AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 82:145-163 (1990)

Archaeoparasitology in North America KARL J. REINHARD Department of Anthropology, University of Nebraska, Lincoln, Nebraska 68588

KEY WORDS

Paleoparasitology, Coprolites, Human parasites

ABSTRACT The study of prehistoric parasitism through analysis of coprolites, mummies, skeletons, and latrine soils is rapidly growing. Its development in North America is interdisciplinary and is derived from the fields of physical anthropology, parasitology, and archaeology. The various parasite finds from North America are reviewed. The data show that prehistoric peoples in North America suffered from a variety of parasitic diseases. The validity of the findings are then considered. Although most finds of parasites from prehistoric contexts result from human infections, some finds cannot be verified as such. However, in combination with data from South America, it is clear that prehistoric peoples in the Americas were host to a variety of human parasites, some of which were not previously thought to be present before historic times. McClary, 1972; Moore et al., 1969, 1974; Zimmerman, 1980;Zimmerman and Aufderheide, 1984) or by anthropologists with training in parasitology and in consultation with parasitologists (Fry, 1970, 1974, 1977, 1980,1985; Fry and Hall, 1969,1975,1986; Hall, 1972, 1977; Williams, 1985). In contrast, paleoparasitological research in South America is done almost solely by pathologists and parasitologists (Allison et al., 1974; Araujo et al., 1981,1983; Confalonieri et al., 1985, 1988; Dalton e t al., 1974; Ferreira et al., 1980,1983a,b, 1984,1987,1988; Horne, 1985).As a result, diagnosis is more rigorous in South America, but in North America parasite data are more often placed in a cultural context. Most parasitological study falls into the realm of coprolite analysis, which is largely a development of North American archaeology (Bryant, 1974a,b, 1986; Bryant and Williams-Dean, 1975; Fry, 1985; Shafer and THE NATURE OF PARASITOLOGICAL RESEARCH Bryant, 1977). Consequently, parasitologiThe impetus for study of parasitism in cal studies are typically integrated with diNorth America has come from archaeology etary data (e.g., Fry, 1977; Fry and Hall, and physical anthropology. The actual exam- 1986; Hall, 1977; Reinhard, 1985a, 1988a; ination of archaeological specimens such as Reinhard et al., 1987; Stiger, 1977).Funding coprolites (desiccated feces), latrine soils, and basic support for such studies come from and mummies is typically done by parasitol- interested archaeologists who often publish ogists or pathologists (Allison et al., 1974; Dunn and Watkins, 1970; Dusseau and Porter, 1974; Hevly et al., 1979; Horne, 1985; Received December 27,1988; accepted May 16,1989. Prehistoric parasitism is a topic of intensifying interest and research. Productive researchers work in many localities in Europe and the Americas. The efforts of these individuals are gradually making an impact on the fields of both parasitology and anthropology. As “paleoparasitological” data are generated and incorporated into these fields, one must emphasize the interpretive limitations inherent in paleoparasitologxal research. This will enable scholars not immediately involved in parasitological analysis to evaluate parasite data sensibly. Presented below is a survey of parasitological studies in North America. The survey is followed by a critique section emphasizing the interpretive pitfalls of the growing data base and assessment of the antiquity of parasitism in North America. The role of such data in physical anthropology is then discussed.

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the parasite data. Thus, in North America the study of archaeological parasitism is jointly derived from the fields of anthropology and parasitology and is largely sponsored by archaeology. The development of the field, its unique relation to both parasitology and anthropology (especiallyparasite ecology and paleopathology), and derivation of specialized analysis techniques (Araujo et al., 1981; Confalonieri et al., 1985, 1988; Fry, 1980; Jones, 1985, 1988; Jones et al., 1988; Reinhard et al., 1986, 1987, 1988) warrant a specific designation of archaeological parasite study. The term paleoparasitology has been applied to this emerging field. As introduced by Araujo et al. (1981),paleoparasitology is defined as an extension of paleopathology, which is the study of ancient disease. The term is gaining acceptance in North America (Reinhard, 1988a; Reinhard et al., 1987). As a matter of opinion, one might object to the application of this term. For New World archaeologists, paleo refers specifically to megafauna hunting cultures that existed up until 9,000 years ago. In genera1,paleo refers to ancient forms or conditions. In North America, the examination of parasite evidence from archaeological sites includes ancient materials (Fry and Moore, 1969; Moore et al., 1969)and recent materials dating into historic times (Reinhard et al., 1986).For the historical material, paleoparasitology is a misnomer, falling out of the range of what is normally considered ancient. Most prehistoric studies have been carried out with materials post-dating New World Upper Paleolithic times. Perhaps archaeoparasitology is a more descriptive term. The term is more general and includes studies of both ancient and recent archaeological remains. It does not imply any specific cultural manifestation. Consequently, for the remainder of this paper I will use the term archaeoparasitology. In the study of archaeoparasitology, research is limited to the helminth and arthropod parasites. Helminths include trematodes (flukes), cestodes (tapeworms), acanthocephalans (thorny-headed worms), and nematodes (roundworms). Helminths are represented by durable reproductive products such as eggs and larvae. Nematode remains also include adult worms. The tough cuticle that surrounds nematode adults allows for their preservation. The tegument of adult cestodes, trematodes, and acantho-

cephalans is too delicate to permit preservation of adult forms in any environmental condition. The most common arthropod remains found in archaeological contexts are lice. These are represented by eggs cemented to hair on mummies or adults from coprolites. The preservation of protozoa has not yet been demonstrated in any North American study. Continued research may eventually reveal techniques that can be used to identify protozoa. In North America, coprolites have been the main focus of archaeoparasitology. Coprolite analysis is historically aimed at the recovery of dietary and ecological data (Bryant, 1974b; Fry, 1980). The techniques of coprolite dietary analysis were first devised by Callen (1967) and Callen and Cameron (1960). Since then, coprolite analysis techniques have been refined by researchers in the Great Basin (Fry, 1977; Hall, 1972,1977; Heizer, 1967; Heizer and Napton, 19691, on the Colorado Plateau (Hevly et al., 1979; Reinhard, 1985a+; Reinhard and Clary, 1986; Reinhard et al., 1987; Stiger, 19771, and in western Texas (Bryant, 1974a,b, 1986;BryantandWilliams-Dean, 1975;Reinhard, 1988b; Shafer and Bryant, 1977; Stock, 1983). For specific application to coprolites, parasitological techniques were devised by Callen and Cameron (1960) and refined by Samuels (19651, Hall (19721, Fry (1977, 1980), and Araujo et al. (19811, as reviewed by Horne (1985). Refinement of technique as applied to coprolites continues jointly between North American and South American researchers (Araujo et al., 1981; Confalonieri et al., 1985; Ferreira et al., 1983a; Reinhard, 1985b,c, 1985b; Reinhard et al., 1987,1988). Mummies are also a source of archaeoparasitological data in North America, especially with regard to arthropod parasitism. However, mummies have not been studied as intensively in North America as in South America, Europe, or Egypt (Cockburn and Cockburn, 1980; Ferreira et al., 1983a). In the future, mummy analysis will play a more important role in North American archaeoparasitology. The study of latrines and soils from cultural deposits has long been an important source of archaeoparasite data in Europe (Gooch, 1983; Herrmann, 1986,1987; Herrmann and Schultz, 1986; Jones, 1985,1988; Jones et al., 1988; Moore, 1981; Pike, 1967, 1975; Taylor, 1955). Relatively few latrine studies have been done in North America,

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although latrines have provided both prehistoric (Hevly et al., 1979) and historic (Reinhard et al., 1986) evidence of parasitism. Such study is becoming increasingly interesting to historical archaeologists. Preservation of remains varies. Coprolites from caves are excellent for the preservation of helminth eggs and larvae (Dusseau and Porter, 1974; Fry, 1977; Reinhard, 1985c; Reinhard et al., 1987). Coprolites from open sites are less well preserved, and parasite eggs within such coprolites can be partially decomposed. The poor preservation of Enterobius vermicularis eggs is specifically noted for fecal remains excavated from open sites (Reinhard and Clary, 1986; Reinhard et al., 1987, 1988). Latrine sites provide suitable conditions for preservation of more durable eggs, but fragile eggs are susceptible to decomposition in latrine environments. This is especiallytrue ofoxyurid eggs (the nematode order including piworms). Mummies provide suitable conditions for helminth preservation, especially if frozen in prehistory (Zimmerman, 1980; Zimmerman and Aufderheide, 1984)or rapidly desiccated (El-Najjar and Molinski, 1980; El-Najjar et al., 1980). Rarely, skeletal analysis reveals evidence of parasitic disease. This occurs exclusively in the form of calcified tapeworm cysts (Ortner and Putschar, 1981; Williams, 1985). Presented below is a summary of North American archaeoparasitology by geographic region. Northern Mexico and the United States are the main foci of research in North America by U S . and Canadian analysts. Since the integration of dietary data and parasite data is the hallmark of North American archaeoparasitology, brief summaries of diet are included. NORTH AMERICAN ARCHAEOPARASITOLOGY

The Arctic and Subarctic To date, the only evidence of prehistoric parasitism from the far north (Fig. 1)comes from the analysis of mummies and skeletons. Feces have been recovered in the permafrost of some sites (R. Holloway, Eastern New Mexico University, personal communication) and show that the potential of parasitological analysis from frozen feces does exist. An interesting case of false parasitism is reported by Zimmerman (1980). False parasitism refers to the find of eg s of a parasite species in a host not susce ti le to infection by that parasite species. n humans, false parasitism occurs when eggs of a parasite

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noninfective to humans are consumed with foods contaminated with the eggs. A frozen mummy dating to about A.D. 400 was recovered on St. Lawrence Island, Alaska (Fig. 1). The eggs of Cryptocotyle lingua (a fluke infective to fish) were found in the colon contents of the mummy. Zimmerman notes that Rausch et al. (1967) report the eggs of this trematode in modern Eskimos, but that true infections have not been found in humans. Instead the eggs were introduced into the human digestive tract by consumption of the fish definitive host. Archaeological evidence indicates that the prehistoric inhabitants of the island subsisted largely by hunting marine mammals. The find of C. lingua demonstrates that fish were also incorporated in the prehistoric diet. Zimmerman also reports the analysis of a mummy recovered from the Aleutians, but notes that parasite examination was negative. At Utqiagvik, near Pt. Barrow, Alaska, an ice surge covered and collapsed an Eskimo winter house about A.D. 1550, killing two women inside. The frozen mummies were recovered in excavations in 1982 and analyzed by Zimmerman and Aufderheide (1984).Cysts resembling those of Trichinella spiralis (the nematode responsible for trichinosis) were observed in diaphragm tissue in one of the women. Unfortunately, the cysts were not sufficiently preserved for definitive identification. The inhabitants of the village subsisted largely on marine mammals, caribou, and migratory birds. Both marine mammals and caribou are modern reservoirs for human Trichinella infection. Trichinosis is a zoonosis, a disease of animals that is transmissible to humans. Another dangerous zoonosis is hydatid cyst disease, which has been reported in the analysis of a female skeleton excavated from Kodiak Island, Alaska (Ortner and Putschar, 1981:232-233). The skeleton predates Russian contact, but a more precise date is not available. This disease is very serious and is caused by the larval stage of the tapeworm Echinococcus, which forms large cysts in somatic tissue. Evidence of the disease consists of calcified cysts excavated with skeletons. Kodiak Island is outside of the modern range of Ech. multilocularis (Rausch, 1958), but Ech. granulosus could have been the infective organism. Human infection often results from close association with dogs. From the archaeological perspective, the find of a cyst in a single skeleton signals infection of many more individuals,

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1. Utqiagvik, Alaska 2. St. Lawrence Island, Alaska 3. Kodiak Island, Alaska 4. 32SN22, North Dakota 5. Schullz Site, Michigan 6. Dirty Shame Rockshelter, Oregon 7. Queen Anne Square, Rhode Island 8. Greenwich Village, New York 9. Upper Salts Cave, Kentucky 10. Daws Island, South Carolina 11. Colonial Williamsburg, Virginia 12. Rio Zape, Durango 13. Frightful Cave, Coahuila I10

I20

I Ill

I

I

I00

VII

1 XI1

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Fig. 1. Site location map ofarchaeoparasitefinds outside ofthe southwesternUnited States.

since only 2% of individuals infected exhibit calcified cysts, and, of these, there is a possibility that some cysts are lost in excavation. The boreal United States Hydatid cyst disease is also reported by Williams (1985) in a female skeleton excavated in North Dakota dating to about A.D.

600. As in the case from Kodiak Island, evidence of hydatid cyst disease is present in the form of a calcified cyst. The cyst is spheroidal and measures 25 mm in diameter. Williams implicates Ech. granulosus, presently endemic to North Dakota. Dietary data are not presented for the site. This is the only intact hydatid cyst thus far

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recovered from a burial to date. It presented the chance of verifying Williams’ diagnosis by examining the contents of the cyst for remains of the durable hooks of the larvae. Unfortunately, it was necessary to return the skeleton to Native Americans for reburial before this could be explored. The modern parasitological literature suggests that the introduction of Echinococcus into the northern Plains is a development of this century. Williams’ find suggests that the genus has some antiquity in the area. Microscopic examination could have definitively verified Williams’ find and therefore modified the view of Echinococcus biogeography. Diphyllobothriasis is yet another zoonosis. Eggs of what may be Diphyllobothrium were recovered from one of 20 prehistoric coprolites excavated from the Schultz site, Michigan (McClary, 1972).The eggs are suggested to be those of D. latum (see critique of Archaeoparasite Finds, below). Diphyllobothrium infects humans who eat raw fish. The coprolites date to the Late Middle Woodlands Period, a prehistoric time when agriculture began to augment a diet based on fishing, hunting, and collection of wild plants. In the case of this analysis, it was not possible to determine whether the coprolites were of human or canid origin. Taeniid tapeworm eggs (family Taeniidae) were also found in one coprolite. Assuming that this coprolite is of canid origin, McClary suggests that the eggs are of Echinococcus or Taenia. The analysis of historic privies excavated in Queen Anne Square, Newport, Rhode Island (Reinhard et al., 19861, provides evidence of parasitism. Historic documentation and archaeological study of three privies shows that one was used by a wealthy merchant who served as a captain in the militia immediately before the Revolutionary War (Mrozowski, 1981, 1983, 1984). Two other privies were used by families of the poorer artisan class. One of these was used by a blacksmiths family before the Revolution, and the other was used by a butcher’s family immediately after the Revolution. The find of Trichuris trichiura and Ascaris lumbricoides in all privies documents parasitism of both of colonial Newport’s social classes. More recently, the author undertook an examination of latrine soils from Greenwich Village, New York. The soils date to the early 19th century. Eggs corresponding in size and shape to Trichur. trichiura were found, but Asc. lumbricoides eggs were absent from the soils.

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The Great Basin and Mojave Desert The Great Basin encompasses a large geographical area that includes Nevada and parts of Utah, California, and Oregon. Our knowledge of prehistoric parasitism in the region comes primarily from the analysis of coprolites recovered from dry caves. Caves in Utah and Nevada have been most extensively excavated, and two prehistoric hunter-gatherer adaptation types are recognized. These are Desertic Adaption and Lacustrine Archaic Adaptation (Fry, 19801, each with a different archaeoparasite assemblage. The Desertic Adaption subsistence strategy was focused in the deserts of Utah and is represented by the excavations of Hogup and Danger Caves (Fig. 2). These caves have a combined time depth of 10,000 years. The ecology of the area is typified by a restricted flora and fauna. Consequently, the prehistoric diet is restricted to 11 plant species, with greatest reliance on Allenrolfea (pickleweed or burroweed) and Opuntia (prickly pear cactus). Insects, reptiles, and small mammals were also eaten. The Lacustrine Archaic Adaptation subsistence strategy typifies prehistoric huntergatherer peoples in Nevada who lived in lake-side habitats. Coprolites from four caves have been excavated and analyzed (Roust, 1967). The most important of these caves is Lovelock Cave (Ambro, 1967; Heizer, 1967; Heizer and Napton, 1969; Fry, 1980). The fauna and flora of that area is richer than that of the Utah desert, and 19 plant species were used, the most important of which were Typha (cattail), Elymus (wild rye), and Scirpus (bullrush). Mollusks, fish, waterfowl, and rabbits were included in the diet, but fish, ducks, and mudhens were the most important prehistoric animal foods. Occupation of the caves began at about 2000 B.C. and lasted to A.D. 1800. The coprolites from Lovelock Cave date between about 500 B.C. and A.D. 1150. Archaeoparasite investigations of Great Basin coprolites show a pronounced difference in helminthiasis between the Desertic Adaption and the Lacustrine Adaption. Fry (1977, 1980) found that six of 46 Danger Cave coprolites and two of 50 Hogup Cave coprolites contained eggs of what is possibly Moniliformis clarki, an acanthocephalan (Fry and Hall, 1969; Moore et al., 1969). In addition, one Danger Cave coprolite and four Hogup Cave coprolites contained eggs ofEnt. vermicularis (Fry and Moore, 1969). Taeniid

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Fig. 2. Sites in the Southwestern United States that have been the focus of archaeoparasite studies.

tapeworm eggs were present in one Danger Cave coprolite and five Hogup Cave coprolites (Fry, 1977). In contrast, few coprolites from Lovelock Cave contained helminth eggs. A fasciolid fluke egg was recovered from one of 50 coprolites (Dunn and Watkins, 1970).Some species of these flukes are infective to humans and utilize snails as intermediate hosts. Another coprolite contained Rhabditis larvae (Heizer, 1967). This genus is nonparasitic and inhabits fecal material. A third coprolite contained Charcot-Leyden crystals, which are associated with, but not specific to, amoebic dysentery (Napton, 1969). Louse nits (Pediculus humanus) were found in coprolites from Danger Cave (Fry, 1977) and Lovelock Cave (Napton, 1969). Louse parasitism occurred in both subsistence types. Helminthiasis was much more common among the Utah desert hunter-gatherers than among those subsisting along the lake

shores in Nevada. Amoebic dysentery and exposure to flukes are common in moist environments. The thorny-headed worm infections in the desert areas were probably related to the consumption of insects (see Critique of Archaeoparasite Finds, below). Pinworm infection is associated with cramped living conditions and poor personal hygiene. Thus, parasitism in the two areas can be related directly to prehistoric lifestyle in different environments of the Great Basin. From southeast Oregon (Hall, 19771, eggs of Ent. vermicularis (the human pinworm) were recovered from one of 13coprolites from Dirty Shame Rockshelter (Fig. 1). This coprolite is 5,900 to 6,300 years old. Five coprolites contain acanthocephalan eggs, tentatively identified as M . clarki. These coprolites are from all occupational levels of the cave, which span a time range of 9,500 to 1,400 years ago. The shelter was occupied in prehistoric times by hunter-gatherers as in-

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dicated by Hall’s dietary analysis. Plant foods include Allium (onion), Artemisia (sage), Chenopodium fremonti (goosefoot), Opuntia (prickly pear), and Helianthus (sunflower). Animal foods include rodents, crayfish, mollusks, and insects. Recently, archaic hunter-gatherer coprolites were excavated from the eastern margin of the Mojave Desert near the east-central border of Arizona at Bighorn Cave (Fig. 2). I examined 35 coprolites from the site, which were excavated from several levels dating from about 200 B.C. to A.D. 400. Dietary analysis of the coprolites demonstrates that Prosopis pubescens (screwbean mesquite), Descurania seeds (mustard), Opuntza (prickly pear pads), S a l k (willow catkins), Yucca, and Ephedra (Mormon tea) were the main plants consumed, although several other species are present in minor quantities. None of the 35 coprolites showed any evidence of helminth or arthropod parasitism. The Colorado Plateau The Colorado Plateau (Fig. 2) is a highelevation area that includes portions of Utah, Colorado, Arizona, and New Mexico. Despite intensive archaeological investigation, coprolites from only one Archaic hunter-gatherer site have been analyzed parasitologically. The site is known as Dust Devil Cave (Ambler, 1984; Lindsay et al., 1968; Reinhard, 1985a, 1988a; Reinhard et al., 1985, 1987) and was intermittently inhabited between 6800 B.C. and 4600 B.C. Dietary analysis of 100 coprolites shows that the inhabitants of the cave subsisted on a variety of plant foods but with special emphasis on Opuntia (prickly pear pads), Chenopodium (goosefoot seed), Yucca, Helianthus (sunflower achenes), and Sporobolis (dropseed grass seeds and influorescences). Sylvilagus (cottontail rabbit) was a primary animal food, although large mammals and rodents were also eaten. The analysis of the coprolites revealed one strongylate egg, which is considered to be a case of false parasitism (Reinhard et al., 1985). The term strongylate refers t o a wide range of nematode parasites of the Strongyloidea and Trichostrongyloidea that produce eggs that are not readily discernible to family, genus, or species. The lack of helminth remains at Dust Devil Cave contrasted with the parasite finds among prehistoric Great Basin huntergathers (Fry, 1980) and Colorado Plateau

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agriculturalists (Samuels, 1965; Stiger, 1977; Hevly et al., 1979; Fry, 1977; Fry and Hall, 1975; Hall, 1972). Reinhard et al. (1985) proposed that the consumption of anthelminthic varieties of Chenopodium, combined with small population size and mobility, limited parasitism at Dust Devil Cave in prehistory. The use of Chenopodium species in prehistory as an anthelminthic was first suggested by Callen and Cameron (1960) and later by Hall (1977). Such use is inferred from the study of Aztec texts (Ortiz de Montellano, 1975) and is further discussed by Reinhard (1988a). Since the publication of the Dust Devil Cave parasite analysis, additional huntergatherer sites have been analyzed, including Bighorn Cave discussed above and Hinds and Baker caves discussed below. These sites show little or no parasite infection, and Chenopodium occurs as a very minor dietary component. This suggests that other aspects of hunter-gatherer life were more important in limiting parasitism among hunter-gatherers than presence of dietary anthelminthics. These other factors include small band size, seasonal movement, and diffuse population size as reviewed by Reinhard (1988a). Maize agriculture was introduced to the ColoradoPlateau about 2,000 years ago. Several cultures emerged on the Colorado Plateau, including the Anasazi, Fremont, and Sinagua. The Fremont culture existed in south-central Utah. The Anasazi were widespread on the Plateau, including the southeast corner of Utah. The Sinagua lived in north-central Arizona. Parasite analyses of Fremont coprolites are presented by Fry (1977, 1980) and Hall (1972). Hall reports on Clyde’s Cavern in east-central Utah. Of 25 coprolites, four contained eggs of Ent. uermicularis, two contained eggs of an unknown acanthocephalan species, one contained larvae of what is probably Strongyloides (hairworm), one contained the embryonated eggs of an unknown nematode, and one contained a fragment of an adult nematode. (Dr. A.W. Grundman identified the helminth remains for Hall.) The acanthocephalan eggs were not identified beyond order level. The photographs and micrometer measurements indicate that two species are present. Human infection is suggested as a possibility, although Hall mentions that false parasitism is an alternate source of the eggs. The identification of Strongyloides was based in part on the morphology of “rhabdi-

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t o i d larvae, specifically, the morphology of the esophagus. The identification is supported by the statement that “Grundman doubts that Rhabditis could have been present in the cave” (Hall, 1972:37). Hall emphasizes that the identification of this worm is only probable. Ten Fremont coprolites from five sites near Glen Canyon were analyzed for parasite remains. One of these contained taeniid tapeworm eggs. Anasazi and Sinagua peoples lived in a variety of habitats on the Colorado Plateau. In these habitats they carried out maize agriculture and foraged for wild plants and animals. Some habitats were decidedly moister than others. For example, the dietary analyses of Antelope House and Inscription House show that foraging in mesic and aquatic environments was common (Fry and Hall, 1986: Reinhard, 1988a). However,

at Salmon Ruin, Mesa Verde, and Glen Canyon, foraging was carried out in xeric pinyon or juniper woodlands. Too many Anasazi sites have been studied to treat each site individually. The helminthological finds from Anasazi sites are presented in Table 1. There is variability in parasite diversity and in the prevalence of helminth eggs in coprolites between sites. Not included in Table 1is Elden Pueblo in northern Arizona (Hevly et al., 1979; Reinhard et al., 1987). This is an open site at which several rooms were used as latrines. Individual coprolites were not preserved in the rooms. Fecal debris was represented by a dense organic strata on top of the room floors. Examination of soil samples from the fecal layers revealed Trichur. trichiura, Asc. lurnbricoides, Ent. vermicularis, hymenolepidid tapeworm, and taeniid tapeworm eggs. It is of significance that hymenolepidid

T A B L E 1. Parasite f i n d s f r o m Anasazi coprolites‘

Site name with number of coprolites studied Human coprolites Antelope House (n = 180) (Reinhard, 1988b) Antelope House (n = 49) (Reinhard et al., 1987) Antelope House (n = 91) (Fry and Hall, 1986) Bighorn Sheep Ruin (n = 20) (Gardner and Clary, 1987) Glen Canyon (n = 30) (Fry, 1977) (Moore et al., 1974) Hoy House, Mesa Verde (n = 56) (Stiger, 1977) Inscription House (n - 17) (Fry, after Horne, 1985) Kin Kletso, Chaco Canyon (n = 5) (Reinhard and Clary, 1986) Peublo Alto, Chaco Canyon (n = 2) Peublo Bonito, Chaco Canyon (n = 13) (Reinhard and Clary, 1986) Salmon Ruin (n = 112) (Reinhard et al., 1987) Step House, Mesa Verde (n = 20) (Samuels, 1965) Turkey Pen Cave (n = 24) (Reinhard et al., 1987) Canid coprolites Antelope House (n = 13) (Reinhard, 1985) Bighorn Sheep Ruin (n = 1) Turkey Pen Cave (n = 1) ‘The three notations for Antelope House represent three separate coprolite samples.

No. of coprolites positive for specified taxa

44 Enterobius uermicularis 2 Strongyloides sp. 4 Strongylate worms 9 Enterobius uermicularis 1 Strongyloides sp. 1 Strongylate worm 1 Hymenolepidid cestode 14 Enterobius uermicularis 8 Rhabditoid (?) larvae

2 Enterobius uermicularis 2 Moniliformis clarki 3 Taeniid cestode 1 Unidentified trematode 4 Enterobius uermicularis 3 Enterobius uermicularis 1 Unidentified nematode egg 1 Unidentified nematode larvae Negative Negative 4 Enterobius uermicularis

9 Enterobius uermicularis 1 Enterobius uermicularis

7 Enterobius uermicularis 2 Strongyloides stercoralis 1 Toxascaris sp. Negative

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eggs appear at one Anasazi site, Antelope House, and a t the one Sinagua site, Elden Pueblo. Rodents are usually the definitive hosts for hymenolepidid species infective to humans, and grain beetles are the typical intermediate hosts. It is probable that grain stores attracted grain beetles and rodents, which resulted in the cycling of hymenolepidids infective to humans (Reinhard et al., 1987).From this perspective, hymenolepidid infection of Anasazi is considered zoonotic. It is also of interest that Asc. lumbricoides (giant intestinal roundworm or maw worm) and Trichur. trichiura (whipworm) make their first appearance in Anasazi agricultural sites. The anal-oral life cycle of these parasites suggests that fecal contamination of agricultural villages occurred. Accepting that Strongyloides is correctly identified, its appearance with strongylate worms and Trichur. trichiuru indicates that agricultural peoples were in frequent contact with moist environments, possibly through irrigation. Acanthocephalan eggs are present in Anasazi sites in southern Utah. One Anasazi coprolite from Glen Canyon contained a fluke egg (Moore et al., 1974). It is probable that this is a case of false parasitism. In Fry’s study of 30 Glen Canyon coprolites (1977),two contained taeniid tapeworm eggs and two contained probable M. clarki eggs. Acanthocephalan eggs from Black Mesa, Arizona, have been found in a coprolite found on the floor of a pithouse (Gummerman et al., 1972:191).The eggs are not identified t o genus or species. Gummerman et al. (1972) note that acanthocephalans are “extremely infectious” and may have been a “severe problem” in prehistory. In actuality, the status of acanthocephalans as parasites of humans is uncertain, and if they did infect humans, they infected only those who happened to eat a parasitized insect. In this light, it appears that the health inferences from the Black Mesa acanthocephalan find are exaggerated. The Southeastern United States Only two prehistoric sites in the southeastern United States have been studied parasitologically (Fig. 1). Upper Salts Cave in Kentucky was the focus of two parasitological studies (Fry, 1974; Dusseau and Porter, 1974). Eight radiocarbon dates indicate that the area was used from 1125 to 290 B.C., and a span from 620 to 290 B.C. is derived from three dates based on coprolites (Watson,

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1974:235-236). Fry reports the find of Asc. lumbricoides in one of eight coprolites recovered from the cave. Dusseau and Porter report finding a larva resembling the infective stage of hookworm or Strongyloides. A burial site on Daws Island off the coast of South Carolina is the focus of the other analysis (Rathbun et al., 1980). A coprolite was recovered from one of the burials and represents the intestinal contents of the burial. The authors report that the coprolite was mineralized and was rehydrated prior to identification. Then “smears” were examined microscopically. They noted the find of nematode adults. The nematodes were morphologically consistent with hookworm adults. However, they note that “species could not be identified even after consultation with a parasitologist.” Unfortunately, photographs of the worn are not available because of poor preservation (Ted Rathbun, personal communication). Recently the author completed an analysis of a historic latrine dating to A.D. 1720 from Colonial Williamsburg. This is the earliest Colonial latrine examined from North America. The fecal layer contained 1,200 eggs per gram of soil. Of these eggs, 83%are identical to the eggs of the human whipworm Trichur. trichiura. The remainder are similar to the eggs of the maw worm Asc. lumbricoides. Mexico and West Texas Although the Tehuacan Valley of central Mexico was the site of the first detailed coprolite analysis (Callen, 19671, parasitological analysis has not been done with Tehuacan coprolites. However, coprolites from Rio Zape in Durango, Mexico, were analyzed recently (Reinhard et al., 1989). The coprolites are from a cave excavated by Brooks et al. (1962). The dietary analysis indicates that maize, Agave, Chenopodium, Helianthus, and Physalis (ground cherry) were the main plant foods at the site, which dates to about A.D. 600, Animal foods included fish, lizards and other small animals. Strongylate eggs were present in one coprolite, and Ent. vermicularis eggs were present in another coprolite. A third coprolite contained an adult louse Ped. humanus. A total of 25 coprolites have been analyzed from the cave to date. Fry (n.d.1studied 32 hunter-gatherer coprolites from Frightful Cave in Coahuilla, Mexico (Fig. 1).These dated from as early as 7500 B.C. to A.D. 300. No evidence of parasitism was found.

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The Lower Pecos of Texas (Fig. 2) was occupied by hunter-gatherers in prehistory, and coprolites have been excavated from contexts as old as 7000 B.P. (Shafer and Bryant, 1977;Williams-Dean, 1978).The coprolites reflect a dependence throughout prehistory on desert succulents, the most important of which are cactus of several species, Yucca, wild grass, and Agave. Animal foods include rabbits, rodents, fish, lizards, and grasshoppers. Williams-Dean (1978) studied 13 coprolites dating to approximately 3700 B.C. and found no evidence of parasitism. Reinhard (1988b) undertook a larger study of 57 coprolites from two Lower Pecos caves (Hinds Cave and Baker Cave) from various contexts dating from 2500 B.C. to A.D. 100 and found no evidence of parasitism. In a current study of 46 Hinds Cave coprolites dating from 2100 to 600 B.C., one contained eggs of Ent. uermicularis (Edwards and Reinhard, current research). Thus, of 116 coprolites studied from the Lower Pecos region of Texas, only one contained evidence of parasitism. Examination of a mummy from Granado Cave in the Rustler Hills of west Texas reveals possible evidence of parasitism. A section of small intestine was examined microscopically, and a small nematode was found in the intestinal mucosa (D.L. Hamilton and N. Dronen, personal communication). Unfortunately, the nematode was not photographed and was eventually lost. Arthropod parasitism Zizner (1934) presents the earliest published summary of prehistoric arthropod parasitism in North America. He notes the find of lice and nits in North American mummies from the southwest United States and also from the Aleutians. The precise locations of the sites where the mummies were excavated are not given, and he does not mention the number of mummies studied. He does note that the morphology of the lice recovered from North America is different from that of lice recovered from Peruvian mummies, although the specifics are not mentioned. Andrews (1977) makes brief mention of lice found on mummified human remains and notes that the lice of Amerindian, Arctic, and mongoloid Asian peoples are morphologically similar. A study of lice from mummies is being carried out by Walter H. Birkby, Human Identification Laboratory, Arizona State Museum. Birkby is examining mummies

from various parts of the southwestern United States. Preliminary data from this study are published by El-Najjar and Molinski (1980). Birkby has examined 18 mummies from five caves. Four Caves are located in Arizona: two on the Colorado Plateau (n = 51, one in the Upper Sonoran Desert (n = 31, and one in the Lower Sonoran Desert (n = 8). The fifth cave is located in west Texas (n = 2). Two mummies from the Colorado Plateau exhibit nits and six from the Lower Sonoran Desert exhibit nits. No adult lice were found. Lice are more likely to be recovered than any other arthropod parasite, because the eggs are cemented to the host’s hair shafts rather than being deposited in the environment. Consequently, the study of mummies is an optimal way of recovering evidence of louse parasitism. Coprolites have also yielded evidence of louse parasitism. Lice are apparently introduced by grooming practices that include consumption of the lice after removal from the head. A nit was found in one coprolite from Danger Cave dating to about A.D. 20. A louse was recovered from one Lovelock Cave coprolite and one Rio Zape coprolite. CRITIQUE OF ARCHAEOPARASITE FINDS

With some archaeoparasite finds, it is difficult to determine whether human infection was involved. For this reason, archaeoparasitologists must be very careful in documenting their finds. A critical, albeit mundane, consideration is whether the coprolites under study were deposited by humans or animals. This problem has been a focus of coprolite research, and criteria for determining probable human origin are presented by several authors (Fry, 1977,1985; Bryant, 1974b; Bryant and Williams-Dean, 1975; Moore et al., 1974; Reinhard, 1985a). Examination of morphology, dietary contents, and other fecal constituents, the reaction of coprolites with rehydration solution, and hair analysis indicate human versus nonhuman origin. The details of determining the origin of coprolites are extensive, and I refer the interested reader to the references above. It is relatively easy to separate human from most nonhuman coprolites. However, it is sometimes difficult to separate human from dog feces, a problem encountered by McClary (1972). One way of empirically determining whether a coprolite is of human origin is based on the finding of humanspecific parasites such as Ent. uermicularis.

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Coprolites excavated from human burials or mummies can be safely assumed to be human. Co rolites excavated from latrine areas are a so probably of human origin. The finding of human-specific parasites in a coprolite leaves no doubt that a true infection occurred. For example, the production of Ent. uermicularis eggs can only result from true human infections. Similarly, the finding of Trichur. trichiura and Asc. lumbricoides eg s indicates human infection. However, in t e Old World specimens, confusion with the eggs of swine trichurids and ascarids with those of humans can occur (Gooch, 1983; Taylor, 1955). The absence of swine and cattle in the prehistoric New World and the demonstrated utility of micrometer measurement in the species identification of Trichur. trichiura from coprolites (Confalonieri et al., 1985,1988) allows for the safe identification of these species in the prehistoric New World. Some of the species reported from the prehistoric New World are traditionally thought to have been introduced into the Americas with historic European colonization. When such finds are made, the archaeoparasitologist must be especially careful to present unambiguous support for diagnoses. Such support is presented by researchers for prehistoric South American ancylostomid (hookworm) infection (Allison et al., 1974; Araujo et al., 1988; Dalton et al., 1974; Ferreira et al., 1983a,b, 1987, 1988). The combined analyses of these researchers resulted in the description of ancylostomid adults, larvae, and eggs. Similarly, the approach to identification of Trichur. trichiura eggs in South America has been rigorous and includes experimental dehydration and rehydration of eggs to determine the feasibility of using egg dimensions in archaeoparasitological diagnosis (Confalonieri et al., 1985, 1988). This experimental work is directly applicable to finds of Trichur. trichiura in North America. The two parasitological analyses from the southeast United States suggest the presence of helminth species generally thought to be historic introductions to North America. Consequently, they are worthy of note. Dusseau and Porter (197459) describe the analysis of 13 coprolites, of which one contained unidentified larvae. They note that the larvae might be “infectivelarvae of either Strongyloides or a hookworm. Available evidence indicates that all human hookworms now present in the Western Hemisphere

Y

a

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were introduced from Europe and Africa in post-Columbian times. . . . [Ilt would be rash to postulate that this presumed nematode is either a hookworm or a Strongyloides.” In the same year, Allison et al. (1974) demonstrated the presence of hookworm in the prehistoric New World by identifying Ancylostoma duodenale in an Inca mummy. As noted before, there was evidence suggesting the presence of Strongyloides on the Colorado Plateau (Hall, 1972) a t the time of the Mummy Cave analysis. Had Dusseau and Porter been aware of these studies, one wonders whether they would have modified their conclusionsto a diagnosis of Strongyloides or hookworm. The identification of possible hookworm in the intestinal contents of the South Carolina skeleton is intriguing (Rathbun et al., 1980). If the nematode can be definitely identified as hookworm, then this area would appear t o have been as endemic to hookworm in prehistory as it is today. Currently, the remains are under reexamination, and clarification of the identification will hopefully be forthcoming. Unfortunately, rigor in diagnosis is sometimes hampered by marginal preservation. McClary’s find of D. latum is a case in point. Traditionally, this species is considered to be an historic introduction. Because he feels that dogs are possibly the source for the coprolites he studied, one immediately wonders whether the related genus Spirometra is involved. This is a common parasite of dogs that rarely infects humans (Schmidt and Roberts, 1981). In this case, clear documentation of the abopercular protuberance would support the claim that it is D. latum. The protuberance is common on D. latum eggs but is not present on Spirometra eggs. Differential diagnosis is not sufficiently addressed by North American researchers, a problem exemplified by McClary’sDiphyllobothrium find. In northeastern North America, humans can be infected with Diphyllobothrium ursi as well as D. latum, a fact not specifically addressed by McClary. With respect to prehistoric parasitism in this area, future differential diagnosis of prehistoric diphyllobothriid tapeworm eggs should focus on separation of Spirometra from Diphyllobothrium and then, if possible, separation of D. latum from D. ursi. The Anasazi and Sinagua were infected by hymenolepidid cestodes (Reinhard et al., 1987).The identification of these remains to genus or species is hampered by the preser-

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vation of only decorticated embryophores (inner portions of the eggs), which lack the outer egg shell or “capsule.”In this case, poor preservation prevents differential diagnosis. Differential diagnosis in South America involves desiccation of modern parasite eggs followed by rehydration (Araujo, 1988; Confalonieri et al., 1985). This procedure provides excellent information regarding the potential alteration of helminth eggs in coprolites. Future research in North America must include dessication studies to clarify diagnostic problems such as that presented by possible hookworms, diphyllobothriid tapeworms, and hymenolepidid tapeworms. In some cases, eggs of different species are morphologically similar and therefore present diagnostic problems. The finds of strongylate eggs in coprolites from Antelope House, Arizona, and Rio Zape, Durango, present a high degree of ambiguity. Initially, the eggs were identified as Trichostrongylus based on the size of the eggs in comparison to ancylostomid eggs and on the absence of ancylostomid genera in the modern Colorado Plateau (Reinhard, 1985b; Reinhard et al., 1987). Later, this identification was modified. As stated by Reinhard (1988a) and Reinhard et al. (19871, it was impossible to determine from the shape of the eggs or from the morphology of the larvae within the eggs whether they are from a trichostrongyle (wire worm) or ancylostomid (hookworm) species. Therefore the eggs were identified as “strongylate.” Recently, paleoparasitological researchers in Brasil have found that the Trichostongyloidea of man can be easily identified based on egg size (Luiz Fernando Ferreira, FIOCRUZ, personal communication). Since the size of the eggs from Antelope House coprolites is more consistent with trichostrongyles, it is probable that they belong in the order Trichostrongyloidea. Continued research will, if possible, clarify the identification. The find of Ent. vermicularis eggs in the same coprolite as trichostrongyle eggs demonstrates that the eggs are associated with human coprolites (Reinhard et al., 1987). Although well preserved, the find of taeniid tapeworm eggs in prehistoric North America also presents interpretive problems. The only taeniid species that are known to use humans as definitive hosts are Taenia solium and Tae. saginata ( = Taeniarhynchus saginatus). They use domestic pigs and cattle, respectively, as intermediate hosts. Since pigs and cattle were not

present in the prehistqric New World, the origin of taeniid eggs in New World coprolites is problematical. Many other taeniid species infect dogs. The eggs were possibly introduced into the human digestive tract with food containing eggs noninfective to humans through close association with dogs. They should not be considered prehistoric human parasites simply on the basis of their presence in human coprolites. It is likely that prehistoric peoples fortuitously consumed the eggs, which were then harmlessly passed. Taeniids are probably of more concern to the modern parasitologists who find the eggs in coprolites than they were to the ancients who produced the coprolites. The finding of parasite eggs of species that normally occur in animals is cause for skepticism. For this reason, the presence of M . clarki in human coprolites warrants attention. Although Moniliformis dubius can infect humans (Noble and Noble, 1982) and M . moniliformis (=M. dubius) can infect man under experimental conditions (Schmidt and Roberts, 1981:552), M. clarki has not been reported as a human parasite. This throws doubt on the finds from Glen Canyon, Danger Cave, and Hogup Cave as cases of human parasitism. As Fry (1980:336) states, the presence of the eggs in the human coprolites “could represent false parasitism by ingestion of adult worms with eggs in the bodies of rodents, or true parasitism by ingestion of the larval stages in the bodies of insects.” The habit of ingesting whole rodents and insects allows for either possibility (35 of the Danger Cave coprolites and 36 of the Hogup Cave coprolites contain bone from the consumption of small animals). The known definitive host range of M. clarki is broad and includes three known orders: Insectivora, Rodentia, and Chiroptera. Known definitive host genera of M . clarki include Sciurus, Glaucomys, Scalopus, Geomus, Spermophilus (=Citellus), Apodemus, Meriones, Tamias, Entamias, Mephitis, and Pitymus. In determining whether or not acanthocephalan eggs represent true infections, examination of dietary components is helpful. If false parasitism occurred, one would expect to find consistently rodent bone in the coprolites that contain the parasite eggs. However, if true parasitism occurred, one would expect eggs t o occur in coprolites that contain bone as well as in coprolites that do not. Hall (1977) found that among the five coprolites containing acanthocephalan eggs

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from Dirty Shame Rockshelter, all contained bone. He concluded that the presence of the eggs was probably due to false parasitism. However, he does not rule out the alternative possibility that true human infections are reflected by the presence of the eggs. In reviewing Fry’s analysis of several Utah caves including Hogup and Danger Caves, nine coprolites contain eggs of M . clarki. Of these, five contain bone and four do not. The absence of bone in four of the coprolites is circumstantial evidence that true infections occurred. Whether acanthocephalans parasitized prehistoric peoples is debatable. It is possible that Moniliformis was a prehistoric human parasite for four reasons. First, considering the common prehistoric habit of eating insects in North America, the potential for human exposure to infective stages of acanthocephalans was high. Second, the other species in the genus infects humans, which suggests a potential for M. clarki to infect humans. Third, the fact that this species has a wide definitive host range underscores the potential that it could infect humans. Finally, acanthocephalan eggs are commonly found in coprolites from Utah,which suggests that humans were often infected by M. clarki or another acanthocephalan species. Planned research in the Coprolite Research Laboratory of the University of Nebraska, in conjunction with the Department of Zoology, will hopefully clarify this issue. The identification of Strongyloides (hairworm) is fraught with difficulty. As stated by Reinhard (1985b,c), confusion with freeliving nematodes such as Rhabditis is possible, even when the larvae are in excellent preservation. Circumstantial evidence, along with morphological analysis, resulted in the probable identification of Strongyloides at Antelope House. The circumstantial evidence included the facts that only first-stage larvae were present, the morphology was identical to Strongyloides, there was an absence of evidence that the coprolite had been colonized by coprophagous organisms, and the coprolite desiccated rapidly. The Strongyloides identification now has support in the recent finding of obvious thirdstage Strongyloides larvae in coprolites from Antelope House (Reinhard, 1988a). Several researchers (Gooch, 1983; Reinhard, 1985b; Samuels, 1965) emphasize the danger of making a specific diagnosis from archaeoparasite remains without due consideration. One should approach archaeo-

parasite analysis with a healthy measure of skepticism. In most cases, identifications are based on helminth reproductive products (eggs and larvae) recovered from co rolites, the human origin of which is often ubious. Modern species identification usually involves examination of the adult worm. Although fragments of ancient adult worms are found in coprolites, intact adult worms are found only in mummies. Consequently, one must be doubly cautious in making diagnoses when working with coprolites. Human parasitism has great antiquity, as documented by parasite finds from prehistoric contexts. Aacanthocephalans, possibly M. clarki, have the greatest antiquity in the Great Basin of Utah. Eggs of this parasite have been found in coprolites that are over 10,000 years old (Table 2). Although pinworm (Ent. vermicularis) has been documented in coprolites that are about 10,000 years old, the eggs are not commonly found until agriculture is established in the Southwest. Until agriculture is established, the prevalence of pinworm eggs is overshadowed by tapeworm and acanthocephalan eggs. Pinworm eggs are especially common in coprolites of Anasazi peoples from about 1,300 years to 700 years ago. The prevalence of this parasite in Anasazi coprolites is far greater than any other species (Table 2) and indicates that agriculture allowed for an increase in human-specific parasitism. Other human-specific parasites that occur late in prehistory among Southwestern agricultural peoples are Trichur. trichiurus and Asc. lumbricoides. Human-specific parasites were apparently established early among southeastern US. hunter-gatherers. Asc. lumbricoides is present as early as 1,500 years ago in Kentucky. The evidence from Daws Island suggests that hookworm (Ancylostomidae) infected people as early as 2,600 years ago. This last finding is especially important if it can be verified. If hookworm is indeed the infective organism, then the arrival of hookworm in North America definitely precedes historic times.

B

DISCUSSION

The nature of prehistoric New World parasitism has been a source of speculation in the parasitological and paleopathological literature. Until fairly recently, it was generally thought that many of the more common human-specific helminths such as Trichur. trichiura, Asc. lumbricoides, and the hook-

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K.J. REINHARD TABLE 2. Dates for prehistoric parasite finds from North America Taxon

Nematodes Adult, species unknown Eggs, species unknown Enterobius uermicularis

Trichuris trichiura

Ascaris lumbricoides

Strongyloides sp Trichostrongylus sp. Possible ancyostomidae Trichinella spiralis Trematodes Cryptocotyle lingua Unidentified egg Unidentified egg Cestodes Diphyllobothriurn latum Hymenolepididae Taeniidae

Echinococcus sp. Acanthocephalans Unidentifiable eggs Moniliformis clarki

Date

Localitv

A.D. 700-1200 A.D. 500-1200 Cd. 8000 R.C. 4800-4300 B.C. 4010 B.C. 2100-600 B.C. 1250 B.C. 650 B.C. A.D. 400 A.D. 600 A.D. 600 A.D. 500-1200 A.D. 920-1020 A.D. 1080-1130 A.D. 1000-1200 A.D. 1075-1140 A.D.llOO-1250 A.D.llOO-1250 A.D.1250-1300 A.D. 1100-1250 A.D. 1,720 A.D. 1760-1776 ca. A.D. 1806 A.D. 1830-1850 570-290 B.C. A.D. 1100-1250 A.D. 1720 A.D. 1760-1776 ca. A.D. 1806 A. D. 500- 1200 A.D. 1075-1140 A.D. 1075-1140 1300-1700 B.C. A.D. 1550

Granado Cave, Texas* Clyde’s Cavern, Utah Danger Cave, Utah Dirty Shame Shelter, Oregon Hogup Cave, Utah Hinds Cave, Texas Hogup Cave, Utah Hogup Cave, Utah Turkey Pen Cave, Utah Antelope House, Arizona* Ria Zape, Durango Clyde’s Cavern, Utah Pueblo Bonito, New Mexico Pueblo Bonito, New Mexico Mesa Verde, Colorado Anetlope House, Arizona Elden Pueblo, Arizona Salmon Ruin, New Mexico Inscription House, Arizona Elden Pueblo, Arizona Colonial Williamsburg, Virginia Newport, Rhode Island Newport, Rhode Island Greenwich Village, New York Upper Salts Cave, Kentucky Elden Pueblo, Arizona Colonial Williamsburg, Virginia Newport, Rhode Island Newport, Rhode Island Clyde’s Cavern, Utah Antelope House, Arizona Antelope House, Arizona Daws Island, South Carolina Point Barrow, Alaska*

A.D. 400 A.D. 1250-1300 500 B.C.-A.D. 1150

St. Lawrence Island, Alaska’ Glen Canyon, Utah Lovelock Cave, Nevada

300 B.C.-A.D. 200 A.D. 1075-1140 A.D. 1100-1250 ca. 4500 B.C. ca. 4200 B.C. ca. 2000 B.C. 300 B.C.-A.D. 200 ca. 20 A.D. A.D.llOO-1250 A.D. 1250-1300 Pre-Contact A.D. 600

Schultz Site, Michigan Antelope House, Arizona Elden Pueblo, Arizona Hogup Cave, Utah Hogup Cave, Utah Hogup Cave, Utah Schultz Site, Michigan Danger Cave, Utah Elden Pueblo, Arizona Glen Canyon, Utah Kodiak Island, Alaska* North Dakota’

A.D. 460-1500 A.D. 900-1100 A.D. 900-1100 10,000-8500 B.C. ca. 8000 R.C. 6400-4856 B.C. 4800-4300 B.C. 4300-5900 B.C. ca. 2000 B.C. 1869 B.C. ca. 20 A.D. A.D. 600-900

Clyde’s Cavern, Utah Black Mesa, Arizona Glen Canyon, Utah Danger Cave, Utah Danger Cave, Utah Hogup Cave, Utah Dirty Shame Shelter, Oregon Dirty Shame Shelter, Oregon Hogup Cave, Utah Danger Cave, Utah Danger Cave, Utah Dirty Shame Shelter, Oregon

‘Asterisks indicate finds in mummies or skeletons. All other finds are from coprolites or latrine remains

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worm genera were historic introductions into the New World (Schmidt and Roberts, 1981; Desowitz, 1981). Prehistoric humans were felt to have been parasitized by very few parasite species. For example, Desowitz (1981) reported that the pinworm (Ent. vermicularis) was the only human-specific helminth parasite among prehistoric New World populations. In essence, from the perspective of helminth parasitism, the prehistoric New World was viewed as relatively worm free, and the morbidity caused by helminths in the Old World was not suffered in the New World. The corpus of archaeoparasite data from North America, combined with data from South America, is large enough to draw some conclusions regarding prehistoric parasitism and debunk the notion that the New World was relatively free of parasitism. Most importantly, the evidence a t hand indicates the presence of certain parasites that were once considered to be introductions from the Old World in historic times. These include at least one hookworm species,Anc. duodenale, and also the whipworm Trichur. trichiura. Other parasites that are commonly found in modern populations, such as hymenolepidid tapeworms, Asc. lumbricoides, and Strongyloides, were present in prehistoric New World peoples. The evidence for certain parasites, such as hookworm, Trichur. trichiura, and Asc. lumbricoides, is much more abundant in South America than in North America (Horne, 1985). This does not reflect a greater prevalence of infection in South America than in North America. The paucity of evidence of these genera is due to a North American research focus on coprolites from desert areas. In deserts, it is unlikely that these genera could complete their life cycles. As coprolites from the southeastern United States are examined in the future, a greater range of parasites will probably be documented for North America. Importantly, it appears that zoonotic parasites had adapted early on to prehistoric human populations. Zoonoses faced by New World populations included acanthocephalan infection, hydatid cyst disease caused by infection with Echinococcus, and trichinosis from Trichin. spiralis. Future research may verify McClary’s Diphyllobothrium find. Considering the close relationship between man and dog among New World peoples, it is probable that the zoonosis resulting from infection with the dog roundworm Toxocaru canis was also a problem (Reinhard, 1985~).

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A related genus, Toxascaris, has recently been recovered from dog feces excavated from an Anasazi site (Gardner and Clary, 1987). Human parasitism in North America has great antiquity. The oldest human coprolites excavated from North America are from Hogup and Danger caves in Utah. These date to about 8000 B.C. and contain eggs of Ent. vermicularis and acanthocephalans. These parasites persist in later populations occupying the caves to historic times (Fry, 1977). Although few coprolites have been examined for this 10,000year period, the numbers of coprolites studied in the southwestern United States are beginning t o reach numbers that can be statistically evaluated. The differences in parasitism between ancient hunter-gatherer populations and agricultural populations is a focus of recent research (Reinhard, 198813; Reinhard and Miller, 1988). From seven hunter-gatherer sites, 357 coprolites have been examined, 14 of which contained helminth remains. From nine agricultural sites on the Colorado Plateau, 513 coprolites have been examined, 89 of which contained helminth remains. A comparative evaluation of these data (Reinhard, 1989) shows that archaic hunter-gatherer groups were relatively free of helminth parasitism in comparison to later agricultural peoples (x2 = 35.9, P < 0.001). THE ROLE OF ARCHAJIOPARASITOLOGY

Until the 1980s, archaeoparasitology focused on observation, description, and documentation of prehistoric helminthiasis. Its affiliation was not defined as a distinct subfield within physical anthropology. Now the archaeoparasitology data base is large enough to make comparative statistical evaluation of prehistoric sites, regions, and lifestyles (Reinhard, 1988a,b, 1989; Reinhard and Miller, 1988). The most significant development of the 1980s is simply that large numbers of coprolites are available to provide a base for quantitative analysis. Archaeoparasitology is beginning to make a significant contribution to the study of human biology, specifically, in the realms of cultural ecology of parasitism (Fry, 1977, 1980; Reinhard, 1985a, 1988a,b), evolution of human parasitism (Kliks, 19831, and paleopathology (Horne, 1985; Reinhard, 1988a,b). The development of archaeoparasitology as a technique for evaluating disease was predicted by Dunn (1972) and Cockburn

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(1971). Dunn recognized that the extant hunter-gatherer life-style was undergoing rapid change. He suggested that one avenue of researching disease of aboriginal huntergatherers is through the study of coprolites. Cockburn recognized the potential of coprolite study in elucidating change in disease through various stages of cultural evolution. As significant numbers of coprolites are analyzed, the potential of coprolite analysis as foreseen by Dunn and Cockburn will be actualized. Archaeoparasite data have long been incorporated in paleopathology. Ruffer’s research (1910) with schistosome eggs in an Egyptian mummy was a major advance in paleopathological diagnosis as noted by Ubelaker (1982).Later developments within the field, however, focused on osseous pathology. Techniques of differential diagnosis and interpretation in archaeoparasitology have been devised relatively recently. Consequently, archaeoparasitology is only now taking a prominent role in paleopathology. Archaeoparasitology can be incorporated into the broad framework of bioarchaeology to assess human adaptive success. Researchers in the Southwest have pointed out that natural ecological conditions and aspects of cultural ecology affect parasitism (Fry, 1980; Reinhard, 1988a,b;Reinhard et al., 1987).As a result, helminthiasis varied in intensity between regional areas (Fry, 1980) and between sites (Reinhard, 1988a,b). The presence or relative absence of parasitism reflects the success of human adaption to various ecological regimes in prehistory. Skeletal paleopathologists are now incorporating parasite data in a bioarchaeological perspective (Akins, 1986; Fink, 1985; Kent, 1986; Walker, 1985). There are many challenges to be faced in North American archaeoparasitology. The most critical of these is identification of parasite species and evaluating their impact on prehistoric community health (Reinhard, 1988a).This approach is especially challenging. The impact of parasitism varies with life conditions of its host, including nutrition, population size, and presence of other diseases. These aspects of prehistoric existence must be evaluated before inferring the health implications of parasitism. Another challenge is the recovery and analysis of truly significant numbers of coprolites that can form the basis of comparative epidemiological study. When these problems are overcome, the study of archaeoparasitology will be able to

add new and significant insights into the evolution and ecology of human parasitism. It will also provide basic bioarchaeological data relevant to human adaptive success to various environments and perhaps provide insights into parasite-related pathology. ACKNOWLEDGMENTS

Debra K. Meier, Cartagraphic Services Unit, Texas A & M University, prepared Figures 1 and 2. Dr. Luiz Fernando Ferreira (Fundaqao Oswaldo Cruz) kindly read the manuscript, and Arthur C. Aufderheide (University of Minnesota Medical School) Douglas H. Ubelaker (Smithsonian Institution), and an anonymous A.J.P.A. reviewer also critiqued the manuscript. Their welcome insights contributed to the revised paper printed here. Elizabeth A. Miller (Department of Anthropology, Arizona State University) and Brian S. Shaffer (Department of Anthropology, Texas A & M University) reviewed and corrected the mechanics of the manuscript. S.K. Edwards generously allowed me to include unpublished data from her thesis on Hinds Cave. LITERATURE CITED Akins NJ (1986) A Biocultural Approach to Human Burials From Chaco Canyon, New Mexico. Reports of the Chaco Center Number 9. Santa Fe: National Park Service. Allison MJ, Pezzia A, Hasigawa I, and Gerszten E (1974) A case of hookworm infection in a pre-Columbian American. Am. J. Phys. Anthropol. 41:103-106. Ambler J R (1984) The Desha Complex: Pre-Altithermal Adaptation at Dust Devil Cave, Utah. A symposium held at the 19th Great Basin Anthropological Conferences, Boise, Idaho (unpublished). Ambro RD (1967) Dietary-technological-ecological aspects of Lovelock Cave coprolites. Univ. Calif. Archaeol. Sum. Rep. 70:34-47. Andrews M (1977) The Life That Lives on Man. New York Taplinger Publishing Company. Araujo A (1988) Dessecacao experimental de fezes contendo ovos de ancilostomideos. In LF Ferreira, A Araujo, and U Confalonieri (eds.): Paleoparasitologia no B r a d Rio de Janeiro: PECENSP, pp. 111-112. Araujo AJG, Ferreira LF, andConfalonieriUEC (1981)A contribution to the study of helminth findings in archaeological material in Brazil. Rev. Brasil. Biol. 41t873-881. Araujo AJG, Ferreira LF, Confalonieri UEC, and Meirelles MN (1988) Microscopia de varredura de larvas de ancylostomideos encontradas ern coprolitos humanos datados DE 3490 t 1 2 0 a 430 -t 70 anos. In LF Ferreira, A Araujo, and U Confalonieri (eds.): Paleoparasitologia no Brasil. Rio de Janeiro: PECIENSP, pp. 109-110. Araujo AJG, Ferreira LF, Confalonieri UEC, and L Nunez (1983) Eggs of Diphyllobothrium pacificum in pre-Columbian human coprolites. Paleopathol. Newslett. 41:ll-13. Brooks RH, Kaplan L, Cutler HC, and Whitaker TW (1962) Plant material from a cave on the Rio Zape, Durango, Mexico. Am. Antiquity 27:356-369.

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