gastrointestinal cestodes and nematodes of coyotes from

43
Eastern Illinois University e Keep Masters eses Student eses & Publications 1981 Gastrointestinal Cestodes and Nematodes of Coyotes from Southeastern Illinois Valerie Keener Eastern Illinois University is research is a product of the graduate program in Zoology at Eastern Illinois University. Find out more about the program. is is brought to you for free and open access by the Student eses & Publications at e Keep. It has been accepted for inclusion in Masters eses by an authorized administrator of e Keep. For more information, please contact [email protected]. Recommended Citation Keener, Valerie, "Gastrointestinal Cestodes and Nematodes of Coyotes from Southeastern Illinois" (1981). Masters eses. 3019. hps://thekeep.eiu.edu/theses/3019

Upload: others

Post on 10-Feb-2022

2 views

Category:

Documents


0 download

TRANSCRIPT

Eastern Illinois UniversityThe Keep

Masters Theses Student Theses & Publications

1981

Gastrointestinal Cestodes and Nematodes ofCoyotes from Southeastern IllinoisValerie KeenerEastern Illinois UniversityThis research is a product of the graduate program in Zoology at Eastern Illinois University. Find out moreabout the program.

This is brought to you for free and open access by the Student Theses & Publications at The Keep. It has been accepted for inclusion in Masters Thesesby an authorized administrator of The Keep. For more information, please contact [email protected].

Recommended CitationKeener, Valerie, "Gastrointestinal Cestodes and Nematodes of Coyotes from Southeastern Illinois" (1981). Masters Theses. 3019.https://thekeep.eiu.edu/theses/3019

TI r F:SIS H EPRODUCTION CERTIFICATE

TO: Graduate Degree Candidates who have written formal theses.

SUBJECT: Permission to reproduce theses.

The University Library is rece 1vtng a number of requests from other institutions asking p ermission to reproduce dissertations for inclu8ion in their library holdings. Although no copyright laws are involved, we feel that professional courtesy demands that permission be obta ined from the author before we allow theses to be copied.

Please sign one of the following statements:

Booth Library of Eastern Illinois University has my permission to lend my thesis to a reputable college or un iversity for the purpose of copying

it for inclusion in that institution's library or research holdings.

_!?J � fff7/ Date

I respectfully request Booth Library of Eastern Illinois Universi ty not allow my thesis be reproduced because

-----�----------

-·--··· ---·-·-----------------------------------

Date Author

in

GASTROINTESTINAL CESTODES AND NEMATODES OF

COYOTES FROM SOUTHEASTERN ILLINOIS (TITLE)

BY

Valerie Keener

THESIS

SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF

Master of Science

IN THE GRADUATE SCHOOL, EASTERN ILLINOIS UNIVERSITY CHARLESTON, ILLINOIS

981J YEAFl

I HEREBY RECOMMEND THIS THESIS BE ACCEPTED AS FULFILLING THIS PART OF THE GRADUATE DEGREE CITED ABOVE

'=I �� ,>?( L/ A""' 4.-ff '' r I DATE

ABSTRACT: Gastrointestinal tracts of 45 coyotes, Canis latrans, from

southeastern Illinois were examined for cestodes and nematodes. The

following parasites and incidences (%)were observed: Taenia sp. (71.1);

Ihysaloptera sp. (53 . 3); Ancylostoma sp. (20.0); Ascarididae (11.1);

Thelazia sp. (2.2); Trichuris sp. (2.2); Uncinaria sp. (2.2). Ancylostoma

sp. occurred significantly more often in juveniles than adults. Parasites

averaged low numbers per coyote, and no ulceration or blood in the

tracts was seen.

ii

ACKNCMLEDGEMENTS

Coyotes were obtained from A. Hasler and D. Fender. J. H. Keener,

J. B. Keener, and s. Priebe assisted with samples and data. M. L. Keener

typed the manuscript. G. Hubert, Jr. radiographed teeth and aged the

coyotes. B. T. Ridgeway, aided by R. Kuye, confirmed the parasite

identifications. K. Kruse offered statistical advice. Members of my

committee, R . D • .Andrews, L . Durham, L. B . Hunt, and B. T. Ridgeway,

critically read the manuscript.

iii

TABLE OF ·CONTENTS

ABSTRACT • • • • • • • • • • • • • • • • • • • • ii

ACKNCJvlLEroEMENTS • • • • • • • • • • • • • • • • • iii

LIST OF TABLES • • • • • • • • • • . . • • •• • • • • v

INTRODUCTION • • • • • • • • • • • • • • • • • • • 1

MN.l'HODS • • • • • • • • • • • • • • • • • • • • • 1

RESULTS • • • • • • . . • • • • • • • • • • • • • 0 3

DISCUSSION • • • • • • • • • • • • • • • • • • • • 3

LITERATURE CITED • • • • • • • • • • • • • • • • • • 7

LITERATURE REVIEW • • • • • • • • • • • • • • • • • 11

LITERATURE CITED • • • • • • • • • 0 • • • • • • • 32

iv

LIST OF TABLES

Table 1 . Incidence of gastrointestinal parasites by sex and age in

coyotes from southeastern Illinois • • • • • • •

Table 2 . Comparison of parasites found in this study, excluding

Thelazia sp., to other m.idwestern studies •

Literature Review

• •

Table 1 . Gastrointestinal helminths, excluding trematodes,

previously found in coyotes • • • • • •

v

• •

4

• 5

12

Coyotes (Canis latrans ) have been·shown by several researchers,.

including Andrews and Boggess (1 978) and Berg and Chesness (1978) , to

consume a wide variety of materials. 'lbeir predatory and scavenging

habits expose them to possible parasite infections from many sources.

Parasite surveys of coyotes have been conducted widely west of the

Mississippi River in the United States including the following states

and reports: California (Voge 1 955) ; Iowa (Franson et al. 1 978) ;

Kansas (Ameel 1 955 ; Gier and Ameel 1 959 ; Gier 1968); Minnesota

(Erickson 1 944) ; Oklahoma (Self and McKnight 1950; Esch and Self 1965);

Texas ('Il1.ornton et al. 1974; Pence and Meinzer 1979); Utah (Butler and

Grundmann 1 954 ; Conder and Loveless 1978). 'Il1.e Canadian provinces of

Alberta, Manitoba, and Ontario were the site of coyote parasite surveys

by Holmes and Podesta (1 968), Samuel et al. (1978), and Freeman et al.

(1 961 ) , respectively. This study of gastrointestinal cestodes and nematodes

from southeastern Illinois was undertaken to add to our knowledge of the

ecology of an increasing coyote population east of the Mississippi River.

METHODS

Skinned and frozen coyote carcasses were obtained from two fur

buyers . 'lhese coyotes were considered to be representative of an area

within 50-mile radii of the two buyers: Louisville (Clay County ) and

Nob'le (Richland County), Illinois. The area is composed of gently rolling

hills with scattered woodlots, most wooded areas occurring in river

bottomland. Agriculture is the major land use.

Carcasses were thawed, and complete gastrointestinal tracts were

removed, placed in plastic bags, and refrozen. Tracts were thawed and

divided into sections- esophagus, stomach , duodenum, jejunum, ileum, and

1

2

cecum.and large intestine . Each section was opened , flushed , and

scraped; contents were searched for parasites . All worms were stored in

vials of alcohol-formalin-acetic acid (AFA).

Some cestode scoleces were stained with Grenacher's Alcoholic Borax­

Carmine, cleared with oil of wintergreen, and mounted on slides in

Permount . Cestodes not requiring staining for identification were stored

in AFA. Nematodes were cleared in lactophenol and stored in 7afo ethanol,

3% glycerine . Identification of cestodes was based on works by Morgan

and Hawkins (1 951 ), Wardle and McLeod ( 1 952) , and Yam.a.guti (1959) .

Nematode identification was made by utilizing Morgan and Hawkins (1 951 ),

Yam.a.guti (1 961 ) , Yorke and Maplestone (1 969), Chabaud (1975), and

Levine (1980).

Age of the coyotes was determined by the size of the canine pulp

cavity . Mandible tips were macerated until the canines fell out . One

canine per coyote was radiographed , with aging following the technique

of Grue and Jensen (1 976) . Juveniles were separated from adults by

applying the following characters: pulp cavity over � the width of

tooth= juvenile; pulp cavity less than� the width of tooth = adult;

pulp cavity approximately equal to � the width of tooth = animal caught

early in second year (about 1 6 months ) .

Significance of relations between age and/or sex of coyotes and

infection was tested by chi-square (with Yate's correction factor ) .

Although expected frequencies were often small, Harnett (1975) states

that the chi-square test may still be u sed.

Representative specimens of parasites obtained in this study have

been deposited in the National Parasite Collection (u .s . n . A . ) at

3

Beltsville, Maryland.

RESULTS

Forty-five coyotes were collected from 20 December 1 980 to 17

January 1981 . Coyotes were aged and sexed as follows: 11 juvenile males;

9 adult males; 20 juvenile females; 5 adult females . Thirty-eight (84.4%)

of the coyotes were infected with parasites from one genus of cestode and

five genera and one family of nematodes (Table 1). The only significant

relation between infection and age and/or sex of the coyote s was that

significantly (P<o.05) more juveniles (9/31) were infected with

Ancylostoma sp. than were adults (0/14).

Multiple infections were found in 65.aofo (25/38) of the infected

coyotes . Twenty coyotes had cestode-nematode infections, while four

had only multiple nematode infections . One coyote contained repre sentatives

of four parasite groups, Taenia sp ., Physaloptera sp., Ancylostoma sp.,

and Uncinaria sp . No significant relations were found between infection

and age and/or sex of the coyotes with re spect to multiple infections,

three or more group infections, or the most common multiple infection

(11/25), Taenia sp . and Ihysaloptera sp .

DISCUSSION

Taenia sp. and Ihysaloptera sp . were the most prevalent parasites

found in this and other midwestern studies (Table 2). Taenia sp. utilize

a variety of rodents and other kind� of herbivores as intermediate hosts

( Olsen 1 974) . Andrews and Bogge ss (1 978) show that rabbits and mice

makeup a considerable amount of the coyote's diet-in Iowa . Observation

of stomach contents in this study indicated the presence of many rabbits

and mice . Other epidemiologic conditions being suitable , it is not

Table 1 . Incidence of Gastrointestinal Parasites by Sex and Age in Coyotes frM Southeastern Illinois

Cestodes

Taenia sp.

Nematodes

Fhysaloptera sp.

Ancylostoma sp.

Ascarididae

Thelazia sp.

Trichuris sp.

Uncinaria sp.

*Sample size.

**Based on scoleces.

Sex

Male (20)*

75.0

45 .0

25. 0

10.0

o.o

o.o

5.0

Female (25)

68.0

60. 0

1 6 .0

1 2 . 0

4. 0

4. 0

o.o

Incidence (%)

Age

Adult (14)

Juvenile (31 )

64.3 74.2

42 . 9 5a.1

o.o 29 .0

1 4 .3 9 .7

o.o 3 . 2

o.o 3.2

o.o 3.2

Total (45)

71 . 1

53 . 3

20. 0

11.1

2.2

2 . 2

2.2

Average Parasite Load

6.6**

3.a

3.3

6 .2

1 . 0

1 4.0

1 .0

Table 2. Comparison of Incidence of Parasites Found in this Study, Excluding Thelazia sp., to Other

Midwestern Studies.

Cestodes

Taenia sp.,

Nematodes

Physalo-ptera sp.

Ancylostoma sp.

Ascarididae

Trichuris sp.

Uncina.ria sp.

IL (45)*

71.1

53.3

20.0

11.1

2.2

2.2

Incidence {%)

IA a (144)

86.8

66. 7

50. 0

3a.9

6.9

o.o

a-Franson et al. 1978; b-Ameel 1955; c-Gier and Ameel 1959.

*Sample size.

KSb (1142)

92.8

60. 4

19.s

19.3

11. 2

o.o

KSc

(1850)

95

51

25

33

6

0

V1

6

surprising that Taenia sp. is prevalent in the coyotes studied. Fhysaloptera

sp. utilize insects of the Orders Orthoptera and Coleoptera as intermediate

hosts and the amphibians, birds and mammals that feed on these insects

may serve as transport hosts (Petri and Ameel 1 950). Coyotes include the

transport hosts in their diet and probably a considerable number of

appropriate insects as well, so the likelihood of transmission i.s great.

One female Thelazia sp., a parasite of the eye, was found in one

coyote's gastrointestinal tract. Possible hosts of Thelazia sp. in Illinois

are cattle, horses, dogs, coyotes, man , cats, and wild birds (Olsen 1 974).

Presence o f the worm in th e gastrointestinal tract must b e considered

accidental. It may have been ingested by the coyote or it may represent

an unusual site for the parasite in its host.

Ancylostoma sp. was found in significantly more juveniles than adults.

Adult coyotes may have been partially immunized by prior infections and

are therefore more resistant, as is the case with dogs ( Levine 1980).

Also Miller (1 965) has demonstrated that true natural age immunity in

canines may exist. These immunologic factors coupled with the opportunity

for very young animals to acquire prenatal and transmamma.ry infections

(Clapham 1 962; Stone and Girardeau 1 968) , in addition to the more common

infections by skin penetration and ingestion, may account for the numbers

of juveniles infected.

The small average parasite load of Illinois coyotes (Table 1 ) suggests

that their health is not being severely affected. Personal observation

also indicated that the coyotes appeared healthy, i.e. no ulceration or

blood in the tracts was seen. Low parasite loads also resulted in the

presence of few male nematodes which inhibited identification beyond genus.

1

LITERATURE CITED

Ameel, D. J. 1955. Parasites of the coyote , Canis latrans Say, in

Kansas. J. Parasit. 41: 325.

Andrews , R. D. , and E. K. Boggess. 1978. Ecology of coyotes in Iowa.

Pages 249-265 !B,M. Bekoff, ed. Coyotes: biology, behavior, and

management. Academic Press , New York.

Berg, W. E., and R. A. Chesness. 1978. Ecology of coyotes in northern

Minnesota. Pages 229-247 !!!, M. Bekoff , ed. Coyotes: biology,

behavior, and management. Academic Press, New York.

Butler , J . M., and A. W. Grundmann . 1954. 'llle intestinal helminths of

the coyote Canis latrans Say in Utah. J. Parasit. 40: 440-443.

Chabaud, A. G. 1975. Keys to genera of the order Spirurida. Part 1.

Camallanoidea, Dracunculoidea, Gnathostomatoidea , Hlysalopteroidea,

Rictularioidea, and 'lllelazioidea. In cm keys to the nematode

parasites of vertebrates. Com • .Agric. Bur., Farnham Royal, Bucks.,

Eng;tand.

Clapham, P. A. 1962. Pre-partum infestation of puppies with Ancylostoma

caninum. Vet. Rec. 74: 754-755·

Conder, G . A., and R. M. Loveless. 1978. Parasites of the coyote in

central Utah. J . Wldl. Dis. 14: 247-249.

Erickson , A. B. . 1944. Helminths of Minnesota canidae in relation to food

habits and a host list and key to the species reported from North

America. Amer. Midl. Nat. 32: 358-372. -

Esch , G . W. , and J. T. Self. 1965. A critical s·tudy of the taxonomy of

Taenia pisiformis Bloch, 1780 ; Mu.lticeps multiceps { Leske , 1780);

and H.vdatigera taeniaeformis Batsch, 1786. J . Parasit. 51: 932-937.

8

Franson, J. c., R. D. Jorgenson, E. K. Boggess, and J. H. Greve. 1978.

Gastrointestinal parasitism of Iowa coyotes in relation to age .

J. Parasit. 64: 303-305.

Freeman, R. S•, A. Adorjan, and D. H. Pimlott. 1961. Cestodes of wolves,

coyotes, and coyote-dog hybrids in Ontario. Can. J. Zool. 39:

527-532.

Gier, H. T. 1968. Coyotescin Kansas. rev. ed. A.gric. Exp. Sta. Kans.

State Univ. Agric. and Appl. Sci., Manhattan, Kansas, Bull. 393.

118 pp.

Gier, H . T., and D. J. Ameel. 1959. Parasites and diseases of Kansas

coyotes. Agric. Exp. Sta. Kans. State Univ. Agric. and Appl. Sci.,

Manhattan, Kansas, Tech. Bull. 91. 34 PP•

Grue, H . , and B. Jensen. 1976. Annual cementum structures in ca.nine

teeth in arctic foxes (Alopex 1§!.gOpus (L.)) from Greenland and

Denmark. Dan. Rev. Game Biol. 10(3): 1-12.

Ha.matt, D. L. 1975. Introduction to statistical methods, 2nd ed.

Addison-Wesley Publ. Co. , Reading, Massachusetts. 522 PP•

Holmes, J . c., and R. Podesta. 1968. The helminths of wolves and coyotes

from the forested regions of Alberta. Can. J. Zool. 46: 1193-1204.

Levine, N. D. 1980. Nematode parasites of domestic animals and man,

2nd ed. Burgess Publ. Co., Minneapolis, Minnesota. 477 pp.

Miller, T. A. 1965. Influence of age and sex on susceptibility of dogs

to primary infection with Ancylostoma caninum. J. Parasit. 51:

701-704.

Morgan, B. B. , and P . A. Hawkins. 1951. Veterinary helminthology.

Burgess Publ. Co., Minneapolis, Minnesota. 400 pp.

9

Olsen, O. W. 1974. Animal parasites: their life cycles and ecology,

3rd ed. Univ. Park Press, Baltimore , Maryland. 562 pp.

Pence , D. B., and w. P. Meinzer. 1979. Helminth parasitism in the

coyote, Canis latrans, from the Rolling Plains of Texas. Int. J.

Parasit. 9: 339-344.

Petri , L. H., and D. J. Ameel., 1950. Studies on the life cycle of

Ih:ysaloptera �Hall and Wigdor, 1918 and Physaloptera praeputialis

Linstow, 1889. J. Parasit. 36(sup.):· 40.

Samuel, w. M., s. Ramalingam, and L. N. Carbyn. 1978. Helminths in

coyotes, wolves, and red foxes of southwestern Manitoba. Can. J.

Zool. 56: 2614-2617.

Self', J. T., and T . J. McKnight. 1950. Platyhelminthes from fur bearers

in the Wichita Mountains Wildlife Refuge, with especial reference to

Oochoristica spp. Amer. Midl. Nat. 43: 58-61.

Stone , w. M., and M. H. Girardeau. 1968. Transma.mmary passage of

Ancylostoma caninum larvae in dogs. J. Parasit. 54: 426-429.

'lhornton, J. E., R. R. Bell, and M . J. Reardon. 1974. Internal parasites

of coyotes in southern Texas. J. Wldl. Dis. 10: 232-236.

Voge, M. 1955. A list of cestode parasites from California mammals .

Amer. Midl. Nat. 54: 413-417.

Wardle, R. A. , and J. A. McLeod. 1952. The zoology of tapeworms. Univ.

Minn. Press , Minneapolis, Minnesota. 780 PP•

Yamaguti , s. 1959. Systema helminthum. Vol . II.- The cestodes of

vertebrates. Interscience Publ., New York. 860 PP•

Ya.maguti, s. 1961. Systema helminthum. Vol. III. The nematodes of

vertebrates. Parts 1 and 2 . Interscience Publ., Inc., New York.

10

Yorke, w., and P. Maplestone . 1 929 . The nematode parasites of vertebrates.

(Reprint 1 969) Hafner Publ. Co ., New York. 536 PP•

11

LITERATURE REVI:EM

The coyote (Canis latrans ) has been surveyed for cestodes and nematodes

in several states west of the Mississippi River in the United States and

some Canadian provinces. Seven genera and 20 species of cestodes have

been identified, as have 1 7 genera and 1 7 species of nematodes, and 2

genera and 2 species of Aca.nthocephala (Table 1 ) .

Table 1 contains specific studies of gastrointestinal parasites of

coyotes. These reports indicate that a number of parasites are commonly

found in coyotes, yet the effect of these parasites on the coyote has not

been well determined. This review sumroa.rizes available knowledge on the

more common parasites presented in Table 1 . The descriptions include

work from coyotes and domestic dogs (.Q.. familiaris).

Taenia pisiformi s is a tapeworm often found in coyotes . Gravid

proglottids from mature worms pass in the coyote ' s feces. If eggs

containing larvae are eaten by the proper intermediate host, such as a

rabbit, squirrel, rat, or other rodent, development to cysticeri continues

in the visceral cavity. After several weeks of development , these larvae

are capable of infecting coyotes when the host is eaten (Olsen 1 974) .

The infected coyote may have diarrhea for two or three days after the

initial infection. Abdominal irritation is sometimes apparent, as the

animal will roll or drag its abdomen over the ground. Anal irritation

from passing gravid proglottids is pre sumed when the animal assumes a

sitting position and drags its a.nus over the ground. If tremendous

numbers of worms are present, intestinal blockage and rupture may occur.

This information was compiled from Morgan and Hawkins (1 951 ) .

The hookworm most commonly reported from coyotes in the southern

Table 1. Gastrointestinal Helminths, Excluding Trematodes, Previously Found in Coyotes

Helminth* Location** Incidence

(%)

Phylum Platyhelminthes

Class Eucestoda

Order Dilepididea

Dipylidium caninum SI 1. 3

Type of Report and*** Geographical Location

case rept., UT II II KS

II II II

gen. rev.

II II

Author

Butler and Grundmann 1954

Ameel 1955

Gier and Aineel 1959

Leiby and Dyer 1971

Gier et al. 1978

*Classification based on Wardle et al. (1974) for cestodes and Levine (1980) for nematodes.

**Abbreviations used: Ce-cecum; Es-esophagus; In-intestines; LI-large intestine; SI-small intestine;

ST-stomach.

***Abbreviations used: case rept.-case report or survey; gen. rev.-mentioned in general review of

..... I\.)

diseases; sp. rev.-review of t:b.at specific disease or a specific parasite description.

Helminth Location Incidence Type of Report and Author

(%) G€ographical Location

Order Mesocestoididea

Mesocestoides carnivoricolus SI - sp. rev. Grundmann 1956

case rept., UT Grundmann 1958

gen. rev. Yamaguti 1959

!:!• corti SI - gen. rev., CA Voge 1955

case rept., KS Gier and .tuneel 1959

II " . ,. Gier 1968 ..... '-" .

gen. rev. J,eiby and Dyer 1971

" " 3ekoff 1977

" II Gier et al. 1978

57. ·5 case rept., TX Fence and :'leinzer 1979

M. kirbyi SI - sp. rev., CA Chandler 1944

gen. rev. :i!irickson 1944

10. 7 case rept., 'JT 3utler and Grundmann 1954

gen. rev., CA Voge 1955

Helminth Location Incidence Type of Report and Author

(%) Geographical Location

D.• kirbyi SI - gen. rev. , CA Ya.1laguti 1959

1 case rept., Alberta Holmes and Podesta 1968

18 !I II UT Conder and Loveless 1978

tl• lineatus SI - gen. rev. Leiby and Dyer 1971

Mesocestoides sp. SI - sp. rev . , KS ',food and F..aldiman 1957

case rept., Ontario .?ree�an et al. 1961

gen. rev. Gier et al. 1978 ...... �

Order Taeniidea

Schinococcus r:r-an�oGus SI 0.5 case :!'.'ept., Jntario Freenan et al. 1961

8 " " Alberta �olLles and Podesta 1968

4 " ,, C1\. Liu et al. 1970

gen. rev. Leiby and Dyer 1971

r• II Clc:r- et al. 1978

a case rept., ?fani toba Sa.�uel et al. 1978 ./

s. mul�iloculaLis SI 4.09 s p • rev. , IA, MN, : IT, Lei'::iy e-t al. 1970 '7'TT'1 :3D -·�,

Helminth

�· mul tilocularis

Hydatigera lvncis

li• taeniaefo::::mis

Nul ticens sp.

Taenia crassicens

!• hydatigena

Location

SI

SI

SI

SI

SI

SI

Incidence

(%)

23

11.4

o.o

31.1

2.6

Type of Report and Geographical Location

gen. rev.

II "

case rept., Hani toba

gen. rev., CA

sp. rev., OK

case re pt., MN

gen. rev.

case rept., Ontari o n II Manitoba

case rept., HN

gen. rev.

case rept., lJT " " "

" " Ontario II II Alberta

Author

Leiby and Dyer 1971

Gier et al. 1978

Samuel et al. 1978

Voge 1955

Esch and Self 1965

Erickson 1944

Erickson 1944

Freeman et al. 1961

Samuel et al. 1978

Erickson 1944

Erickson 1944

..... \.J1

Butler and Grundmann 1954

Grundmann 1958

7reeman et al. 1961

Holmes and Podesta 1968

Helminth Location Incidence

(%)

:r_. hydatigena SI -

7

:;g_. krabbei* SI 3.2

1

l• laticollis SI -

*Listed oy \:l2Xd2.e et al. (1974) as T. ovis kraobei. - ��- --------

Type of Report and Author Geographical Location

gen. rev. Leiby and Dyer 1971

case rept., UT Conder and Loveless 1978

gen. rev. Gier et al. 1978

case · re pt., Hani toba Samuel et al. 1978

case rept., YllI Erickson 1944

gen. rev. Erickson 1944

case rept., TJT :Butler and Grundmann 1954 _.. O'\

II " Ontario FTeeman et al. 1961

II " Alberta Holmes and Podesta 1968

gen. rev. Leiby and Dyer 1971

case rept., U'J.' Conder and Loveless 1978

gen. rev. Zrickson 1944

case rept., Ontario Freeman et al. 1961

gen. rev. Leiby and Dyer 1971

Helm in th Location Incidence Type of Report and Author

(%) Geographical Location

!• laticollis SI - gen. rev. Gier et al. 1978

!• macrocystis* SI - gen. rev. Leiby and Dyer 1971

!• ovis SI - gen. rev. Gier et al. 1978

!• pisifomis SI - gen. rev. Hall 1914 II " Hall 1929

39.3 case rept., HN Erickson 1944

gen. rev. Erickson 1944 ..... -.J

100 case rept., OK Self and McKnight 1950

92.s 11 " tr� 1\.0 P..meel 1955

11 II \J-r:L' Grundman..'1 1958

95 It " KS Gier and Ameel 1959

gen. rev. Yamaguti 1959

case rept., Ontario Freeman et al. 1961

spo rev., OK Esch and Self 1965

95 case rept., KS Gier 1 c;68

*Listed by \�ardle et al. (1974) as Eydatigera macrocYstis.

Helminth

:f.• pisif omis

!• pseudolaticollis*

i1 � � .=_. rueyi**

!· serialis

Location

SI

SI

SI

SI

Incidence

(%)

31

67

38.7

4 ') . /

2.7

Type of Report and

Geographical Location

case rept., Alberta

gen. rev.

case rept. , KS

gen. rev.

case rept., UT

gen. rev.

case rept., r·lani toba

" " TX

gen. rev.

case rept., '.·I;;

gen. rev.

case rept., T.JT

gen. rev.

sp. rev.

*Not listed as ;-l, sriecies oy Yamaguti (1959) or ·:!ardle et al. (1974).

**Listed as a species by Yanaguti (1959).

Author

Holmes and Podesta 1968

Leiby and Dyer 1971

Hirsch and Gier 1974

Bekoff 1977

Conder and Loveless 1978

Gier et al. 1978

Samuel et al. 1978

Fence and Meinzer 1979

Gier et alo 1978

Erickson 1944

Erickson 1944

� CD

:Butler and Grundmann 1954

Gier et al. 1978

Eall 1911

Helminth Location Incidence Type of Report and Author

(%) Geographical Location

!• serialis SI - sp. rev. Hall 1912

1.6 case rept., MN Erickson 1944

gen. rev. Erickson 1944

5.3 case rept., UT 3utler and Grundmann 1 954

gen. rev., CA Voge 1955

case rept., UT Gru...'1.d.mann 1958

gen. rev. Yamagiti 1959 � '°

sp. rev., CK 3sch ar.d. Self 1965

4 case rept., AlbeYta Ifolrr:es and Podesta i}68

gen .. -::ev. -- ·- d fy 1°r ueioy an � er / i

47 case rept., ;;rp Ccnder and Loveless 1978 J ...

gen. rev. Gier et al. 1978

4.0 case rept., �,.r ?ence and Heinze-r 1979 ...... \.

!• twi tchelli* SI 1 case rept., ·\lberta �i:ol:ne s anC. Podesta 1 c;6a

gen. rev. Gier et al. 1978

*Linted by '•ia-rc:'...e et al. (1974) as Multicens -twitchelli.

Helminth

Taenia sp.

Class Cotyloda

Order Diphyllidea

DiPhyllobothrium 3p.

Phylum Nematoda

Class Secernentasida

Order Strong::,rlorida

Location

SI

.�.,-;) l.

Family Ancylostaraatidae

Ancvlostorna caninum SI

Incidence

(%)

4.9

76

e6.s

9

5

3.2

Type of Report and GeogTaphical Location

case rept. , MN

gen. rev.

II II IA

case rept., UT II II IA " 11 Nanitoba

case rept., Alberta

gen. rev.

case re pt., I-TN

Author

Erickson 1944

::!:rickson 1944

Andrews and Boggess 1978

Conder and Loveless 1978

FTanson et al. 1978

Samuel et al. 1978

I\) 0

�olmes and Podesta 1968

Hall 1914

Erickson 1944

Helminth Location Incidence Type of Report and Author

(%) Geographical Location

Ancylostoma caninum SI - gen. rev. Erickson 1944

19.s case rept., KS Arneel 1955

25 " " " Gier and Arneel 1959

28 II II " Gier 1968

1 " II Alberta Holmes and Podesta 1968

" . " KS Hirsch and Gier 1974

89.3 " " TX Mitchell and Beas om 197 4 I\) ..... 100 " II " Thornton et al. 1974

gen. rev. 3ekoff 1977

" " IA Andrews and 3oggess 1978

12 case rept., U'I' / Conder and Loveless 1978

50.0 " II IA �anson et al. 1978

gen. rev. Gier et al. 1978

89.3 case rept., TX Pence and Meinzer 1979

gen. rev. Levine 1980

Helminth Location Incidence Type of Report and Author (%) Geographical Location

Uncinaria stenocephala SI 16 case rept., Alberta Holmes and Podesta 1968

g-en. rev. Gier et al. 1978

28 case rept., Manitoba Samuel et al. 1978

Family Trichostrongylidae

Molineus patens SI - gen. re''• Erickson 1944

I I II Gier et al. 1978

Order Ascaridorida fl.)

F2.mily Ascarididae fl.)

Toxascaris leonina '�T .J� - g-en. rev • Erickson 1944

37.3 case rept., UT Butler and Grundrr.anr: 1954

:9.3 II " KS .;.meel 1955

" II U1I' Grundmann 1957

33 II II KS Gier and Ameel 1959

g-en. rev. -f'amaguti 1961

� 'Z case rept., t<S '::;.ier 1968 ,;_,

52 II " Alberta 2olmes and Podesta·1968

Helm in th Location Incidence Type of Report and Author

(%) Geographical Location

!• leonina SI - case rept., KS Hirsch and Gier 1974

gen. rev. Bekoff 1977 II II Hi. Andrews and 3oggess 1978

29 case rept., 'J� Conder and Loveless 1978

38.9 II II IA Franson et al. 1978

gen. rev. Gier et al. 1978

65 case re pt., Manitoba Sa;:iuel et alo 1978 I\) \.)'j

1.3 II " TX Fence and Heinzer 1979

gen. rev. Levine 1980

'?oxascaris sp. - gen. revo Erickson 1944

To:rncara canis SI - gen. rev. 28.ll 1914

1 ' I.) case rept., TjT ::3utler and Grundmann 1954

1 " " ... .\lberta ::olir.e:.; and Podesta 1'.?'68

6 " II TjT Conder and Loveless 1?78

gen. rev. Gier et al. 1978

Helminth Location Incidence

(%)

Toxocara canis SI -

Family Subuluridae

Subulu....� sp. SI 28

Family 0x'JUridae

Denna tOX"� veligera SI -

o.6

Passalurus nonannulatus SI -

3yphacia sp. St o.6

Ox<.[Uridae St 15

Order Spirurorida

Family Spi!."Uridae

Easto:phorus ;-;:uri::: SI -

Type of Report and Geographical Location

gen. rev.

case rept., TX

case rept., UT II II TX

gen. rev.

case rept., UT

gen. rev.

II II

case rept., TX

case re pt., ·u

gen. rev.

Author

Levine 1980

Pence and !1einzer 1979

Butler and Grundmann 1954

Pence and Meinzer 1979

Erickson 1944 f\) �

Butler and Grundmann 1954

Ya.maguti 1961

Gier et al. 1978

Pence and 'Neinzer 1979

'"Ohornton et al. 1974

Zrickson 1944

Helminth Location Incidence Type of Report and Author

(''/) /'O Geographical Location

Hastophorus muris SI - gen. revo Yamaguti 1961

!I II Gier et al. 1978

Protosnirura numidica SI - gen. rev. Erickson 1944

1.3 case rept., UT "Butler and Grundmann 1954

gen. rev. Ya..11aguti 1961

II II Gier et al. 1978

?a.mily 'I'helaziidae I'\) \,JI

'Rictularia cahirensis SI - sp. rev., CO l-Iall 1914

gen. rev. Erickson 1944

" " Yamaguti 1961

36.6 case rept., :::'X Pence and Meinze r 197S·

Spiroce:r."ca luni Es 23 case rept., I'X 'I'hornton et al. 1974

gen. rev. 3ekoff 1977

" " Gier et al. '.978

7 case :r."ept., 3( Pence and 1·1einzer 1 ':79

Helminth Location Incidence (%)

Spirocerca luni Es -

Family PJ1ysalopteridae

I'hYsalol'tera fe'J.idis St and SI -

f• praenutialis St -

f. pseudoprC?.epqtialis St -

f·� St 4.6

50.0

60

Physaloptera sp. St 12.3

Type of Report and Geographical Location

gen. rev.

gen. rev.

case rept. , KS

gen. rev.

case rept., NN

gen. rev.

case rept., XS

gen. rev.

+ •r.-. case rep.,., i�:J

gen. rev.

" "

J.. rrl "" "' case rep v• , lA

. g-en. rev.

+ .. ,.. ...... case rep "o, .·,�i

Author

Levine 1980

Ya.maguti 1961

Ameel. 1955

Yamaguti 1961

;;:;rickson 1944

�rickson 1944 I'\) O'\ Am.eel 1955

"lanaguti 1961

Gier 1968

:.::ckoff 1977

G:ie:::- et al. 1978

Fence and I1einzer 1979

Levine 1980

T:ri�kson 1944

Helminth

Ph.ysaloptera sp.

Class )_denc;horci.sids.

Order Dor:,rlai.":lorida

Family .='richuridae

Location

St

Capillaria sp. LI

'::richuris vul TJL Ce

Incidence

(%)

60.4

51

23

6

66.7

'7 i

') c.

1:.2

"Z (") ,/. /

Type of Report and

Geographical Location

gen. rev.

case rept., KS II !I II

II II II

II II TX

gen. rev., L\

case rept. , UT II II u

II II i-lanitoba

case rept., r1:ani toba

case rept., "70 ,\.o.j

II 11 II

Author

Erickson 1944

A.'r!eel 1955

Gier and Ao.eel 1959

Hirsch and Gier 1974

�ho:rnton et al. 1974

Andrews and Boggess 1978

Conder and Loveless 1978 I\) -l

_,_-·::-ar.son et al . 1978

:�c::n-riel et al. 1978

�-�a::-�uel et al. 1978

..;\ .. ::t0JCl 1955

Gier and 11.IDeel 1 �'59

Helminth Location Incidence Type of Report and Author

(%) Geographical Location

-

Trichuris vulpis Ce 6 case rept., KS Gier 1968

gen. rev. Bekoff 1977

II " IA Andrews and Boggess 1978

6.9 case rept., IA Franson et al. 1978

gen. rev. Gier et al. 1978

II It Levine 1980

Pence and Meinzer 1979 I\)

case rept., TX CX> Trichuris sp.

Family Trichinellidae

Trichinella spiralis 12� 5 sp. rev. , AK Rausch et al. 1956

0.5 II II CQ Olsen 1 960

4.3 " II IA Zimmerman and Hubbard 1969

gen. rev. Zimmerman 1971

case rept., British Schmitt et al. 1 976 Colombia

gen. rev. Gier et al. 1978

Helminth Location Inc1d.ence Type of Report and Author (%) Geographical Location

Ihylum Acanthocephala

Oncicola canis SI - sp . rev. , TX Price 1 928

gen. rev. Erickson 1 944

sp. rev. Van Cleave 1953

gen. rev. Yamaguti 1 963

1 1 I I DeGuisti 1 971 I\) II II Bekoff 1 977 \.0

II II Gier et al . 1 978

26 case rept. , TX Pence and Meinzer 1 979

Pachysentis canicola In 22 . 7 case rept . , TX Pence and Meinz er 1 979

30

half of their range is Ancylostoma caninum. The life history is

relatively simple (Olsen 1974) . Infection may occur in several ways,

such as by ingestion of larvae directly or in food, skin penetration,

.fetal infection, or from the colostrum or milk , but ingestion is the

most frequent manner of infection. Partially embryonated eggs hatch on

the soil and undergo a series of molts for about a week resulting in the

third stage infective larvae . If swallowed by the coyote , the larvae

molt in the stomach and shortly thereafter migrate to the small intestine

where the final molt occurs and maturity is reached in about five weeks .

The major problem with hookworms is that the adults feed on blood.

Levine ( 1980) reports a mean blood loss of 0. 07 ml/worm/day with a

maximum of 0. 12 ml./worm/day. Young dogs and dogs on inadequate diets

seem to be most susceptible to the worms ; there is some evidence that

dogs gain imnrunity as they grow older (Morgan and Hawkins 1 951 ) .

Necrosis may occur at the point of hookworm attachment. Blood loss

continues after the worm has moved to a new location because of the

presence of an antic oagulant ( Olsen 1974). Levine ( 1 980) lists anemia,

emaciation, edema , weakness , listlessness , rough hair coat, foul breath ,

and diarrheic feces which are bloody or tarry as signs of hookworm

infection.

Toxascaris leonina is an ascarid commonly found in coyotes . As

Levine ( 1980) indicates , coyotes can be infected by ingesting eggs -

containing second stage larvae. The eggs hatch and the larvae spend

about 10 days in the intestinal wall, after which they return to the

lumen and mature . If an infected mouse , a conunon transport host, is

eaten, digestion of the mouse releases larvae, usually third s tage ,

31

which develop in the intestinal lumen. Animals with :;r_. leonina

infections often show restlessness and twitching. Infected pups may

show unthriftiness , enlargement of abdomen, digestive disturbances , dull

hair coat , alternating periods of diarrhea and constipation , vomiting,

and sweetish breath. If enough worms are present , intestinal blockage

and/or rupture may occur.

Members of the genus Ihysaloptera , especially P. m_, are

spiruroids often found in coyote s . Th e intermediate hosts o f Physaloptera

sp . are insects of the Orders Orthoptera and Coleoptera. The intermediate

hosts for f• m, are the German cockroach (Blatella germanica) , the

field cricket (Gryllus assimilis ) , the flour beetle (Tribolium confusum) ,

and ground beetles (Harpalus sp. ) . Development of the worms in coyotes

is the same regardless of whether the infective third stage larvae are

acquired by swallowing infected insects or by eating parasitized pa.ratenic

hosts such as frogs, birds , or mice . Adults develop in the stomach ,

duodenum, or both ( Olsen 1 974) . As these worms attach to the mucosa ,

they may cause small erosions , gastritis , enteritis , and mucus formation

(Levine 1 980) . Gier and Alneel ( 1 959 ) report that these spiruroids feed

largely on blood , and that laboratory infections of coyote pups with 50

or more P. rn regularly caused anemia and poor growth .

Whipworms, Trichuris sp . , are parasites of the coyote cecum. Levine

( 1 980) reports that coyotes become infected by directly inge sting eggs

containing infective first stage larvae . Larvae hatch in the small

ihtestine and spend eight to ten days in the intestinal muco sa before

returning to the lumen and passing to the cecum. Most infections are

light and cause little , if any, damage although it is believed that

32

!• vulpis is a blood sucker . Affec ted animals wi th heavy in.fections · may

be unthrifty , lose weight , have diarrhea , dysentery , or anemia. 'lli.e

colon and cecum may become in.flamed and even hemorrhagic .

LITERATURE CITED

Ameel , D . J. 1 955 . Parasite s of the coyote , Canis latrans Say , in

Kansas . J. Parasi t . 41 : 325.

Andrews , R . D . , and E . K . Bogge s s . 1 978 . Ecology of c oyotes in Iowa.

Page s 249-265 in M. Bekoff , ed. Coyote s : biology , behavior , and

management . Academic Pre s s , New York.

Bekoff , M. 1 977 . Canis latrans . Amer . Soc . Ma.mm. , Mamm. Spec i e s No . 79 .

Butler , J . M. , and A . w . Grundmann. 1 954. The intestinal helminths of

the coyote Canis latrans Say in Utah . J. Parasit . 40 : 440-443 .

Chandler , A. c . 1 944. A new species of Mesoc e s toide s , !:!• kirbyi , from

Canis latrans . J . Parasit . 30 : 273 .

Conder , G. A . , and R . M. Lovele s s . 1 979 . Parasite s of th e coyote in

central Utah . J. Wldl. Dis . 1 4: 247-249 .

DeGuisti , D. L. 1 971 . Acanthocephala. Pages 1 40-1 57 in J . W . Davi s ,

and R. c . Anderson , eds . Parasitic di seas e s of wild mammals . Iowa

State Univ. Pre ss , Ames , Iowa.

Erickson , A. B . 1 944 . He lminths of Minne sota canidae in re lation to food

habits and a host list and key to the spec i e s reported from North

America. Amer . Midl . Nat . 32 : 358-372 .

Esch , G. w. , and J. T . S e lf . 1 965. A critical study of the taxonomy of

Taenia pisiformis Bloch , 1780 ; Multiceps multiceps ( Le ske , 1 780) ;

and flldatigera taeniaeformi s Batsch , 1 786 . J . Parasit . 51 : 932-937 .

Franson , J . c. , R . D . Jorgenson , E . K. Bogge s s , and J. H . Greve . 1 978 .

33

Gastrointestinal parasitism of Iowa coyote s in relation to age .

J. Parasit . 64 : 303-305 .

Freeman, R . s. , A . Adorjan , and D. H . Pimlott. 1 961 . Cestodes of wolves ,

coyotes , and coyote-dog hybrids in Ontario. C an. J . Zool. 39 :

527-532 .

Gier , H . T. 1 968 . Coyotes in Kansas. rev. ed. Agric. Exp . Sta. Kans .

State Univ. �ic . and Appl . Sci . , Manhattan, Kansas , Bull . 393 .

1 1 8 PP•

Gier , H . T . , and D. J • .Amee l . 1 9 59 . Parasites and di sease s of Kansas

coyotes . Agric . Exp . Sta. Kans . State Univ. Agric . and Appl . Sci . ,

Manhattan , Kansas , Tech. Bull . 91 . 34 PP•

Gier , H . T. , s . M. Kruckenberg, and R. J. Maxler . 1 978. Parasites and

di seases of coyote s . Pages 37-71 in M . Bekoff , ed. Coyote s :

biology, behavior , and management. Academic Press, New York.

Grundmann , A. W . 1 956 . A new tapeworm , Mesocestoides carnivoricolis ,

from carnivores of the Great Salt Lake desert region of Utah. Proc .

Helm. Soc . Wash. 23 : 26-28.

Grundmann, A. W. , 1957. Nematode parasites of the Great Salt Lake desert

of Utah . J. Parasit . 43 : 1 05-1 1 2 .

Grundmann, A. w . 1 958. Cestodes of mammals from the Great Salt Lake

desert region of Utah . J. Parasit . 44 : 425-429 •

Hall , M. c. 1 91 1 . The coyote as a host of Multiceps multiceps. Science

33 : 975.

Hall , M . c . 1 91 2 . A second case of Multiceps multiceps in the coyote.

Science 35 : 556.

Hall , M . c . 1 91 4. A new nematode , Rictularia splendida, from the coyote ,

34

with notes on other coyote parasites . Proc . U. S . Natl . Mus . 46 :

73-84.

Hall , M. c. 1 929 . 'lbe adult taenioid cestodes of dogs and cats , and of

related carnivores in North America. Proc . u . s. Natl . Mus . 55 :

1 -94.

Hirsch , R . P. , and H . T . Gier . 1 974. Multiple-species infections of

inte stinal helminths in Kansas coyotes . J . Parasit. 60 : 650-653 .

Holmes , J . c. , and R. Podesta. 1 968. The helminths of wolve s and coyote s

from the forested regions of Alberta. Can. J. Zool . 46 : 1 1 93-1 204.

Leiby, P. D. , w. P. Carney , and c. E . Woods . 1 970 . Studies on sylvatic

echinococcosi s . III . Hos t occurrence and geographic di stributi on of

Echinococcus mu.ltiloculari s in the north c entral Uni ted State s . J.

Parasit. 56 : 1 1 41 -1 1 51 .

Leiby , P. D. , and w . G. Dyer . 1 971 . Cyc lophyllidean tapeworms of wild

carnivora. Pages 1 74-234 in J. w.. Davis , and R. C . Anderson , eds .

Parasitic disease s of wild mammals . Iowa S tate Univ. Pres s , Ame s ,

Iowa.

Levine , N. D. 1 980. Nematode parasites of dome stic animals and man, 2nd

ed. Burge s s Publ. C o . , Minneapolis , Minnesota. 477 PP •

Liu , I . K. M. , c . w . Schwabe , P . M . Schantz , and M . N. Allison. 1 970.

The occurrence of Echinococcus granulosus in coyote s in the central

valley of California. J. Parasit. 56 : 1135-1138.

Mitchell , R . L . , .and s . L . Beasom.

and bobcats . J. Wldl . Mgmt .

Morgan, B . B . , and P. A. Hawkins.

1 974. Hookworms in south Texas coyote s

38 : 455-458 .

1 951 . Veterinary helminthology.

Burgess Publ . Co. , Minneapolis , Minnesota. 400 PP•

35

Olsen , o . w . 1 960. Sylvatic trichino sis in carnivorous mammals in the

Rocky Mountain region of Colorado . J. Pa.rasit . 46 : 22 .

Olsen , o . w. 1 974. Animal parasite s : their life cycles and ecology ,

3rd ed. Univ. Park .Pre s s , Baltimore , Maryland . 562 pp .

Pence ' D. B . , and w . P . Meinz er . · 1 979 . Helminth parasitism in the coyo te ,

Cani s latrans , from the Rolling Plains of Texas . Int . J. Parasit .

9 : 339-344.

Price , E. w. 1 928. 'lhe c oyote (Canis latra.n:s texensis ) , a new host for

Oncicola canis (Kaupp) and Oslerus o sleri (C obbold) . J. Pa.rasi t .

1 4 : 1 97 .

Rausch , R . , B . B . Babero , R . v . Rausch , and E . L . Schiller. 1 956 .

Studies on the helminth fauna of Alaska. XXVII. The occurrenc e of

larvae of Trichinella spirali s in Alaskan mammals . J . Parasi t .

42 : 259-271 .

Samuel , w . M. , s . Ramalingam, and L. N . Carbyn. 1 978. Helminths in

coyotes , wolve s , and red foxe s of southwe stern Manitoba. C an . J.

Z.ool. 56 : 261 4-261 7 .

Schmitt , N . e t al . 1 976 . Can. J. Pub . Health . 67 ( 1 ) : 21 -24.

Self, J. T . , and T . J. McKnight. 1 950. Platyhelminthes from fur bearers

in the Wichita Mountains Wildlife Refuge , with e special reference to

Oochori s tica spp . Amer . M.idl . Nat . 43 : 58-61 .

Thornton , J . E . , R . R . Bell , and M. J . Reardon. 1 9 74. Internal parasites

of coyote s in southern Texas . J. Wldl . Di s . 1 0 : 232-236.

Van C leave , H. J. 1 953 . Acanthocephala. of North American mammals .

Ill . Biol . Monogr . , Univ. Illinoi s Pre s s , Urbana. Vol . 23. No . 1 -2 .

1 79 PP •

36

Voge , M. 1 955 . A li st of c e s tode parasi t e s from California mammal s •

Amer o Midl . Nat . 54 : 41 3-41 7 .

Wardle , R . A . , J . A . McLeod , and s . Radinovsky. 1 974. Advance s in the

z oo logy of tapeworms , 1 950-1 970. Univ. Minn . Pre s s , Minneapolis,

Minne sota . 274 PP •

Wood , I. B . , and J. T . Haldiman. 1 957 . New locality and host records for

the cestode genus Me soc e s toide s . J . Parasi t . 43 : 598.

Yamaguti , s . 1 959 . Sys terna helminthurn. Vol . II . Th e c e s tode s of

vertebrate s . Interscience Publ . , Inc . , New York . 860 pp .

Yamaguti , s. 1 961 . Systerna helminthurn . Vol . III . The nematode s of

vertebrate s . Part 1 . Interscienc e Publ . , Inc . , New Yorko 679 PP •

Yamaguti , s . 1 963 . Systema helminthurn. Vol . V . Acanthocephala.

Interscience Pu.bl . , Inc . , New York . 423 pp .

Zimmerman , w . J. 1 971 . Trichino si s . Page s 1 27-1 39 !!!. J. W . Davi s , and

R . c . Anderson , eds . Parasitic di seas e s of wild mammals . Iowa

State Univ. Press , Ame s , Iowa .

Zimmerman , w. J . , and E. D. Hubbard. 1 969 . Trichiniasi s in wildlife

of Iowa. Amer . J. Epidemiol . 90 : 84-92 .

\