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Archaeoparasitology in North America.

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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.
@
1990 WILEY-LISS. INC
146
K.J. REINHARD
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,
NORTH AMERICAN PARASITISM
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
e"
147
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,
148
K.J. REINHARD
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
711
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
NORTH AMERICAN PARASITISM
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.
149
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
150
K.J. REINHARD
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-
NORTH AMERICAN PARASITISM
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
151
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-
152
K.J. REINHARD
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
NORTH AMERICAN PARASITISM
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,
153
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.
154
K.J. REINHARD
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.
NORTH AMERICAN PARASITISM
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
155
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-
156
K.J. REINHARD
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
157
NORTH AMERICAN PARASITISM
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-
158
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
NORTH AMERICAN PARASITISM
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~).
159
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
160
K.J. REINHARD
(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.
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