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Anthropology Faculty Publications Anthropology, Department of

1-1-2008 Archaeoparasitology Karl J. Reinhard University of Nebraska at Lincoln, [email protected]

Adauto Araújo Fundação Oswaldo Cruz

Reinhard, Karl J. and Araújo, Adauto, "Archaeoparasitology" (2008). Anthropology Faculty Publications. Paper 22. http://digitalcommons.unl.edu/anthropologyfacpub/22

This Article is brought to you for free and open access by the Anthropology, Department of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Anthropology Faculty Publications by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Published in Encyclopedia of , p. 494-501, ed. by Deborah M. Pearsall. ©2008, Elsevier, Academic Press, New York. Used by permission.

Archaeoparasitology Karl J. Reinhard, University of Nebraska, Lincoln, NE, USA ([email protected]) Adauto Araújo, Escola Nacional de Saúde Pública, Fundação Oswaldo Cruz, Rio de Janeiro, Brazil

Glossary archeoparasitology The study of parasite evidence from ar- cheological sites. acanthocephalan Any of various worms in the phylum cestode Any of the parasitic flatworms of the class Ces- Acanthocephala, also called thorny-headed worms, living toidea, including the tapeworms, having a long, segmented, in intestines of vertebrates having a retractile proboscis cov- flat body equipped with a specialized organ of attachment ered with many hooked spines. at one end. anthelminthic A compound that affects and causes the ex- ectoparasite Parasites such as lice and flies that live on the pulsion of parasitic intestinal worms. body’s outer surface.

494 Archaeoparasitology 495 endoparasite Parasites such as blood flukes and pinworms and also by certain gross pathology they left in their that infect the internal parts of the body. mummified hosts. Helminths are parasitic worms in- helminth Worm that is parasitic on vertebrates, especially cluding nematode roundworms, cestode tapeworms, roundworms and tapeworms, thorny-headed worms, and trematode flukes, and acanthocephalan thorny- flukes. An organism that provides food and shelter to a para- headed worms. Helminth eggs from some species are site. microparasites laid in thousands within their hosts. Helminth eggs A microscopic organism of medical importance including from humans and domestic contaminated an- bacteria, viruses, fungi, and . cient villages. The eggs are very durable and are eas- infestation Parasites that are present on the outside of the ily retrieved from archaeological sediments, copro- hosts, such as ectoparasites, or the contamination of a habitat lites, and . and lice can be found with arthropods such as mosquitoes, bed bugs, and . on mummies and also in archaeological sediments. infection Parasites that are present inside the host organism, Lice are especially important in studies be- Including helminths and microparasites. nematode Any of phylum Nematoda of elongated cylindri- cause the eggs are cemented on hair shafts. There- cal worms some of which are parasitic in animals and plants, fore, examination of scalps from mummies provides a and others of which are free-living in soil or water. method of quantifying infestations between individ- parasite An organism that lives at the expense of its host by uals and sites. taking energy from the host and sometimes causing pathol- The discipline that focuses on the relationships be- ogy in the host. tween behavior, environment, and parasite infection prevalence Number of hosts in a population infected with a is archaeoparasitology. This field developed from the parasite at any one time. need for a fine-tuned analysis of prehistoric ecologi- protozoa (parasitic) Single-cell organisms, some of which are parasitic and can only reproduce within a host organism. cal and behavioral conditions to assess the factors that Malaria is caused by a protozoa, Plasmodium. Other proto- affected disease. Archaeoparasitology depends on ar- zoan parasites are Giardia and Toxoplasma. chaeological information regarding community size, trematode Referring to flukes, phylum Trematoda, which trade patterns, water sources, subsistence practices, are parasitic flatworms having external suckers for attach- social stratification, environment, medicine use, and ing to a host. many other lines of modern archaeological investiga- vector An , usually a biting insect, that is responsible tion. It also depends on biological understanding of for the transmission of a parasitic organism. complex parasite life cycles and other dimensions of parasite ecology. When broadly applied, archaeopar- Introduction asitology defines the rise in parasitic disease -associ ated with the development of complex societies and Parasites are the major cause of ill health and early changes in subsistence strategies. In a more restricted death in the world today. Malaria, sleeping sickness, application, archeoparasitology sheds light on the amoebic dysentery, and hookworm infection are ex- health impact of urbanization and empire expansion. amples of commonplace parasitic diseases that are en- When tightly applied to a single burial or mummy, ar- demic in most parts of the world (see Health, Healing, cheoparasitology shows how habits promote disease and Disease). They were significant threats in prehis- on an individual basis. tory, especially in cultures whose social complexity outstripped the development of effective sanitation, History and Major Themes hygiene, and germ theory awareness. Parasites are organisms that live in or on other or- Aidan Cockburn explored the origins of disease and ganisms called hosts. Parasites derive sustenance and generated interest in archeoparasitology. Cockburn shelter from their hosts and carry out reproduction in theorized that there was a relation between human host tissues and structures. There is a wide amount cultural development and the of infectious of taxonomic diversity among parasites. They range diseases. In the first archeoparasitological study, Re- from single-celled protozoa, such as amoeba, to mul- inhard compared Colorado Plateau Archaic parasit- ticelled arthropods such as fleas. Strictly defined, par- ism to agricultural Puebloan sites. Reinhard verified asites do not include bacteria and viruses. However, Cockburn’s hypothesis that occasional hunter-gath- some epidemiologists refer to bacteria and viruses as erer infections became major agricultural health haz- microparasites. There are two general types of par- ards (Figure 1). The reasons for the emergence of par- asites: ectoparasites such as lice and endoparasites asitic disease were many. was limited in such as intestinal worms. hunter-gatherer societies, called bands. Hunter-gath- All types of parasites can be found in archeological erer parasitism was limited by small band size, dif- sites. Protozoa can be identified by traces of antigens fuse regional populations, high band mobility, and 496 Reinhard and Araújo in Encyclopedia of Archaeology (2008)

presence of natural anthelminthics in hunter-gatherer to control their parasite burden very effectively while diets.The one factor that could have promoted hunter- others were simply overwhelmed by their pathogens. gatherer parasitism was the consumption of uncooked This topic was explored by a comparison of pinworm vertebrate meat and insects. Parasitism was promoted (Enterobius vermicularis) prevalence in by in descendent agricultural Puebloan communities a group of specialists in pinworm disease. Pinworm by contaminated water sources, concentrated popu- was chosen as an indicator of general infectious dis- lations, more sedentary life, apartment-style living, ease because it is transferred from person to person absence of effective sanitation, activities centered on and by contamination of living quarters and food water (agriculture), and activities that expanded wet- (Figure 2). Some ancestral Pueblo communities were lands including irrigation of all types. extremely parasitized. In a clinical setting, only 5% of Reinhard recognized that the parasite variation be- feces from pinworm-infected people are positive for tween agricultural Puebloan villages nearly equaled pinworm eggs. The percentages of coprolites positive the variation between agriculturalists and hunter- for pinworm from several sites exceed this and range gatherers. This means that some settlements managed up to 29% (Figure 3). The lowest prevalence was found Archaeoparasitology 497

in small cave sites not containing walled villages. The heavy infections. In such populations, pinworm in- highest prevalence came from large, walled villages fection prevalence reflects serious health risks, when built in rock shelters (Figure 4). Hugot et al. concluded one considers that other pathogens are spread by the that poor air circulation in large populations living same means. Reinhard showed that the prevalence of in complex apartment-style communities resulted in parasitism co-varied with porotic hyperostosis prev- truly impressive levels of pinworm parasitism. In fact, alence at ancestral Pueblo sites where both some sites have the highest levels of pinworm infec- and skeletons were studied (Figure 5). Porotic hyper- tion recorded for ancient or modern peoples. ostosis is a general skeletal pathology indicator long The data indicate that pinworm parasitism was used to assess maternal-infant health. unavoidable and that in some villages people had In Brazil, the cultural transfer of parasites has been 498 Reinhard and Araújo in Encyclopedia of Archaeology (2008)

a focus for many years. The discovery of hookworm and whipworm in prehistoric South American mum- mies and coprolites was sensational. These are human- In historic archeology, archeoparasitology focuses specific nematodes that cannot parasitize humans in on sociological and urbanization concerns. Historic the Arctic and subarctic because they require warm, archeologists can define the ethnicity, economic level, moist soils for maturation once the eggs are defecated. and social status of people associated with archeologi- Therefore, the discovery of these parasites in prehis- cal features. Therefore, archeoparasitologists have the toric South America mummies and coprolites was opportunity to examine the effect of social differenti- sensational. These are human-specific nematodes that ation on parasitism. The role of urbanization on the cannot parasitize humans in the Arctic and subarctic emergence of parasitic disease is a common theme in because they require warm, moist soils for maturation historic context along with the development of sani- once the eggs are defecated. Therefore, the discovery tation in controlling parasitism. Ascarid roundworms of these parasites in prehistoric South America indi- (Ascaris lumbricoides ) and whipworms (Trichuris trichi- cated that there was a nonarctic migration of humans ura) are the main indicators for assessing the parasitic from the Old World to the Americas. state of historic sites. These two species are most asso- The most long-standing debate in archeoparasitol- ciated with fecal contamination, crowding, and poor ogy revolved around the discovery of hookworms in sanitation. These parasites are used for comparative prehistoric sites. One species of hookworm, Ancyslos- evaluation of the threat of parasitism between neigh- toma duodenale, was diagnosed from examination of borhoods, villages, and cities. Other parasites, espe- adult worms in prehistoric Peruvian mummies, and cially tapeworms and flukes associated with different later larvas were discovered in coprolites and mum- types of meat, are useful indicators of ethnicity. mies from Brazil, and the United States. Hookworms In the twenty-first century, the discipline of ar- are host specific, which means that one species of cheoparasitology became global. Researchers pub- worm only infects one species of host. Ancyslostoma lished parasitological finds from Japan, Korea, Ger- duodenale only infects humans. Hookworms require many, Peru, Chile, Brazil, and many other countries. tropical or subtropical environments for their eggs to The intellectual foci of these studies are diverse. In Ja- hatch and larvas to mature to infective stage. Finally, pan, parasites were analyzed in context of the devel- hookworms have their evolutionary origins in the Old opment of sanitation and food practices. They were World. Therefore, to reach the New World, they had also used to identify areas used by foreign ambassa- to migrate with human populations from a tropical or dors who hosted parasite species exotic to Japan but subtropical environment (Figure 6). The conventional endemic to China. In Korea, archeoparasitology was wisdom of the twentieth century was that hookworms used to trace the origins of indigenous species, es- arrived in the New World in historic times with Euro- pecially trematodes. The impact of the expansion of pean colonists and African slaves. This conventional the Inca Empire was defined in Chile. There, the Inca wisdom has been so strong that over a dozen papers compelled indigenous people to move from small, have appeared in anthropological and parasitologi- scattered communities to large towns, resulting in in- cal journals debating the validity and meaning of the creased infection with certain parasite species. Also, hookworm finds. parasitism of the oldest hunter-gatherers, the Chin- Archaeoparasitology 499 chorro, was characterized. Chinchorro consumption of undercooked fish resulted in heavy cestode infec- tion. Archeoparasitology in Peru examines the dis- eases of humans and domestic animals, and especially the transfer of deadly protozoa from animals to hu- mans via insect vectors. One theme that crosscuts the global diversity of modern archeoparasitology is defining the distribu- tion of parasites. In a Brazilian mummy, Sianto et al. discovered eggs of hookworm and of a trematode ge- nus, Echinostoma. Echinostoma has never been found in people from the Americas and shows that an indig- enous species has the ability to infect humans. This adds to the medical knowledge of the diversity of par- asites infective to humans. On the Texas–Coahuila border, Reinhard et al. discovered the gross pathology of megacolon which is often associated with Chagas’ disease. Infection with Trypanosoma cruzi causes Cha- gas’ disease. Previously, Chagas’ disease was thought to have originated in the high Andes and then spread to lowland South America in historic times. The dis- covery of Chagas’ disease in prehistoric border of Mexico and Texas shows that the disease spread fur- ther and earlier than generally believed.

Data Sources and Methods Archeoparasitologists find their data in a variety of ar- cheological contexts. Historical medical texts provide information about ancient parasitological knowledge and treatment. Artifacts provide rare glimpses of the pathology caused by certain parasites. For example, potters of the Peruvian Moche culture portrayed the facial disfigurement resulting from Leishmania infec- tion (Figure 7). This protozoan parasite can cause de- struction of the soft and hard tissue of the face. Skeletal remains can reveal hard tissue pathology caused by parasites. Calcification of soft tissue of the urinary tract often results from Schistosoma hemato- bium (blood fluke) infection. Cysts of the tapeworm Echinococcus granulosis calcify and are preserved in burials (Figure 8). Destructive bone lesions resulting from Leishmania infection are evident in skeletal re- mains from Peru (Figure 9). Sediments such as soil Mummies preserve the hard tissue and soft tissue within burial pelvic girdles (see Burials: Dietary Sam- pathology caused by parasites as well as the parasites pling Methods) are an important source of informa- themselves. Ectoparasites such as fleas, head lice, body tion about parasites. For example, German research- lice, and crab lice are easily recovered from mummies ers were able to recover liver fluke (Fasciola hepatica) and the clothing buried with mummies. Parasites of from the sediment of human and cattle pelvic gir- the lungs, intestinal tract, liver, and urinary tract are dles. This showed that this debilitating parasite was a evident. Molecular biological diagnosis can recover threat to humans and their domestic livestock. the DNA of ancient parasites from mummified tissue,

500 Reinhard and Araújo in Encyclopedia of Archaeology (2008)

rarely found with the microscope. Molecular biolog- ical characterization of ancient parasite DNA is very useful in making definitive diagnosis of ancient para- sites and identifying genetic strains of single species. The power of modern archeoparasitology is based based on its ability to quantify infections for compara- tive evaluation of disease. There are a variety of meth- ods evaluation of disease. There are a variety of meth- ods for quantification of eggs per milliliter or gram of archeological sediments and coprolites. These include dilution methods derived from clinical techniques and concentration methods derived from palynolog- ical techniques. Quantifying eggs allows comparative evaluation of parasitism between sites and features within sites.

The Future of Archeoparasitology Several trends in archeoparasitology are evident by comparing the nature of studies in the last century and the current century. Many studies from 1960 to even when the parasites themselves have decom- 1990 are focused on the recovery and diagnosis of posed. Therefore, each mummified corpse is- ex ancient parasites. With the exception of Brazilian re- tremely important in the analysis of parasitic diseases. search into migration and Southwestern research into Also, mummified animals are a wonderful source epidemiology, few researchers answered behavioral about parasites that may have threatened the vitality questions with archeoparasitological data. This pre- of domestic animals. liminary stage is over. Now archeoparasitologists In the twentieth century, coprolites provided most place their data in behavioral context to reveal aspects of the information about parasitic disease. The eggs of migration, food preparation, effect of social status and larvas of parasites that disperse their offspring on disease, cross-infection between humans and an- are easily found in coprolites. However, Bain de- imals, and many other topics of anthropological in- scribes the increasing importance of analysis of do- terest. There is also a new interest in the influence of mestic archeological sediments in archeoparasitology. parasitic disease on the vitality of cultures and site Sediments from latrines, sewers, drains, streets, yards, abandonment. Finally, there is now a solid nexus be- and living floors contain parasite eggs. Parasites were tween biological parasitology and archeoparasitol- so abundant in medieval and colonial villages that ogy in exploring questions of parasite hundreds of parasite eggs per milliliter of house sed- and endemicity that have relevance to modern health. iments are commonly found. In latrines, drains, and These areas will continue to expand as archeoparasi- sewers, the numbers of eggs range into hundreds of tology becomes a standard archeological discipline. thousands per milliliter of sediment. The analysis of sediments from domestic context will be increasingly See also: Americas, North; American Southwest, Four important in the future. Corners Region; Burials; Dietary Sampling Methods; The microscope is the main tool of the archeopar- Health, Healing, and Disease; New World, Peopling of. asitologist. Most diagnoses of helminths and arthro- pods have been made with compound or binocular Further Reading microscopes. However, molecular biology and en- Aidan CT (1971) Infectious diseases in ancient popu- zyme diagnostic methods have expanded the range of lations. Current Anthropology 12: 45–62. parasites identified from archeological sites. Enzyme- Bain A (2001) Archaeoentomological and archaeopar- linked immunoassay is a new, proven method for the asitological reconstructions at îlot Hunt (CeEt-110): identification of parasite antigens that can be applied New perspectives in historical archeology (1850– to coprolites,mummies, and all types of archeological 1900). In: BAR International, p. 153. Series 973, Ox- sediments. So far, protozoa have been effectively diag- ford: Archaeopress. nosed. This is very important because protozoa cysts Fisher CL, Reinhard KJ, Kirk M, DiVirgilio J, and are ephemeral in archeological remains and are only Miller TS (in press) Archeoparasitology of Historic Archaeoparasitology 501

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