Southwest with recovered from 10,000 year-old coprolites. Stiger the establishment of a link between the emergence of (1977) and Fry (1977) integrated dietary and parasitic disease and Ancestral Pueblo cultural parasitological analyses to present a picture of development (Reinhard 1988b). Later, this approach pathology and diet for Pueblo and Fremont cultures. In was applied to the biarchaeological problem of the this period, Bryant (1974a, 1974b, 1974c; 1975) etiology of anemia resulting in porotic hyperostosis emerged as the first coprolite specialist who fostered (Reinhard 1992a). From 1990 through 2000, much of many graduate student studies at Texas A&M the coprolite research, especially University (Bryant and Williams-Dean 1975; Edwards archaeoparasitology, shifted from the American 1990; Jones 1988; Reinhard 1988a; Reinhard and Southwest to South America (Bouchet et al. 2003; Bryant 1992; Sobolik 1988; Stock 1983; Weir et al. Chaves and Reinhard 2006; Dittmar et al. 2003; 1988; Williams-Dean and Bryant 1975). Gonçalves et al. 2003; Iñiguez et al. 2003; Reinhard In other parts of the world, coprolite analyses and Buikstra 2003; Reinhard and Urban 2003; Sianto were applied to bioarchaeology (Anderson 1967; Callen et al. 2005), as research groups developed in Peru, 1967; Weir et al. 1988) while in the Southwest coprolite Chile, and Brazil. The Escola Nacional de Saúde analyses usually fell into the realm of paleoethnobotany Pública, Fundação Oswaldo Cruz in Rio de Janeiro and dietary reconstruction. However, there was became a coprolite research hub for the Western considerable bioarchaeological influence on coprolite Hemisphere. During this period new methods for research at Texas A&M University and at Northern diagnosing disease based on gross pathology, Arizona University which resulted in the application of immunology, and molecular biology were developed coprolite studies to explaining osteological expressions and a paleoepidemiological approach also emerged of disease (Reinhard 1985a, 1988a). At the University (Aufderheide et al. 2004; Reinhard and Buikstra of Utah and the University of Colorado, parasitology 2003). began to be incorporated in thesis research resulting in Along with these new methods, Martinson et al. Fry’s (1977) and Stiger’s (1977) applications of (2003) codified the concept of “pathoecology” to parasitology to evaluate the symptoms that would have explain patterns of parasitic disease in archaeological been suffered by prehistoric people. Bioarchaeologists sites in the Moquegua Valley of southern Peru. were quick to incorporate the new coprolite data, Martinson et al. (2003) showed that the parasitism at especially archaeoparasitology, into their models of several villages was defined by occupation, trade, disease causation (Akins 1986; Merbs and Miller 1985; status, presence of domestic animals, and site Stodder 1984, 1987; Stodder and Martin 1992). location relative to fresh water access. In one village, Unfortunately, the numbers of theses and which specialized in fishing, the inhabitants were dissertations devoted to coprolite studies has parasitized by a tapeworm species, Diphyllobothrium declined during the last 15 years. Only four graduate pacificum that survives when fish is undercooked research projects with a focus on Southwestern and then eaten. In another village further inland, coprolites have been completed since 1991 (Androy where the main economy specialized in the herding 2003; Danielson 1993; Hansen 1994; Nelson 1999). of llamas, some of the herders may have suffered Perhaps what is needed is a new theoretical approach from dermatitis caused by a species of ectoparasites that will attract young researchers to consider recovered in ancient llama coprolites. Trade also coprolite analyses and bring them “back into the seems to have spread parasites. As fish was traded to laboratory.” Bioarchaeology is a natural future host inland groups, Diphyllobothrium infection spread for coprolite research, providing that coprolite inland to areas where agriculture was the main researchers in the Southwest can develop a subsistence base. Overall lower levels of conceptual framework applicable to bioarchaeology. Diphyllobothrium infection are found in mummies from a higher status site, indicating that social status PATHOECOLOGY: A NEW also impacted parasitism. Local site conditions also APPROACH affected parasitism. The coastal site examined had no source of clean drinking water, which explains the Pathoecology is the study of the environmental evidence of fecal-borne disease. The presence of determinants of disease (Martinson et al. 2003; domesticated guinea pigs and dogs in human Reinhard 2008a, 2008b). These include human habitations promoted the spread of Chagas disease. Pathoecology and the Future of Coprolite Studies in Bioarchaeology 207

Population aggregation in sites occupied during Inca Salmon Ruin located on a terrace above the San Juan times was yet another factor (Santoro et al. 2003), River. The subsistence data from the coprolites since in populated areas crowd diseases spread verifies that Antelope House inhabitants used wet rapidly by person-to-person contact and fecal-borne areas extensively for various economic activities and pathogens. were thus exposed to wireworms and hookworms In the late nineties, these pathoecological (Reinhard 1985b; Reinhard et al. 1987). The studies of past human populations in the Peruvian sanitation system at Salmon Ruin was also more Andes began to influence Southwestern coprolite efficient at sequestering feces and therefore reducing studies. The methods developed in South America the occurrence of infections from fecal remains. were applied to a limited number of Ancestral Hugot et al. (1999) addressed pinworm Pueblo coprolites (Gonçalves et al. 2002, 2004; infection and its variation among Southwestern sites. Reinhard et al. 2001) after the discovery that Pinworm is a crowd disease evident in 0 percent to Ancestral Puebloans were infected by protozoa and 25 percent of coprolites from Anasazi sites. In hookworms. In combination with previous work modern clinics, tests reveal that only five percent of (Reinhard 1990) these newer studies revealed that pinworm-infected patients pass pinworm eggs in the Ancestral Puebloans were infected with Giardia feces. If this equivalent rate was also present in lamblia, (cause of “beaver fever “), Entomoeba ancient human populations, then the very high histolytica (cause of amoebic dysentery), Trichuris prevalence of pinworms at some sites begs trichura (whipworms), Ascaris lumbricoides (giant explanation. Hugot et al. (1999) compared the intestinal roundworms), Ancylostomidae pathoecological influences of site location and (hookworms), Acanthocephala (thorny headed architecture against the prevalence of pinworm worms), Strongyloides stercoralis (threadworm), infection. They found that sites in caves had a higher taeniid tapeworms, hymenolepidid tapeworms, prevalence of pinworm–infected coprolites than open Enterobius vermicularis (pinworm), ticks, lice, and sites. Large pueblos had a higher prevalence of possibly flukes. Relative to other parts of the infection than small pueblos. Large pueblos built in Americas where there is excellent preservation and rock shelters had the highest prevalence of infection intensive archaeological research (central Mexico, of any sites studied. Therefore, it appears that human northeastern Brazil, Chile, and Peru), the Southwest crowding, combined with the pinworm’s spread via has evidence for a greater diversity of parasites airborne contamination, promoted infection to levels infecting Ancestral Puebloans than any other known that were unprecedented and are unequaled in the prehistoric culture. The variation, diversity, and modern clinical literature regarding pinworm prevalence of infections between pueblo sites is ideal infection. for the application of pathoecology as an interpretive The lessons learned from the Hugot et al. framework. (1999) study should be important to The pathoecological approach is exemplified in bioarchaeologists interested in spread of airborne the comparative study of the parasite ecology of diseases such as tuberculosis. Since pinworm and Salmon Ruins, New Mexico, occupied from A.D. tuberculosis infections are both airborne, one might 1088 to the mid-thirteenth century (Reed 2006; anticipate that large communities, especially those in Reinhard 2008b) and Antelope House (occupied rock shelters like those with the highest pinworm from A.D. 500 to 1250) located in Canyon de Chelly, rates, would also have been more prone to Arizona (Morris 1986; Reinhard 1996). Reinhard tuberculosis infection. (1996; 2008b) reviewed the pathoecology of the Pueblo III (A.D. 1100 – 1300) occupations of the APPLYING THE NIDUS CONCEPT IN sites. This study incorporates data on site ecology, subsistence, sanitation, food storage, and other data THE STUDY OF PREHISTORIC revealed by the archaeological work to explain the HEALTH dramatic difference in parasitism between the two sites. The elevated levels of parasitism at Antelope A paper currently in press (Reinhard 2008a) House resulted in higher rates of anemia, as indicated proposes that bioarchaeologists adopt the concept of by porotic hyperostosis in cranial vaults in the the nidus (Pavlovsky 1966) as a tool in burials from Antelope House: 88 percent of reconstructing the pathoecology of infectious disease subadults affected (El-Najjar 1986:219), compared to in prehistoric communities. The nidus is a 43 percent in subadults from Salmon Ruin (Berry geographic or other special area containing 1983). In essence, Antelope House was built in a pathogens, vectors, reservoir hosts and recipient canyon bottom location that was more conducive to hosts that can be used to predict infections based on the life cycles of a greater variety of parasites than one’s knowledge of ecological factors related to 208 Reinhard and Bryant

infection. Ecological factors include the presence of Puebloan pathoecology. They found that in West vectors, reservoir hosts, humans, and external Bengal, villagers defecated in areas around the environment favorable for the transmission of village peripheries. People used the same areas day parasites. An individual nidus therefore reflects the after day for this purpose. As a result, hookworm limits of transmission of a given parasite or pathogen larvae proliferated in those areas and the time spent within specific areas of interaction: bedbugs in a by humans in the contaminated areas was sufficient bedroom, for example. Thus, a nidus is a focus of for hookworm infection. At Antelope House, infection. A nidus can be as confined as a single hundreds of small concentrations of coprolites were room containing a bed and with access to the room found within the rock shelter. This pattern of by rodents carrying plague-infected fleas. However, coprolite remains shows that the Antelope House a nidus can also be as large as the community and its residents had a similar sanitation pattern as the surrounding area in which there is a transmission of people living in West Bengal, but used abandoned hookworms. rooms rather than the peripheral regions around the Pueblos are a complex of overlapping nidi. For village. Therefore, the nidi for hookworm and example, at Antelope House the grain storage and wireworm infections could have been in the processing rooms could be identified as a nidus for peripheries of the village or in the abandoned rooms. tapeworm infections. The habitation rooms within Wireworm larvae were found in Antelope House dog the shelter formed a nidus for pinworm infection. coprolites (Reinhard 1985b, 1985c) suggesting that The water sources were nidi for whipworm, Giardia, dogs were a reservoir within the Pueblo for and amoeba infections. Finally, the defecation and/or wireworm infection. agricultural areas were nidi for hookworm and By building a multi-room, stone-walled village wireworm transmission. within a rock shelter, Puebloans at Antelope House Hymenolepis nana is a tapeworm that has only established a large nidus for the transmission of been recovered from agricultural sites and most pinworm. Pinworms are transmitted both by hand-to- commonly uses grain beetles as intermediate hosts hand contact between humans and by airborn- and rodents as definitive hosts. Humans become dissemination when a population is occupying a infected when they eat food made with grain closed space. The apartment-like pueblos with their contaminated with beetles. The beetles and other closed spaces and stagnant air would have been ideal insects become infected when they feed on the feces areas for the dissemination of eggs (Hugot et al. produced by infected rodents. Some grain beetles are 1999; Reinhard 2008a). Proof for this is noted at quite small, only 2-3 mm long, therefore it is quite Antelope House, which has one of the highest easy to overlook them when selecting grain for prevalences of pinworm infections due to airborne processing and eating. Alternatively, H. nana contamination. evolved a direct life cycle without intermediate Whipworm, Giardia, and amoebas are hosts. Nevertheless, it is very likely that rodents and transferred in contaminated water, although hand-to- grain beetles in or around grain storage areas hand transfer of Giardia is known. This would have promoted the cycle of the tapeworm infections. Once been a serious problem during a period of drought in humans processed the contaminated grain, human the Canyon de Chelly region in Pueblo III times infection was promoted. (Morris 1986; Reinhard 2007c). During the drought Hookworm and wireworm larvae penetrate the the number of Pueblos in the canyon grew as people skin of humans. Nidi for these parasites probably moved closer to the remaining reliable sources of were in agricultural fields and/or in or around toilet water in the canyon floor. As a result of areas in the sites. The most common method to overcrowding and poor sanitation the local water control for hookworms is to use footwear. Ancestral sources became contaminated and thus could be Puebloans wore sandals, but Behnke et al. (2000) considered as nidi for fecal-borne diseases. document problems with the wearing of sandals among modern agriculturalists in Mali. They state PARASITES, DISEASE, AND (2000: 352) “soil sticks to sandals, making them uncomfortable and frustrating to wear when tilling ANCESTRAL PUEBLOAN soil, and risking damage. As a result of this practice, DEMOGRAPHY those who often wore shoes still became infected through bare skin.” This same situation may have Parasitic infection does not always provoke plagued Puebloan horticulturalists. Therefore, larval disease. In fact, some parasite species rarely cause skin penetration could have occurred in cornfields. disease. Disease results from the pathogenicity and Schad et al. (1983) suggest another scenario for virulence of the parasite infection, combined with hookworm infection that is also applicable to characteristics of the host. Virulence refers to a Pathoecology and the Future of Coprolite Studies in Bioarchaeology 209

parasite’s ability to multiply. Pathogenicity refers to immune status, and the behavioral patterns of the the parasite’s capacity to cause disease symptoms population that increase or decrease risk. Two of and mortality in host populations. Disease may be these, susceptibility to infection and the nutritional passed by vertical transmission, from parent to child, needs of the host, can be assessed through or lateral transmission from one human to another examinations of the archaeological record. An (except from parent to child). example of this can be seen in the studies of sites in Among the various worm types of parasites that the American Southwest. A number of coprolite infected Ancestral Puebloans, the most virulent studies reported that the breadth of Ancestral Pueblo included whipworms which have the potential for diets was generally nutritionally sound (Cummings each female to lay as many as 20,000 eggs per day, 1994; Fry 1980; Minnis 1989). However, as early as and giant intestinal roundworms which lay 200,000 1992, Reinhard began finding coprolite evidence for eggs per day. Perhaps the least virulent worm substantial variation in the consumption of wild parasite was wireworm. Female wireworms lay only plants at Ancestral Puebloan sites, suggesting that hundreds of larvae per day. However, the pathogenic agricultural failures at some locations may have potential of these worms is reversed. Whipworms forced a much higher reliance on wild foods. and giant intestinal roundworms rarely cause To test this assumption we compared the debilitating long-term problems or death. In contrast, ancient diets reconstructed for the populations living wireworms are more pathogenic. Wireworms enter at Antelope House and at Salmon Ruin to determine the body by penetrating the skin. In their migration whether or not we could identify clues reflecting toward the intestine they work their way through the starvation in coprolite data (Reinhard 2008b). These heart and lungs and can cause serious pathology in coprolite studies included identification and these organ systems. When they enter the intestine evaluation of starvation foods based on phytoliths they plow through the intestinal mucosa and damage (Reinhard and Danielson 2005), pollen the intestinal tract. Mothers infected with wireworms concentrations (Reinhard et al. 2006a), and starch can potentially infect their nursing babies in vertical and macrofossils (LeRoy-Toren and Reinhard 2004; transmission via breast milk. Whipworms and giant Reinhard 2008b; Reinhard et al. 2006b; Sutton and intestinal roundworms can be transferred only by Reinhard 1995). Data from both sites covered the fecal contamination of the environment. These are time period during the recorded environmental “geohelminths” which refers to parasites whose eggs deterioration in Pueblo III Canyon de Chelly need to mature to infective stage in soil. Of all the (Reinhard 2004, 2008b). We found that starvation worm type parasites that infected the Ancestral foods, including yucca leaf bases and prickly pear Puebloans, only the wireworm and hookworm had pads, became very common foods at Antelope House the potential for vertical transmission. but not so at Salmon and other Pueblo sites in Utah The number of parasite species infecting a host and New Mexico (Reinhard and Danielson 2005). population relates to their ability to combine into Pollen concentrations in coprolites reveal that stored conditions that create a disease. If only one species maize was eaten less often and in smaller quantities of parasite infects a host population, then disease at Antelope House than at Salmon Ruins, and that conditions are not likely to become a serious people at Antelope House were relying more heavily problem for that population. However, as more and on wild foods like cattail heads and horsetail stroboli. more parasites become established in the host Finally, we found that starch grains were much less population, the population will begin to experience common in the coprolites from Antelope House than disease conditions caused by the symptoms and those found at Salmon Ruin. Macrofossil remains stress from the infection of multiple parasites. show the widest variety of wild plant use at Antelope Therefore, it is important to document the diversity House. When combined, all of these data support the of parasite species found in a host population. One apparent dietary impact of environmental advantage of coprolite analyses is that they provide deterioration in the Antelope House region of the potential for identifying the number and diversity Canyon de Chelly, which undermined local of parasites that are infecting a single host (Reinhard agricultural yields and forced the residents to et al. 1988). This potential should be fully explored diversify and increase their reliance on wild plant in future coprolite work. Currently, the identification foods (Reinhard 1996, 2008b). of multiple types of parasites found in a single During that same stressful period a greater coprolite is documented only for Antelope House. diversity of parasite species infected the population The level and intensity of parasite-caused living at Antelope House. Some of these parasites, diseases in a population is affected by many like hymenolepidid tapeworms and pinworms, are conditions. These include the population’s past neither very virulent nor pathogenic but their experience with parasitic infection, nutritional and presence suggests that the population, perhaps 210 Reinhard and Bryant

already suffering from various forms of malnutrition physiological needs of pregnancy put women at was generally in poor health. Whipworms were also much higher risk. present and although this parasite is virulent, it is not When multiple infections occur in a pregnant very pathogenic. Hookworm and wireworm woman, as it most probably did at Antelope House infections were also common at Antelope House during this period of severe stress, serious during this period. Neither of these is particularly consequences result. Of the parasites at Antelope virulent, but they both have the capability of vertical House, hookworm, G. lamblia, and E. histolytica transmission from mother to fetus or newborn and were the most serious challenges to maternal and the ability to infect the most susceptible portion of infant health. In pregnant women hookworm causes the population (infants) with regard to helminthes. In severe iron deficiency anemia, nutrient addition, two species of diarrhea-causing protozoan malabsorption, alimentary bleeding, fatigue, parasites were found that are both highly virulent and diarrhea, preeclampsia, and heart failure in labor. E. pathogenic, Entomaeba histolytica and Giardia histolytica and G. lamblia cause iron deficiency lamblia. Overall, the image that emerges of the anemia, nutrient malabsorption, diarrhea, people living at Antelope House during this period is dehydration, and shock. one of a physiologically stressed, highly infected These parasites also cause problems for fetuses. population among whom some individuals in the Hookworm has a vertical transplacental infection poorest health were more susceptible to infection by mode and is associated with spontaneous abortion, many parasites. still birth, and premature birth. Infants born to The widespread infection levels indicated by infected mothers often have a low birth weight. E. the Antelope House coprolites raise other important histolytica and G. lamblia infections of the mother questions, such as the level of interaction between also result in fetal reduced growth, low birth weight, the host’s nutritional needs and parasitism and at abortion, still birth, and premature birth. what point those levels become critical or even life- The presence of these three parasites in threatening. One example of this important nutritionally stressed Puebloan populations is an relationship between nutritional needs and parasitism important factor in the prevalence of porotic is discussed by Crompton and Whitehead (1993). In hyperostosis in Ancestral Puebloan skeletal their discussion they constructed a model predicting assemblages, especially in concert with nutritional the effects of hookworm infection on a non-pregnant stress when changing environmental conditions vs. a pregnant woman. Their model predicts that resulted in reduced food and nutrient intakes hookworm infection will deplete stored iron in the (Reinhard 1992a). Long-term droughts, such as host’s body because of the hookworm’s destruction recorded in the Antelope House region resulted in an of red blood cells thus reducing their density per aggregation of human populations around the milliliter of blood. Thus the effect of this iron dwindling water sources. This in turn led to a depletion becomes much more acute in pregnant proliferation of crowd diseases and diseases women because they must also provide iron for their associated with contaminated water and inadequate growing fetus. Each hookworm in a person’s body sanitation. Bioarchaeologically, these phenomena are can consume 0.27 ml of blood per day (MacLeod expressed as elevated levels of porotic hyperostosis 1988). After only 20 weeks of hookworm infection in skeletal remains. the host may show symptoms of hypochromic and macrocytic anemia. A major problem among pregnant women is THE PATHOECOLOGY OF CHAGAS that the minor symptoms of hookworm infection are DISEASE IN THE TEXAS ARCHAIC often indistinguishable from normal complaints of pregnancy such as epigastric pain, heartburn, etc. Chagas disease, which is caused by the When the hookworm infection is more severe, the protozoan parasite Trypanosoma cruzi, has a pregnant woman may show symptoms of a low grade multifaceted pathoecology. Its basic life cycle fever, fatigue dyspnea, heart palpitations, flow involves many species of reduviid bugs in the family murmurs, and anemia. In cases of heavy infection, Triatominae -- winged insects called “kissing bugs” constipation or diarrhea, jaundice, emaciation, that act as vectors for the disease. The disease cardiac failure, or preeclampsia can occur. If an normally cycles among a wide variety of host infected woman survives labor, she often cannot animals including marsupials, edentates (such as the recover easily from post-partum hemorrhage, which armadillo), carnivores, rodents, and bats. Humans can contribute to maternal death. Overall, in most often become infected with T. cruzi when hookworm-infected populations, the increased infected triatomines (assassin bugs) emerge Pathoecology and the Future of Coprolite Studies in Bioarchaeology 211

nocturnally to feed on sleeping people (Reinhard et potential transmission of Chagas disease to humans. al. 2003; Schmidt et al. 2005). One of the most common animals killed and eaten by The symptoms of Chagas disease are diverse. these Late Archaic cultures were woodrats that could There are two stages of infection with distinct easily be caught by women and children foraging for symptoms. In about 1 percent of cases, acute other foods. The unfortunate result of these actions symptoms occur one to two weeks after infection. was that people reduced the number of normal hosts These include fever, fatigue, facial swelling around for the insects by killing woodrats and then the bite site, and enlarged lymph glands. These inadvertently constructed ideal alternative habitats in symptoms last from four to eight weeks and then the middens and debris in and around their disappear. Chronic disease develops 10 to 20 years habitations. after initial infection in about one-third of infected A bioarchaeological examination of ancient people. Cardiac problems such as cardiomegaly, skeletons recovered from the Lower Pecos region arrhythmia, and cardiac arrest are common indicators (Reinhard et al. 1989) reveals that over 40 percent of of the chronic stage of this disease. Problems with those people had abscesses. Analysis of human the digestive system, including megaesophagus and coprolite remains from these same rock shelters and megacolon, are also symptoms of the chronic stage, caves suggests that woodrats may not have been which in some cases causes death. completely cooked before being eaten (Reinhard et Recently, oral transmission of Chagas disease al. 2006c). These findings raise the possibility that has been discovered (Prata, 2001; Shikanai-Yasuda Chagas infection could have been transmitted et al. 1991). This occurs when humans eat infected through consumption of infected animals. In this food that is either contaminated with the feces of possibility, the protozoa in the blood of the woodrats infected triatomines, or they accidentally eat the may have entered the blood of the humans directly infected bugs. Either can occur inadvertently in through oral lesions from abscesses. We also know contaminated products that are processed into that bedrock mortars and grinding stones were used consumable foods. Another potential method of in the same rock shelters and caves where the transmission for Chagas disease is eating the meat of triatomines lived. This raises the possibility that the infected animals when the meat has not been fully actual insects or their feces could have been cooked. inadvertently ground and mixed into prepared foods, Bioarchaeological diagnosis of Chagas Disease resulting in infection. is based on gross pathology (Martinson et al. 2003) Megacolon is a very rare symptom of Chagas and molecular analysis (Aufderheide et al. 2004). A disease. The discovery of this one case suggests that case of “megacolon” was discovered in a mummy many more Archaic age people were infected than found in a dry rock shelter located on the Texas- previously suspected. Mexico border in the lower Pecos area west of Del Rio (Reinhard et al. 2003). The Late Archaic age mummy was about 1,000 years old and the INSECTS AND PARASITISM IN THE preliminary diagnosis of Chagas disease was verified PREHISTORIC SOUTHWEST by molecular tests (Dittmar et al. 2003). In an effort to understand why this might have happened during Some of the most intriguing pathoecological the Late Archaic period, a pathoecological data come from areas and archaeological sites where reconstruction of the individual’s lifestyle and ancient people ate the intermediate insect hosts that Chagas disease ecology was attempted. Studies of were carrying acanthocephalan - thorny-headed or the Lower Pecos region revealed that there are seven spiny-headed worms – and tapeworms. In these types of insects (tratomine vectors) that can carry cases, specific aspects of prehistoric diet or hygiene Chagas disease. Four of these live in woodrat nests resulted in their infection. and feed primarily on woodrats. The mixture of rock Acanthocephala rarely parasitize humans in the and vegetation in the woodrat nests attracts the modern world. Acanthocephala are named for their triatomines. Thus, the woodrat nest is the natural proboscis which is usually covered with spiny hooks nidus for Chagas disease transmission. Human that are often arranged in rings of horizontal rows. occupation of the rock shelters and caves in the Using this proboscis, these worms attach themselves region resulted in the accumulation of vegetal debris, to the tissues of its host. The hooks on the proboscis including grass-lined beds. Other debris that attracted pierce the intestinal wall of the host and allow the triatomines included rocks and vegetal remains from worm to derive nourishment while it also completes roasting agave and from the deposition of plant its life cycle. remains in trash. In essence, humans created artificial Acanthocephalans have complex life cycles. nidi in caves for the insects and thus increased the Adults produce eggs that are released into the 212 Reinhard and Bryant

intestines of the host and thus become part of the an infected dog. Since there is coprolite evidence that host’s feces. The feces are then eaten by an prehistoric people did eat fleas and lice, we suspect invertebrate and the dormant eggs hatch in the that the taeniids eggs found in Southwestern human intermediate host. There, the acanthocephalan worm coprolites may be from D. caninum infections. penetrates into the body cavity of the intermediate host and encysts. This then becomes the worm’s infective cystacanth stage. When the invertebrate is THE PATHOECOLOGY OF DENTAL then eaten by a vertebrate predator, such as a human, DISEASE the cystacanth develops into a mature adult in the intestine. The connection between dental disease and We know that ancient Americans were infected coprolite studies is obvious. The carbohydrates that because we have found acanthocephalan eggs in are associated with dental caries such as starch can human coprolites. These finds are most common in be recovered from coprolites in the form of starch the Great Basin from coprolites found at Archaic grains. Fruits are represented in coprolites by seeds sites including Dirty Shame Rockshelter in Oregon and epidermis. Habits that can cause dental fractures (Hall 1977), Clyde’s Cavern in Utah (Hall 1972), including cracking nuts or retouching stone tools and Danger and Hogup Caves in Utah (Fry 1977) with one’s teeth are evidenced in coprolites in the (for review see Reinhard 1990). Acanthocephalan form of nutshells and tiny chips of flint or obsidian. eggs have also been recovered from Ancestral Abrasive plant foods full of tough fibers, such as Pueblo coprolites found in sites located in Glen yucca, agave, and prickly pear cactus pads, leave Canyon, Utah, and Black Mesa Arizona (Fry 1977; behind traces of fiber and phytoliths in coprolites. Gummerman et al. 1972:191; Reinhard 1990). The Finally, grit from grinding stones is often included in Ancestral Pueblo infections resulted from accidental the resulting flour. When chewed, the grit tends to or purposeful consumption of flour beetles, camel wear down the cusps on molar teeth. Grit has been crickets, roaches, and perhaps other similar insects. found in coprolites. A different type of infection may have resulted From previous studies (Hartnady 1986; from eating ectoparasites (parasites outside the body Hartnady and Rose 1991; Turpin et al. 1986) we such as lice, fleas, and ticks). Fry (1977) and Napton know a lot about dental disease in the Lower Pecos (1969) found lice in human coprolites recovered region of Texas. Analysis of teeth from skeletal from Danger Cave, Utah and Lovelock Cave, assemblages reveal that dental caries and extreme Nevada. Recently, I (Reinhard) found a tick in a dental attrition led to abscesses and antemortem human coprolite from Antelope Cave, an Ancestral tooth loss (Turpin et al. 1986; Marks et al. 1985). Pueblo Site in northwestern Arizona near the Virgin The molars of these people tended to have smooth, River. Apparently, the tick had been bitten off and polished occlusal surfaces and rounded occlusal swallowed and then became part of a coprolite. Fry margins from chewing tough fibers and grit in their (1977) presented a case suggesting that prehistoric foods. Some people had little or no enamel left on people ate ectoparasites as a hygienic measure. This their front teeth. SEM analyses of the dentition of assumption makes good sense and we believe that Lower Pecos peoples reveals microwear in the form Fry made a good case for this type of behavior. of gouges, striations, and compression fractures in Taeniid tapeworm eggs have been found in the polished enamel surfaces. coprolites from Danger Cave, Hogup Cave, Glen Hartnady (1986) and Turpin et al. (1986) Canyon, and Antelope Cave. The exact species of completed separate studies of large samples from taeniid that is represented by these finds is unknown. ancient teeth recovered from the Lower Pecos region In the modern world, humans are most commonly spanning the full temporal range of the Archaic infected by two species, Taenia solium from pork period (8,000 B.C. to A.D. 1000). They found that and T. saginata from beef. These species of taeniids for many of those early inhabitants, by age 25 were almost certainly absent in the prehistoric New essentially all molars were lost, and by age 40 adults World because the host animals are native to the Old were virtually toothless. World. There is one other known taeniid that can Review of published dental pathology data infect humans, although it is much more common to indicates that dental caries, dental wear, abscesses, find it as a parasite of dogs. This is Dipylidium and tooth loss were much more significant problems caninum (cucumber tapeworm). D. caninum uses for Archaic peoples in the Lower Pecos region than fleas and lice as intermediate hosts. Therefore, if a for contemporaneous groups living along the Texas person ate fleas and lice, in an effort to prevent the coast, on the coastal plains, or on the Edwards ectoparasites from feeding on humans, it is possible Plateau (Reinhard et al. 1989). Several dietary they could have eaten an ectoparasite that had fed on explanations for the excessive dental problems in the Pathoecology and the Future of Coprolite Studies in Bioarchaeology 213

Lower Pecos region have been proposed. Hartnady followed by remains from Bighorn Sheep Ruin from and Rose (1991) suggest that grinding stone grit was Grand Gulch of Utah. Coprolites from Salmon Ruin the main cause, while Turpin et al. (1986) suggest in New Mexico had the lowest frequency of cactus that a diet high in plant fiber or a dietary dependence and agave phytoliths of the Puebloan samples. In on hard seeds was the primary cause. their earlier paper Reinhard and Danielson (1998) Studies of human coprolites from the Lower noted these phytolith data correlated with similar Pecos region (Bryant 1969; Danielson 1993; variations in dental wear reported from Ancestral Danielson and Reinhard 1998; Edwards 1990; Pueblo human remains. In summary, it appears that Reinhard 1988a; Sobolik 1988, 1994; Stock 1983; increased use of cactus and agave as foods is Williams-Dean 1978) reveal important reflected in increased dental pathology and in dental characteristics of prehistoric diet and shed light on wear patterns. the causes of dental pathology. Macroscopic remains During the 1970s, when one of us (Reinhard) in coprolites include fragments of shells from hard worked at Salmon Ruin, the abundance of worn out seeds and nuts including pecans, walnuts, mesquite manos and metates led the archaeologists to and yucca seeds, and hackberries. Although some of speculate that the average Salmon villager probably those were probably broken open initially with “ate the grit from seven manos and three metates stones, chewing on the hard fragments could have during his/her lifespan.” In the analysis of coprolites caused microfractures in teeth. Danielson and from Salmon Ruin it is very difficult to determine Reinhard (1998) explored one of the other potential whether or not the recovered grit came from grinding causes, the high fiber hypothesis. They discovered stones, from eating wind-blown sand, or from the high amounts of phytoliths in all of the Lower Pecos exterior of the feces what were often deposited on coprolites they examined; as much as 20 percent of sand and then sometimes covered with sandy soil. the total volume of some Lower Pecos coprolites was This problem can only be resolved through a careful composed of agave and prickly pear phytoliths. SEM study of the morphological similarity between the studies revealed that the distance between the sharp recovered coprolite grit and the composition of chisel-shaped phytoliths embedded in agave fibers various grinding stones. was the same as the distance between the parallel micro-striations observed on teeth of Lower Pecos region people (Danielson and Reinhard 1998). Those COPROLITES AND THE observations support the hypothesis that a high fiber CANNIBALISM CONTROVERSY diet including agave probably caused much of the lower Pecos dental pathology. Although we cannot Any discussion of Southwest coprolite studies be certain, it appears that grinding stone grit was not must address the world’s most notorious coprolite one of the major factors causing the high incidence which was deposited in a hearth at a small residential of dental disease in these Archaic cultures of the site in Cowboy Wash, Colorado by, some say, a region. cannibal (Billman et al. 2000; Diamond 2000; In a more recent study the same researchers Dongoske et al. 2000; Lambert et al. 2000; Marlar et (Reinhard and Danielson 2005) expanded the study al. 2000a, 2000b). These publications focus on the of phytoliths to include coprolites from other parts of discovery of disarticulated skeletons with butchery the Southwest including some hunter-gatherer sites marks, human myoglobin residue in cooking pots, in the northern Sonoran Desert, the Colorado Plateau human bone fragments polished by boiling, and the of Utah, and three Ancestral Puebloan sites in occurrence of human myoglobin in one human Arizona, New Mexico, and Utah. Phytoliths from coprolite. The coprolite was determined to be human genera and species in the cactus and agave plant based on the presence of human-linked digestive groups were as abundant in the Arizona and Utah enzymes (Marlar et al. 2000a, 2000b). This human hunter-gatherer coprolites as in the coprolites form coprolite containing the remains of human muscle Lower Pecos region sites. Although no human protein is considered the most convincing evidence skeletal remains from these hunter-gatherer sites of cannibal behavior in the Southwest. have been recovered, we suspect that if found the None of the major protagonists in the debates teeth of those people would show severe dental about Anasazi cannibalism compared the results of microwear and related dental disease, similar to the the dietary analysis of the Cowboy Wash coprolite Archaic peoples of the Lower Pecos. with any of the extensive published records of other Cactus and agave phytoliths were also present Southwestern coprolites. The problem, as I in the coprolites from Ancestral Pueblo sites. (Reinhard) noted when the coprolite was first Coprolites from Antelope House, northeastern discovered and examined, is that the Cowboy Wash Arizona, had the highest phytolith frequency coprolite is absolutely abnormal in terms of its 214 Reinhard and Bryant

content. A recent essay (Reinhard 2006) addresses was to gain some type of magical benefit from eating the central issue of what some have suggested was a a small portion of flesh; neither was an attempt to case of drought-induced cannibalism. This summary replace animal protein with human flesh! of the coprolite evidence for 10,000 years of subsistence in the American Southwest indicates that PATHOECOLOGY AND THE FUTURE neither environmental collapse nor agricultural failure were ever associated with cannibalism or OF SOUTHWESTERN increased reliance on meat. Instead, the coprolite BIOARCHAEOLOGY record documents that in times of stress the Ancestral Puebloans increased their reliance on the As demonstrated by our discussion of past wild plants which had at one time been the primary studies, the combined theoretical approach to diet of their hunter gatherer ancestors. Based on the pathoecology combined with the data and coprolite evidence, there is no reason to assume that methodologies derived from coprolite research are any of the Ancestral Puebloans would have altered essential elements for Southwestern bioarchaeology. their diet patterns at any time, even during periods of Currently, the greatest need is to refine certain areas extreme stress, to systematically rely on eating the of this research. flesh of other humans. One area of research that needs strengthening is It seems to us that the methods and conclusions the study of paleonutrition. The forte of coprolite presented by Marlar et al. (2000a, 2000b) were analysis is the potential to identify chemical, accepted as fact and reified before the method was microscopic, and macroscopic remains of past tested or proven. For such a controversial discovery, human diets. The weakness of this approach at an independent, blind study of the Cowboy Wash present is our inability to determine exactly how coprolites should have been performed. One problem much of a potential food product was actually eaten! with the original study is that a positive ELISA test What does one seed, one piece of plant fiber, one will confirm the presence of a target protein even if it leaf, or one fragment of a nut shell reflect? Does it is present in very tiny amounts. It is a qualitative test mean a person ate only one tiny portion of each of without any type of quantitative measure. Thus, even these items, or do these data mean that these meager very small amounts of a protein will result in positive fragments are only slight traces of meals composed outcomes. Previous studies using the ELISA test almost entirely of those items? Because we cannot demonstrate this aspect. For example, Gonçalves et determine the “quantity of food eaten,” we can only al. (2002, 2004) successfully used the ELISA method interpret the nutritional value of a diet qualitatively. to identify the presence of parasitic protozoa proteins For example, the range of 40 foods discovered in a in coprolites, which would occur in very tiny coprolite may include all the elements of a amounts when a human was infected. Therefore, the nutritionally sound diet. However, because we do not most pressing question that should be asked is “how know the exact amounts that were actually eaten, we much human muscle must be consumed before the cannot verify that the represented diet was really ELISA test will confirm a positive result?” Would nutritionally adequate. This is a problem that should someone need to eat one milligram, or one gram, or be addressed in future research by experimental as much as a kilogram of human muscle before the consumption of known amounts of traditional foods test would return positive results? We believe that by volunteers and the controlled recovery and this is the most important issue in this discussion. analysis of the volunteers’ feces. The method for Consumption of one milligram does not imply the conducting this type of research, as it applies only to type of rampant cannibalism that has been suggested the flow of pollen that is eaten, was first developed by some, such as Christy and Jacqueline Turner by Kelso (1977; Kelso and Solomon 2006) and later (1999) in their book Man Corn: Cannibalism and by Williams-Dean (1978; 2006). Both used modern Violence in the Prehistoric American Southwest. volunteers who ate specific diets and were able to We believe that the issue of cannibalism must record times and amounts of each component that be addressed with cautious, hypothesis-driven was eaten. research that takes into account many possibilities. If the quantification of paleodiets can be For example, one possibility comes from achieved, then the pathoecology approach can have ethnographic accounts that note the widespread use important applications to ancient and modern Native of ritualistic cannibalism. Records suggest that ritual American health. The combination paleonutritional cannibalism occurred among Native American and archaeoparasitological data will enable a more groups where warriors either ate a small portion of a nearly precise estimation of the nutritional and killed enemy, or a small portion of some departed physiological stresses experienced by the ancient relative. In both of these ritualistic cases the intent populations we study. For modern populations, this Pathoecology and the Future of Coprolite Studies in Bioarchaeology 215

approach could be very useful in addressing the abrasives (phytoliths and fibers), and processing origins of Type II diabetes in modern Southwestern dietary abrasives (grit from manos and metates) Native Americans and whether it is more closely and the causation of micro-wear and dental related to the influence of environmental (Bennett pathology. This type of study is perhaps the most 1999) or diet (Neel 1999) change. direct way where coprolite data and pathoecology Archaeoparasitology is well developed, but its are linked to bioarchaeology. It is potentially the study needs to be applied broadly over wider regions most fruitful area of research because of the wide of the American Southwest. The newest methods of variation of diets at different times and in using data from ELISA assay for protozoa parasites different environments throughout the American and broader searches for cryptic evidence of Southwest. hookworm and wire worm infection in ancient There are other potential bioarchaeological groups could provide useful clues for interpreting the applications that can be gained from studies of etiologies of osteological and dental indicators of coprolite analyses. Nevertheless, before stress including porotic hyperostosis, enamel bioarchaeologists can discover and apply these hypoplasias, growth arrest lines, and periosteal applications to answer questions, they will need to reactions. These types of stress, which are recorded become familiar with the pathoecological approach in skeletal remains, could be from physiological to coprolite analysis. Once this “marriage” has been stresses caused by forms of parasitism or bacterial achieved, questions can be asked and testable transmitted infections from the same types of entry hypotheses developed for coprolite studies. routes. In summary, we firmly believe that the field of We can make the link between dental caries, bioarchaeology must broaden its scope to include abscesses, and tooth loss and the types of foods that coprology as one of its central tools. In essence, it were eaten. What we need now is new research to must become the “host” for the further development explore the relationship between plant dietary of coprolite research.

216 Reinhard and Bryant

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