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Archaeoparasitology'''Archaeoparasitology''', a multi-disciplinary field within paleopathology, is the study of parasites|parasites in Archaeology|archaeological contexts.[Reinhard, K.J. and A. Araújo (2008) Archaeoparasitology. In: D.M. Pearsall (ed) ''Encyclopedia of Archaeology''. Elsevier: Amsterdam, pp. 494-501.] It includes studies of the protozoan and animal|metazoan parasites of humans in the past, as well as parasites which may have affected past human societies, such as those infesting domesticated animals.
Reinhard suggested that the term "archaeoparasitology" be applied to "... all parasitological remains excavated from archaeological contexts ... derived from human activity" and that "the term 'paleoparasitology' be applied to studies of nonhuman, paleontological material." (p. 233)[Reinhard, K.J. (1992) Parasitology as an interpretive tool in archaeology. ''American Antiquity'' 57(2):231-245.] Paleoparasitology includes all studies of ancient parasites outside of archaeological contexts, such as those found in amber,[Poinar, G.O., Jr. and H. Poinar (2004) ''Paleoleishmania proterus'' n.gen., n.sp. (Trypanosomatidae: Kinetoplastida) from Cretaceous Burmese amber. ''Protista'' 155(3):305-310][Wier, A., M. Dolan, D. Grimaldi, R. Guerrero, J. Wagensberg, and L. Margulis (2002) Spirochete and protist symbionts of a termite (''Mastotermes electrodominicus'') in Miocene amber. ''Proceedings of the National Academy of Sciences of the USA'' 99(3):1410-1413.] and even dinosaur parasites.[Poinar, G., Jr. and A.J. Boucot (2006) Evidence of intestinal parasites of dinosaurs. ''Parasitology'' 133(2):245-249.]
The first archaeoparasitology report described calcified eggs of Schistosoma haematobium|''Bilharzia haematobia'' (now ''Schistosoma haematobium'') from the kidneys of an ancient Egyptian mummy.[Ruffer, M.A. (1910) Note on the presence of ''Bilharzia haematobia'' in Egyptian mummies of the Twentieth Dynasty (1250-1000 BC). ''British Medical Journal'' 1:16.] Since then, many fundamental archaeological questions have been answered by integrating our knowledge of the host (biology)|hosts, parasitic life cycles|life cycles and basic biology of parasites, with the archaeology|archaeological, anthropology|anthropological and history|historical contexts in which they are found.
Parasitology basics
Parasitism|Parasites are organisms which live in close association with another organism, called the host, in which the parasite benefits from the association, to the detriment of the host (biology)|host. Many other kinds of associations may exist between two closely allied organisms, such as commensalism or mutualism.
Parasitism#Types_of_parasitism|Endoparasites (such as protozoans and parasitic worm|helminths), tend to be found inside the host, while Parasitism#Types_of_parasitism|ectoparasites (such as ticks, lice and fleas) live on the oustide of the host body. parasitic life cycles|Parasite life cycles often require that different developmental stages pass sequentially through multiple host species in order to successfully mature and reproduce. Some parasites are very host (biology)|host-specific, meaning that only one or a few species of hosts are capable of perpetuating their life cycle. Others are not host-specific, since many different hosts appear to harbor and pass on the infective stages of the parasite.
Most archaeoparasitology reports involve species which are considered to be true parasites of humans today. However, incidental parasitism (referred to by some authors as "pseudoparasitism", "false parasitism" or "accidental parasitism") occurs when a parasite which does not normally utilize a host for the perpetuation of its lifecycle is found in that host incidentally. One example is finding the eggs of ''Cryptocotyle lingua'' (a fish parasite) in the stomach contents of an Eskimo mummy.[Zimmerman, M.R. (1980) Aleutian and Alaskan mummies. In: A. Cockburn and E. Cockburn (eds) ''Mummies, Disease and Ancient Cultures''. Cambridge University Press: London, pp. 118-134.] It is estimated that 70% of the "parasite" species reported from present-day humans are actually only incidental parasites.[Ashford, R.W. (1991) The human parasite fauna: Towards an analysis and interpretation. ''Annals of Tropical Medicine and Parasitology'' 85(1):189-198.] Some incidental parasites do cause harm to the infested pseudohosts.[Sing, A., K. Tybus, and I. Fackler (2008) Acute urticaria associated with ''Dicrocoelium dendriticum'' infestation. ''Indian Journal of Medical Microbiology 26(1):97-98.]
Sources of material
In archaeological contexts, endoparasites (or their eggs or cysts) are usually found in (i) fossilized human or animal dung (coprolites), (ii) the tissues and digestive contents of mummies|mummified corpses, or (iii) soil samples from latrines, cesspits, or middens (dumps for domestic waste). A cyst of ''Echinococcus granulosus'' was even retrieved from cemetery soil in Poland.[Gladykowska-Rzeczycka, J.J., A. Wrzesinska and J. Wrzesinski (2003) Rzadkie znalezisko torbieli pasozyta z wczesnosredniowiecznego cmentarzyska w dziekanowicach / A rare finding of a parasitic cyst from an early mediaeval cemetery in Dziekanowice. Archeologia Polski 48(1-2):65-76] Ectoparasites may be found on the skin or scalp, as well as wigs, clothing, or personal grooming accessories found in archaeological sites.[Mumcuoglu, Y.K. and J. Zias (1988) Head lice, ''Pediculus humanus capitis'' (Anoplura, Pediculidae) from hair combs excavated in Israel and dated from the first century BC to the eighth century AD. ''Journal of Medical Entomology'' 25(3):545-547][Mumcuoglu, K.Y., J. Zias, M. Tarshis, M. Lavi and G.D. Stiebel (2003) Body louse remains found in textiles excavated at Masada, Israel. Journal of Medical Entomology 40(4):585-587] Ectoparasite eggs may be found attached to individual hairs.[Capasso, L. and G. Di Tota (1998) Lice buried under the ashes of Herculaneum. ''Lancet'' 351(9107):992][Rivera, M.A., K.Y. Mumcuoglu, R.T. Matheny, and D.G. Matheny, D.G. (2008) Huevecillos de ''Anthropophthirus capitis'' en momias de la tradición Chinchorro, Camarones 15-D, norte de Chile / Head lice eggs, ''Anthropophthirus capitis'', from mummies of the Chinchorro tradition, Mamarones 15-D, northern Chile. ''Chungará - Revista de Antropología Chilena'' 40(1):30-39.] The International Ancient Egyptian Mummy Tissue Bank in Manchester, England, provides tissue samples for a variety of uses, including parasitological studies.[Lambert-Zazulak, P.I., P. Rutherford, and A.R. David (2003) The International Ancient Egyptian Mummy Tissue Bank at the Manchester Museum as a resource for the pelaeoepidemiological study of schistosomiasis. ''World Archaeology'' 35(2):223-240.]
Since 1910, parasite remains have been found in archaeological samples from Africa, the Americas, Asia, Europe, the Middle East, and New Zealand. The age of archaeological sites yielding human parasite remains ranges from approx. 25,000-30,000 years ago[Bouchet, F., D. Baffier, M. Girard, P. Morel, J.-C. Paicheler, and F. David (1996) Paléoparasitologie en contexte pléistocène premières observations à la Grande Grotte d'Arcy-sur-Cure (Yonne), France. ''Comptes Rendus des Seances de l'Academie des Sciences. Série III. Sciences de la Vie'' 319(2):147-151.] to late 19th-early 20th century.[Reinhard, K.J., A. Araújo, L. Sianto, J.G. Costello, and K. Swope (2008) Chinese liver flukes in latrine sediments from Wong Nim's Property, San Bernardino, California: Archaeoparasitology of the Caltrans District Headquarters. ''Journal of Parasitology'' 94(1):300-303.] Parasite remains have also been found in domestic animal remains at archaeological sites.[Dittmar, K. and W.R. Teegen (2003) The presence of ''Fasciola hepatica'' (liver-fluke) in humans and cattle from a 4,500 year old archaeological site in the Saale-Unstrut Valley, Germany. ''Memórias do Instituto Oswaldo Cruz 98(Suppl 1):141-143.][Schelvis, J. and C. Koot (1995) Sheep or goat? ''Dalaminia'' deals with the dilemma. ''Proceedings of the Section of Experimental and Applied Entomology of the Netherlands Entomological Society'' 6:161-162]
Human skeletal remains may exhibit indirect evidence of parasitism. For example, hookworm (''Ancyslostoma duodenale'') parasitism may lead to anemia, and anemia is one factor associated with the skeletal changes of cribra orbitalia and porotic hyperostosis. Thus, hookworm parasitism ''may'' be a causal factor in observed cribra orbitalia and porotic hyperostosis,[Carlson, D., G. Armelagos, and D. Van Gerven (1974) Factors influencing the etiology of cribra orbitalia in prehistoric Nubia. ''Journal of Human Evolution'' 3(3):405-410.] though dietary factors may also lead to anemia.[Holland, T.D. and M.J. O'Brien (1997) Parasites, porotic hyperostosis, and the implications of changing perspectives. ''American Antiquity'' 62(2):183-193.]
Information on the presence of host (biology)|intermediate hosts, required for life cycle completion by many parasites, is also useful in determing the likelihood that a parasite may have infected a particular ancient society. One example is the identification of molluscan intermediate hosts of schistosomiasis in an Islamic archaeological context.[Insoll, T. and E. Hutchins (2005) The archaeology of disease: Molluscs as potential disease indicators in Bahrain. ''World Archaeology'' 37(4):579-588]
Artifacts depicting the appearance of individuals may also indicate cases of parasitism. Examples include the characteristic facial deformities of leishmaniasis found on pre-Columbian moche|Mochica pottery,[Serarcangeli, C. and A. Pennica (1996) Testimonianze di una malattia autoctona nella ceramica del Perú precolombiano / Testimonies of an autochthonous illness on the anthropomorphic pottery in ancient Peru. ''Medicina nei Secoli'' 8(1):125-141.] and morphological features of certain ancient Egyptian figurative art.[Hoeppli, R. (1973) Morphological changes in human schistosomiasis and certain analogies in ancient Egyptian sculpture. ''Acta Tropica'' 30(1):1-11] Literary sources also provide valuable information regarding not only the parasites present in historic societies, but also the knowledge and attitudes that the people had towards their parasitic infestations.[Moule, Léon (1911) La parasitologie dans la litterature antique. II. Les parasites du tube digestif. Archives d'Parasitologie 14:353-383.][Sandison, A.T. (1967) Parasitic diseases. In: D. Brothwell and A.T. Sandison (eds) ''Diseases in Antiquity''. Charles C. Thomas: Springfield, IL, pp. 178-183.][Beavis, I.C. (1988) Insects and Other Invertebrates in Classical Antiquity. University of Exeter: Devon, England. xv, 269 pp.] However, specific parasitological diagnoses reported in ancient and medieval texts must always be read with some degree of skepticism.[Bondeson, J. (1998) The bosom serpent. ''Journal of the Royal Society of Medicine'' 91(8):442-447.]
Techniques and methods
Parasite remains in archaeological samples are identified by a variety of techniques. Very durable remains, such as eggs and cysts, may remain intact for many thousands of years. In some cases, relatively intact soft-bodied adult helminths[Allison, M.J., A. Pezzia, I. Hasegawa, and E. Gerszten (1974) A case of hookworm infestation in a precolumbian American. ''American Journal of Physical Anthropology'' 41(1):103-106.][Ferreira, L.F., A. Araújo, and A.N. Duarte (1993) Nematode larvae in fossilized animal coprolites from Lower and Middle Pleistocene sites, central Italy. ''Journal of Parasitology'' 79(3):440-442.] and ectoparasitic arthropods[Araújo, A., L.F. Ferreira, N. Guidon, N.M.S. Freire, K.J. Reinhard, and K. Dittmar (2000) Ten thousand years of head lice infection. ''Parasitology Today'' 16(7):269.][Kenward, H. (2001) Pubic lice in Roman and medieval Britain. ''Trends in Parasitology'' 17(4):167-168] have been found. All of these forms can be identified to the family, genus or species level by compound or electron microscopy.
In cases where the intact bodies of parasites are not found, protein or DNA from the parasite may still be present. Antigenic and immunological assays (including enzyme-linked immunoassay - ELISA,[Deelder, A.M., R.L. Miller, N. de Jonge, and F.W. Krijger (1990) Detection of schistosome antigen in mummies. ''Lancet'' 335(8691):724-725.][Gonçalves, M.L.C., A. Araújo, R. Duarte, J.P. da Silva, K. Reinhard, F. Bouchet, and L.F. Ferreira (2002) Detection of Giardia duodenalis antigen in coprolites using a commercially available enzyme-linked immunosorbent assay. ''Transactions of the Royal Society of Tropical Medicine and Hygiene'' 96(6):640-643.][Mitchell, P.D., E. Stern, and Y. Tepper (2008) Dysentery in the crusader kingdom of Jerusalem: An ELISA analysis of two medieval latrines in the City of Acre (Israel). ''Journal of Archaeological Science'' 35(7):1849-1853.] ), and DNA sequencing[Aufderheide, A.C., W. Salo, M. Madden, J. Streitz, J. Buikstra, F. Guhl, B. Arriaza, C. Renier, L.E. Wittmers, Jr., G. Fornaciari, and M. Allison (2004) A 9,000-year record of Chagas' disease. ''Proceedings of the National Academy of Sciences of the USA'' 101(7):2034-2039.][Dittmar, K., U. Mamat, M. Whiting, T. Goldmann, K. Reinhard, and S. Guillen (2003) Techniques of DNA-studies on prehispanic ectoparasites (''Pulex'' sp., Pulicidae, Siphonaptera) from animal mummies of the Chiribaya culture, southern Peru. ''Memórias do Instituto Oswaldo Cruz'' 98(Suppl 1):53-58.][Iñiguez, A.M., K. Reinhard, M.L.C. Gonçalves, L.F. Ferreira, A. Araújo, A.C.P. Vicente (2006) SL1 RNA gene recovery from ''Enterobius vermicularis'' ancient DNA in pre-Columbian human coprolites. ''International Journal for Parasitology'' 36(13):1419-1425.][Loreille, O., E. Roumat, O. Verneau, F. Bouchet, and C. Hänni (2001) Ancient DNA from ''Ascaris'': Extraction, amplification, and sequences from eggs collected in coprolites. ''International Journal for Parasitology'' 31(10):1101-1106.][Raoult, D., D.L. Reed, K. Dittmar, J.J. Kirchman, J.M. Rolain, S. Guillen, and J.E. Light (2008) Molecular identification of lice from pre-Columbian mummies. ''Journal of Infectious Diseases'' 197(4):535-543.][Zink, A.R., M. Spigelman, B. Schraut, C.L. Greenblatt, A.G. Nerlich, and H.D. Donoghue (2006) Leishmaniasis in ancient Egypt and Upper Nubia. ''Emerging Infectious Diseases'' 12(10):1616-1617.] are used to identify the source of these chemical remains, often to the species level.
Fundamental questions
Archaeoparasitological studies have provided information on many fundamental archaeological, historical, and biogeographical questions. These questions may be grouped into the following broad categories: past dietary and farming practices,[Reinhard, K.J., R.H. Hevly, and G.A. Anderson (1987) Helminth remains from prehistoric Indian coprolites on the Colorado Plateau. ''Journal of Parasitology'' 73(3):630-639.] animal domestication,[Nansen, P. and R.J. Jørgensen (1977) Fund af parasitaeg i arkaeologisk materiale fra det vikingetidige Ribe / Parasite eggs identified in material from archaeological excavations in Ribe (the Viking Age). ''Nordisk Veterinaermedicin'' 29(6):263-266.][Sadler, J.P. (1990) Records of ectoparasites on humans and sheep from Viking-age deposits in the former western settlement of Greenland. ''Journal of Medical Entomology'' 27(4):628-631] migration patterns,[Araújo, A., K.J. Reinhard, L.F. Ferreira, and S.L. Gardner (2008) Parasites as probes for prehistoric human migrations? ''Trends in Parasitology'' 24(3):112-115.][Montenegro, A., A. Araújo, and M. Eby (2006) Parasites, paleoclimate, and the peopling of the Americas: Using the hookworm to time the Clovis migration. ''Current Anthropology'' 47(1):193-200.] climate change,[Araújo, A., A. Rangel, and L.F. Ferreira (1993) Climatic change in northeastern Brazil: Paleoparasitological data. ''Memórias do Instituto Oswaldo Cruz'' 88(4):577-579.] sanitation|sanitary practices,[Faulkner, C.T. (1991) Prehistoric diet and parasitic infection in Tennessee: Evidence from the analysis of desiccated human paleofeces. ''American Antiquity'' 56(4):687-700.] cultural contacts,[Araújo, A., L.F. Ferreira, U. Confalonieri, and M. Chame (1988) Hookworms and the peopling of America. ''Cadernos de Saúde Pública'' 4(2):226-233.][Bouchet, F., S. Harter, J.C. Paicheler, A. Araújo, and L.F. Ferreira (2002) First recovery of ''Schistosoma mansoni'' eggs from a latrine in Europe (15th-16th centuries). ''Journal of Parasitology'' 88(2):404-405.] ethnomedicine,[Reinhard, K.J., J.R. Ambler, and M. McGuffie (1985) Diet and parasitism at Dust Devil Cave. ''American Antiquity'' 50(4):819-824.][Chaves, S.A. de M. and K.J. Reinhard (2006) Critical analysis of coprolite evidence of medicinal plant use in Piaui, Brazil. Palaeogeography Palaeoclimatology Palaeoecology 237(1):110-118.] and the overall health of various human societies[Sallares, R. (2002) Malaria and Rome: A History of Malaria in Ancient Italy. Oxford University Press: Oxford, England. xv, 341 pp.]. Archaeoparasitology data, combined with our knowledge of present host-parasite associations, also contributes to our understanding of the co-evolution of human host-parasite interactions.[Ashford, R.W. (2000) Parasites as indicators of human biology and evolution. ''Journal of Medical Microbiology'' 49:770-771.] Our understanding of the geographic origins, evolution and biogeography of the parasites themselves and human diseases associated with them[Hugot, J.P., K.J. Reinhard, S.L. Gardner, and S. Morand (1999) Human enterobiasis in evolution: Origin, specificity and transmission. ''Parasite'' 6(3):201-208.][Nozais, J.-P. (2003) The origin and dispersion of human parasitic diseases in the Old World (Africa, Europe and Madagascar). ''Memórias do Instituto Oswaldo Cruz'' 98(Suppl 1):13-19][Hoberg, E.P. (2006) Phylogeny of ''Taenia'': Species definitions and origins of human parasites. ''Parasitology International'' 55(Suppl):s23-s30.] has also benefitted tremendously from archaeoparasitological studies.
References
Category: Archaeology
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