11

NOAA Technical Report NMFS 11

Taxonomy of North American

Fish Eimeriidae

Steve J. Upton, David W. Reduker, William L. Current,

and Donald W. Duszynski

August 1984

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service
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St., Washington, DC 20235.
NOAA Technical Report NMFS 11

Taxonomy of North American

Fish Eimeriidae

Steve J. Upton, David W. Reduker, William L. Current,

and Donald W. Duszynski

August 1984

u.s. DEPARTMENT OF COMMERCE

Malcolm Baldrige, Secretary

National Oceanic and Atmospheric Administration

John V. Byrne, Administrator

National Marine Fisheries Service

William G. Gordon, Assistant Administrator for Fisheries
The National Marine Fisheries Service (NMFS) does not approve, recommend or endorse any proprietary product or proprietary material

mentioned in this publication. No reference shall be made to NMFS, or

to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or imply that NMFS approves, recommends

or endorses any proprietary product or proprietary material mentioned

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publication.
CONTENTS

Introduction ... '. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Taxonomic description. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Eimeria anguillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Eimeria aurati . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Eimeria brevoortiana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Eimeria catostomi :............... 3

Eimeria duszynskii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Eimeria etheostomae 3

Eimeriafernandoae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Eimeriafreemani . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Eimeriafunduli .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Eimeria gasterostei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Eimeria glenorensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Eimeria haneki . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

Eimeria hoffmani . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Eimeria hybognathi .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Eimeria ictaluri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Eimeria iroquoina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Eimeria laureleus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Eimeria micropteri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Eimeria moronei 6

Eimeria myoxocephali . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Eimeria ojibwana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Eimeria osmeri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Eimeria pungitii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Eimeria salvelini 7

Eimeria squali . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Eimeria tedlai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Eimeria truttae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Eimeria sp. Davis, 1946 8

Eimeria sp. Fantham and Porter, 1948 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Eimeria sp. Fantham and Porter, 1948 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Eimeria sp. Munson, 1974 8

Eimeria sp. Orias and Noble, 1971 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Goussia caseosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Goussia degiustii 9

Goussia gadi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Key to Eimeriidae of North American fishes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Figures 1-30 13

iii

Taxonomy of North American Fish Eimeriidae

STEVE J. UPTON,1 DAVID W. REDUKER,2 WILLIAM L. CURRENT,3

and DONALD E. DUSZYNSKP

ABSTRACT

Taxonomic descriptions, line drawings, and references are given for the 30 named and S unnamed species of

North American fish Eimeriidae. In addition, a key was developed based on available morphologic data to

distinguish between similar species. Taxa are divided into two genera: Eimeria (27 species) which are tetr&sporocystic with dizoic, nonbivalved sporocysts, and Goussia (3 species) which are tetrasporocystic with dizoic, bivalved sporocysts that lack Stleda bodies and have sporocyst walls composed of two longitudinal valves.

INTRODUCTION

The phylum Apicomplexa Levine, 1970, comprises about

4,000 species of totally parasitic protozoa which include the

gregarines, haemogregarines, coccidia, malaria, and piroplasms

(see Levine 1982 for taxonomic review). The largest family in the

phylum is the Eimeriidae Minchin, 1903, which contains nearly

1,400 named species; over 75% of which belong to a single genus

Eimeria.

Traditionally, genera within the Eimeriidae have been

classified according to the number of sporocysts within the oocyst

and the number of sporozoites within the sporocysts. The Eimeria

are characterized by an oocyst which contains four sporocysts,

each with two sporozoites.

Most Eimeriidae are species (or at least genus) specific and

have similar life cycles. After an appropriate host ingests a

sporulated oocyst, sporozoites are released into the intestinal

lumen by the action of digestive enzymes and bile salts upon the

oocyst and sporocysts. Sporozoites of most species enter intestinal

epithelial cells where they reproduce asexually by merogony

(schizogony), forming numerous merozoites by multiple fission.

Usually merozoites enter new host cells and initiate one or more

merogonous cycles. The number of asexual generations is believed

to be genetically determined and the last generation merozoites

invade new host cells to form gamonts. Most gamonts enlarge

without karyokinesis into macrogametes, while others undergo

multiple fission giving rise to numerous flagellated microgametes.

When released, microgametes apparently fertilize macrogametes

and the resulting zygote forms an oocyst wall to protect it from

adverse environmental conditions. Usually, oocysts are passed out

of the host unsporulated. Under the appropriate conditions of oxygen, temperature, and moisture, sporulation will occur and the

sporulated oocyst is then infective for the next host via fecal-oral

contamination. Thus, most Eim.eriidae develop directly (utilize

one host) and produce self-limited infections.

About 130 named species and over two dozen unnamed species

of fish coccidia have been reported. All of these species except

three, Cryptosporidium nasoris Hoover, Hoerr, Carlton, Hinsman,

and Ferguson, 1981, from a naso tang; Sarcocystis salvelini FanIDepartment of Zoology-Entomology. Auburn University, Ala.; present address:

Biology Department, University of Texas, EI Paso, TX 79968.

2Department of Biology, University of New Mexico, Albuquerque, NM 87131.

'Department of Zoology-Entomology, Auburn University, Ala.; present address:

Animal Health Discovery, Eli Lilly Research Laboratories, P.O. Box 708, Greenfield,

iN 46140.

tham and Porter, 1943 from a trout; and Sarcocystis sp. Fantham

and Porter, 1943 from an eel pout, have been placed within the

Eimeria (Fantham and Porter 1943; Hoover et al. 1981).

Although little is known about the life cycles of fish coccidia,

most developmental stages appear morphologically similar to the

more well known species infecting higher vertebrates. However,

'two important differences can be noted for the coccidia of fish.

First, about one-third of the species of fish coccidia develop and

sporulate in nonintestinal sites and apparently remain in situ until

the host dies. Second, whether oocysts remain in host tissues or

exit via the feces, fish coccidia usually sporulate endogenously

and, presumably, are immediately infective when they contact the

external environment. Although the significance of these

peculiarities is not known, readers are directed to a recent review

by Dykovil and Lorn (1981) for an excellent overview of the

biology of fish coccidia.

Virtually nothing is known about the incidence of fish coccidia

in North America and little is known about the impact that these

coccidia have on their fish hosts. Solangi and Ogle (1981)

reported lower weight gains for killifish infected with the hepatic

coccidium Eimeria funduli than for uninfected, control fish.

Odense and Logan (1976) believed heavy swim bladder infections

of Goussia gadi might interfere with the ability of haddock to

spawn. In other countries, however, some fish coccidia have been

shown to be econom.ically important pathogens. Eimeria sinensis,

E. subepithelialis, and Goussia carpelli may cause enteritis and

death of commercially reared carp (Molnar 1979; Dykovil and

Lorn 1981). New Zealand eels infected with E. anguillae may

become emaciated and weak (Hine 1975). Sterility can result in

male clupeids infected with the testicular coccidium E. sardinae

(Pinto 1956). MacKenzie (1981) has reported that an unnamed

species of hepatic coccidium causes blue whiting to weigh less

than uninfected fish of similar age.

Recently, Dykovil and Lorn (1981) have split the Eimeria of

fish into four genera based on available morphologic and life

cycle data. These include: 1) Eimeria, which retains most of the

species and is characterized by being tetrasporocystic with dizoic,

nonbivalved sporocysts with or without Stieda bodies; 2) Goussia

Labbe, 1896 containing about 20 species which are tetrasporocystic, dizoic, lack Stieda bodies, and have sporocyst walls consisting of two valves joined by a longitudinal suture (Dykovil and

Lorn 1981; Lorn and Dykovil 1982a, b); 3) Crystallospora Labbe,

1896 contains a single species, Crystallospora cristalloides

(Thelohan, 1893) Dykovil and Lorn, 1981, which is tetrasporocystic, dizoic, and has dodecahedral sporocysts composed of two
hexagonal, pyrimidal valves joined at their bases by a suture; and

4) Epieimeria Dykova and Lorn, 1981 which was created for those

tetrasporocystic, dizoic Eimeriidae of fish that possess Stieda

bodies and undergo merogony and gametogony on the lumenal

surface of the intestinal tract; three species are included in this

genus (Dykova and Lorn 1981; Lorn and Dykova 1982a).

Innate problems exist with portions of this classification

scheme; particularly for those unfamiliar with the pecularities of

fish coccidia. Crystallospora has been reported only once, and only

briefly, since the original description in 1893 (Dogiel 1948). We

believe a more accurate and up to date description is needed to retain this genus. Some species (e.g., Eimeria anguillae Leger and

Hollande, 1922 and Eimeria hexagona Lorn and Dykova, 1981)

also appear hexagonal in cross section and, thus, the only difference between the sporocysts of Crystallospora and those of

several other species is the location of the suture. A more detailed

examination of the sporocyst, particularly in the placement of the

suture, is needed before the use of the genus Crystallospora can be

warranted.

The Epieimeria, which are probably not epicellular but rather

develop in a parasitophorous vacuole on the lumenal side of the

endothelium, require histological and cytological examination to

differentiate them from the Eimeria. Additionally, Eimeria pigra

Leger and Bory, 1932, a coccidium infecting the intestinal

epithelium of the rudd, Scardinius erythrophthalmus, not only

develops "epicellularly," but occasionally intracellularly and

lacks a Stieda body, unlike other Epieimeria. This genus also requires additional research to warrant proper recognition.

Pellerdy (1974) provided descriptions of most of the known

fish coccidia through 1970. Since then, many new species have

been reported. Because of the numerous species of fish coccidia,

and because many of the original descriptions are in languages

other than English, a detailed summary of each species is beyond

the scope of this paper. Instead, our purpose is to provide succinct

taxonomic descriptions of the known Eimeriidae from North

American fish and to provide line drawings and original

references that would be beneficial. These are presented below,

along with literature known to us on North American fish coccidia. Taxa are divided into two genera, Eimeria and GOUSSla, and

a taxonomic scheme is presented that we believe provides for

rapid and accurate means of diagnosing fish coccidia.

TAXONOMIC DESCRIPTION

Eimeriidae Minchin, 1903

Genus Eimeria Schneider, 1875

Oocysts contain four sporocysts, each with two sporozoites;

Stieda body mayor may not be present; sporocyst wall nonbivalved.

Eimeria anguillae Leger and Hollande, 1922

(Fig. 1)

Synonyms: Eimeria sp. Hine and Boustead, 1974; Epieimeria

anguillae (Leger and Hollande, 1922) Dykova and Lorn, 1981.

Hosts: Anguilla anguilla "European eel," A. australis "shortfinned eel," A. dieffenbachii "Iongfinned eel," A. rostrata

"American eel" (Anguilliformes) (C3).

'All measurements are in micrometers ().<m). Abbreviations: (B) brackish water;

(C) catadromous; (F) freshwater; (M) marine.

2

Description: Oocysts spherical, 9.8 (9.1-10.4) (see footnote 3).

Wall smooth and colorless, composed of a single layer'" 0.1.

Micropyle, polar granule, and oocyst residuum absent. Sporocysts

ovoid, hexagonal in cross section, 7.8 by 4.1 (7.5-8.0 by 3.9-4.4),

in contact with oocyst wall. They are usually arranged so that

parallel pairs of sporocysts cross each other. Wall thin, '" 0.2;

Stieda body present, formed by 4 papillae situated at tapered end

of sporocyst. Sporocyst residuum present, spherical or ellipsoid,

compact, 2.3 by 1.6 (1.9-2.6 by 1.3-1.9). Sporozoites vermiform,

reflexed at one end, 6.8 by 1.5 (6.5-7.0 by 1.3-1.8) (without

reflexed portion), arranged head to tail in sporocyst. Refractile

bodies absent.

Sporulation: Endogenous.

Geographical locations: Corsica; New Zealand; Quebec, Canada.

Location in host: Anterior intestinal epithelium; feces.

References: Leger and Hollande (1922); Schulman and Shtein

(1962); Molnar and Hanek (1974); Hine and Boustead (1974);

Hine (1975); Dykova and Lorn (1981).

Remarks: Merogony and gamogony occur on the lumenal border

of the endothelium in the anterior portion of the intestine. The

description is based predominately on that given by Molnar and

Hanek (1974).

Eimeria aurati HotTman, 1965

(Fig. 2)

Host: Carassius auratus "goldfish" (Cypriniformes) (F).

Description: Oocysts ovoid, occasionally spherical, 20.1 by 16.3

(16.0-24.0 by 14.0-17.0). Wall thin, '" 0.3. Micropyle, polar

granule, and oocyst residuum absent. Sporocysts ellipsoid,

11.0-13.0 by 6.5-8.0. Stieda body and sporocyst residuum absent.

Sporozoites sausage-shaped, 10.0-13.0 by 2.0-2.5, lying lengthwise and nearly filling the sporocyst entirely. Refractile bodies absent. Older oocysts may contain free sporozoites.

Sporulation: Exogenous.

Geographical location: Pennsylvania, U.S.A.

Location in host: Anterior intestinal epithelium; feces.

Reference: Hoffman (1965).

Remarks: Unsporulated oocysts were numerous in long, whitish,

opaque fecal casts.

Eimeria brevoortiana Hardcastle, 1944

(Fig. 3)

Host: Brevoortia tyrannus "Atlantic menhaden" (Clupeiformes)

(M).

Description: Oocysts spherical, 25.1 (17.5-30.0), or occasionally ovoid, 26.2 by 22.7 (21.2-30.0 by 15.0-27.5). Micropyle absent; oocyst residuum described but not depicted. A polar granule

is not described or depicted. Sporocysts elongate, 16.4 by 6.3.
Sporocyst residuum small; a Stieda body is not described.

Sporozoites cigar-shaped, slightly curved, 15.0 by 2.5 and arranged head to tail within the sporocyst. Each sporozoite contains a

large, centrally located nucleus and (possibly) a small, anterior,

refractile body.

Sporulation: Endogenous.

Geographical location: Beaufort, North Carolina, U.S.A.

Location in host: Testes.

Reference: Hardcastle (1944).

Remarks: Hardcastle (1944) found that merogony and

gamogony occurred in the endothelium of the pyloric caeca of

both male and female fish, however, oocysts were only found in

the testes of males. He postulated that after fertilization, a reduction division occurred involving the entire zygote resulting in two

migratory ookinetes which carried the infection to the testes. Two

separate species may be involved.

Eimeria catostomi Molnar and Hanek, 1974

(Fig. 4a, b)

Hosts: Catostomus commersoni "white sucker," Hypentelium

nigricans "northern hog sucker" (Cypriniformes) (F).

Description: Oocysts spherical, 7.0 (6.5-7.5). Wall smooth,

colorless, composed of a single layer::: 0.1. Micropyle and oocyst

residuum absent. Polar granule present, centrally located, 0.8

(0.7-0.9). Sporocysts ovoid, flattened on one side, 5.5 by 3.8

(5.2-5.9 by 3.6-4.0), in contact with oocyst wall. Sporocysts

arranged so that three lie in one plane with fourth overlying. Wall

single layered, ::: 0.2; Stieda body absent. Sporocyst residuum

spherical, finely granular, 2.3 (2.0-2.6). Sporozoites vermiform,

slightly curved with one end reflected, 4.6 by 1.4 (4.4-4.8 by

1.3-1.6) and lie head to tail within the sporocyst. Refractile bodies

absent.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Anterior intestinal epithelium; feces.

Reference: Molnar and Hanek (1974).

Eimeria duszynskii Conder, Oberndorfer,

and Heckmann, 1980

(Fig. 5)

Host: Cottus bairdi "mottled sculpin" (Perciformes) (F).

Description: Oocysts irregular in shape, 12.2 (11.6-12.9). Wall

very thin, adhering closely to sporocysts. Micropyle, polar

granule, and oocyst residuum absent. Sporocysts ovoid, 9.1 by 6.0

(8.4-10.0 by 5.2-6.3), with one side slightly flattened. Stieda and

substieda bodies absent. Sporocyst residuum present, compact and

coarsely granular, occasionally dispersed. Sporozoites 2.1 wide,

lying head to tail in sporocyst with one end reflexed. Each sporozoite contains a large, oblong, refractile body near one end. A

3

spherical nucleus and transverse banding occur at the end opposite

from the refractile body.

Sporulation: Endogenous.

Geographical location: Provo river, Utah, U.S.A.

Location in host: Intestinal epithelium; feces.

Reference: Conder et al. (1980).

Eimeria etheostomae Molnar and Hanek, 1974

(Fig. 6)

Hosts: Etheostoma exile "Iowa darter," E. nigrum "Johnny

darter" (Perciformes) (F).

Description: Oocysts spherical, 9.4 (9.1-10.4). Wall smooth and

colorless, composed of a single layer::: 0.1. Micropyle, polar

granule, and oocyst residuum absent. Sporocysts coffee-bean

shaped, 8.5 by 5.0 (7.8-9.1 by 4.5-5.4), in contact with oocyst

wall. Sporocysts usually arranged so that three lie in one plane

with fourth overlying. Stieda body absent. Sporocyst residuum

present, spherical, and finely granular, 3.6 (3.1-3.9). Sporozoites

banana-shaped, 8.4 by 1.6 (7.8-9.0 by 1.4-1.8), arranged head to

tail in sporocyst. Rarely, one end of the sporozoite may be reflexed. Each sporozoite has a spherical refractile body.

Sporulation: About one-half of the oocysts sporulate endogenously; the rest exit semisporulated.

Geographical location: Ontario, Canada.

Location in host: Anterior intestinal epithelium; feces.

Reference: Molnar and Hanek (1974).

Eimeria fernandoae Molnar and Hanek, 1974

(Fig. 7)

Hosts: Catostomus commersoni "white sucker," Hypentelium

nigricans "northern hog sucker" (Cypriniforrnes) (F).

Description: Oocysts ellipsoid, 8.3 by 6.6 (7.8-9.0 by 6.5-7.0).

Wall smooth and colorless, composed of a single layer::: 0.1.

Micropyle, polar granule, and oocyst residuum absent. Sporocysts

ellipsoid, 7.2 by 3.0 (6.8-7.5 by 2.6-3.4), in contact with oocyst

wall. All four sporocysts lie parallel in oocyst. Wall 0.2; Stieda

body absent. Sporocyst residuum spherical and finely granular,

2.6 (2.3-3.2). Sporozoites banana-shaped, 5.8 by 1.2 (5.2-6.5 by

1.0-1.3), arranged head to tail in sporocyst. Each sporozoite has a

spherical refractile body.

Sporulation: Endogenous.

Geographical location: Ontario, Canada

Location in host: Anterior intestinal epithelium; feces.

Reference: Molnar and Hanek (1974).
Eimeria freemani Molnar and Fernando, 1974

(Fig. 8)

Host: Notropis cornutus "common shiner" (Cypriniformes) (F).

Description: Oocysts ellipsoid, 22.0-24.0 by 17.0-18.0. Wall

smooth and colorless, composed of a single layer::: 0.1. Oocyst

residuum absent, a micropyle is not described. Polar granule present, 1.0-2.0. Sporocysts ellipsoid, 16.0-17.0 by 5.0-6.5, arranged

parallel within oocyst. Wall::: 0.2; a Stieda body is not described.

Sporocyst residuum spherical, compact, and finely granular,

2.5-5.0. Sporozoites banana-shaped, 14.0-15.5 by 2.5-3.5. Each

sporozoite has a large, posterior, refractile body and a smaller

refractile body more centrally located.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Kidney, renal parenchyma, renal tubules.

Reference: Molnar and Fernando (1974).

Eimeriafunduli Duszynski, Solangi,

and Overstreet, 1979

(Fig. 9)

Hosts: Fundulus grandis "gulf killifish," F. heteroclitus

"mummichog," F. pulvereus "bayou killifish," F. simi/is

"longnose killifish," Menidia beryllina "inland silverside"

(Atheriniformes) (B).

Description: Oocysts spherical, 24.5 (20.0-31.0). Wall smooth

and colorless, ::: 1.0, composed of two layers of equal thickness.

Outer layer transparent; inner layer opaque. Micropyle, polar

granule, and oocyst residuum absent. Sporocysts ovoid, 10.0 by

6.0 (9.0-11.0 by 5.0-7.0). Shape index 1.68 (1.45-2.02). Stieda

and substieda bodies present. Thin membrane encloses sporocysts

completely forming a transparent matrix, 1.7 (1.0-3.0) wide.

Membrane supported by 15 (10-25) footlike structures termed

sporopodia attached to outer surface of sporocysts. Sporocyst

residuum present, consisting of 1 to 4 refractile granules between

the sporozoites. Each sporozoite contains a single refractile body

near the posterior end.

Sporulation: Endogenous.

Geographical locations: Estuaries of Mississippi, Alabama, and

Louisiana, U.S.A.

Location in host: Liver (rarely intestinal mucosa and submucosa, ovary, testes, caudal peduncle).

References: Duszynski et al. (1979); life-cycle, Solangi and

Overstreet (1980), Upton and Duszynski (1982), Fournie and

Overstreet (1983); Hawkins, Fournie, and Overstreet (1983b); effect of temperature, Solangi et al. (1982); prevalence, Fournie and

Solangi (1980); effects on fish, Solangi and Ogle (1981), Hawkins

et al. (1981); transmission studies, Fournie and Overstreet (1982);

ultrastructure of sexual stages, Hawkins, Solangi, and Overstreet

4

(1983a, b); ultrastructure of oocysts, Hawkins, Fournie, and

Overstreet (l983a).

Eimeria gasterostei (Thelohan, 1890) Dotlein, 1909

(Fig. 10)

Synonym: Coccidium gasterostei Thelohan, 1890

Hosts: Gasterosteus aculeatus "threespine stickleback" (Gasterosteiformes) (F,M).

Description: Oocysts spherical, 16.0-18.0, with smooth thin

wall. Oocyst wall greatly separated from sporocysts. Oocyst

residuum absent; micropyle and polar granule not described.

Sporocysts elongate, narowing at poles, 10.0-14.3 by 4.0-6.5.

Sporocyst residuum large and coarsely granular. Sporozoites

botuliform, each with a large refractile body.

Sporulation: Endogenous.

Geographical locations: Vancouver, British Columbia; Kamchatka, Paratunka river basin.

Location in host: Liver.

References: Thelohan (1890); Schulman and Shtein (1962);

Lester (1974).

Remarks: Description is based on that given by Thelohan

(1890), Schulman and Shtein (1962), and Lester (1974).

Eimeria glenorensis Molnar and Fernando, 1974

(Fig. 11)

Host: Morone americana "white perch" (Perciformes) (F).

Description: Oocysts spherical, 10.5-12.0. Wall smooth and

colorless, composed of a single layer::: 0.1. Oocyst residuum absent; a micropyle is not described. One or two polar granules present, 1.0-2.0. Sporocysts ovoid, 8.0-9.5 by 5.7-6.0, in contact with

oocyst wall. Stieda body present, consisting of a thickening of the

sporocyst wall at tapered end, ::: 0.5 high; rest of wall ::: 0.3.

Sporocyst residuum spherical and compact, refractile, 2.0-2.6.

Sporozoites vermiform, one end reflexed, 5.0-5.3 by 2.0-2.2

(without reflexed portion). Reflexed portion 2.5-2.7 long.

Sporozoites arranged head to tail in sporocyst. Refractile bodies

absent.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando (1974).

Eimeria haneki Molnar and Fernando, 1974

(Fig. 12)

Host: Culaea inconstans "brook stickleback" (Gasterosteiformes) (F).
Description: Oocysts spherical, 7.1-7.8. Wall smooth and colorless, composed of a single layer:::: 0.1. Oocyst residuum absent; a

micropyle is not described. One or two polar granules present,

1.0-1.5. Sporocysts ellipsoid, 5.6-6.5 by 3.9-4.5, compact. and in

contact with oocyst wall. Sporocysts usually arranged so that pairs

of parallel sporocysts cross each other. Wall:::: 0.2. A Stieda body

is not described. Sporocyst residuum spherical and compact,

nongranular, refractile, 1.3-1.5. Sporozoites vermiform, one end

reflexed, 5.6-6.0 by 1.3-2.0 (without reflexed portion), arranged

head to tail in sporocyst. Each sporozoite has a spherical refnctile

body.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando (1974).

Eimeria hoffmani Molnar and Hanek, 1974

(Fig. 13)

Host: Umbra limi "central mudminnow" (Salmoniformes) (F).

Description: Oocysts ellipsoid, 11.5 by 9.2 (11.0-12.2 by

9.1-9.6). Wall smooth and colorless, composed of a single layer ::::

0.1. Micropyle and oocyst residuum absent. Polar granule present,

0.8 (0.7-0.9). Sporocysts ellipsoid, 10.C by 3.6 (9.6-10.4 by

3.4-3.9), arranged lengthwise and paralle!. in oocyst. Wall :::: 0.2;

Stieda body absent. Sporocyst residuum ellipsoid and finely

granular, 6.4 by 2.0 (6.2-6.6 by 1.8-2.2). In older oocysts the

residuum is either less elongate or dispersed. Sporozoites vermiform, one end reflexed, 8.6 by 1.4 (8.4-8.7 by 1.3-1.5), (without

reflexed portion), arranged head to tail in sporocyst. Refractile

bodies absent.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Anterior intestinal epithelium; feces.

Reference: Molnar and Hanek (1974).

Eimeria hybognathi Molnar and Fernando, 1974

(Fig. 14)

Host: Hybognathus hankinsoni "brassy minnow" (Cypriniformes) (F).

Description: Oocysts spherical, 14.0-14.5. Wall of uniform

thickness, composed of a single layer:::: 0.1. Polar granule and

oocyst residuum absent. A micropyle is not described. 3porocysts

ellipsoid, 9.5-10.0 by 5.7-6.0, with a narrow space separating

them from oocyst wall. They are usually arranged so that pairs of

parallel sporocysts cross each other. Sometimes three lie in one

plane with the fourth overlying. Wall:::: 0.2. A Stieda body is not

described. Sporocyst residuum ovoid, compact, and finely

granular, 5.8-8.0 by 4.5-5.2. Sporozoites banana-shaped, one end

slightly reflected, 9.0-9.2 by 2.0-2.6, lying head to tail in sporo5

cyst. A single spherical refractile body is present in each

sporozoite.

Sporulation: Endogenolls.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando (1974).

Eimeria ictaluri Molnar and Fernando, 1974

(Fig. 15)

Host: Ictalurus nebulosus "brown bullhead" (Siluriformes) (F).

Description: Oocysts spherical, 9.0-10.5. Wall colorless and of

uniform thickness, composed of a single layer:::: 0.1. Polar granule

and oocyst residuum absent; a micropyle is not described.

Sporocysts ellipsoid, rarely coffee-bean shaped, 7.8-8.4 by

3.6-4.2, in contact with oocyst wall. Sporocysts usually arranged

so that pairs of parallel sporocysts cross eaclJ other. Wall 0.2; a

Stieda body is not described. Sporocyst residuull! ovoid, compact,

and coarsely granular, 5.0-5.3 by 3.0-3.3. Sporozoites vermiform,

one end reflexed, 7.1-7.8 by 1.5-1.8, without reflexed portion, arranged head to tail in sporocyst. Each sporozoite has a spherical

refractile body.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando (1974).

Eimeria iroquoina Molnar and Fernando, 1974

(Fig. 16)

Hosts: Nocomis biguttatus "hornyhead chub," Notropis comutus

"common shiner," N. heterolepis "blacknose shiner," N. rubel/us

"rosyface shiner," Pimephales notatus "bluntnose minnow," P.

promelas "fathead minnow," Rhinichthys atratulus "blacknose

dace," Semotilus atromaculatus "creek chub" (Cypriniformes) (F).

Description: Oocysts spherical, 9.0-11.0. Wall colorless and of

uniform thickness, composed of a single layer:::: 0.1. Polar granule

and oocyst residuum absent; a micropyle is not described. Sporocysts ellipsoid, 7.8-8.0 by 4.0-4.5, in contact with oocyst wall.

Sporocysts usually arranged so that pairs of parallel sporocysts

cross each other. Wall :::: 0.2; a Stieda body is not described.

Sporocyst residuum ovoid, finely granular, 2.3-4.0 by 2.2-3.0.

Sporozoites vermiform, one end reflexed, 7.5-7.7 by 1.5-1.7

(without reflexed portion), arranged head to tail in sporocyst. Each

sporozoite has a spherical refractile body.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.
References: Molnar and Fernando (1974); Molnar and Hanek

(1974); Paterson and Desser (l981a, b, c, 1982).

Eimeria laureleus Molnar and Fernando, 1974

(Fig. 17)

Host: Perca flavescens "yellow perch" (Perciformes) (F).

Description: Oocysts spherical, 11.0-12.0. Wall colorless and of

uniform thickness, composed of a single layer::: 0.1. Polar granule

and oocyst residuum absent; a micropyle is not described. Sporocysts coffee-bean shaped, 9.2-11.0 by 5.0-5.8, in contact with

oocyst wall. Sporocysts usually arranged so that pairs of parallel

sporocysts cross each other. A Stieda body is not described.

Sporocyst residuum finely granular and dispersed. Spvrozoites

vermiform, one end reflexed, 9.0-10.0 by 1.5-2.0, without reflexed portion, arranged head to tail in sporocyst. Each sporozoite has

a spherical refractile body.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando (1974).

Eimeria micropteri Molnar and Hanek, 1974

(Fig. 18)

Host: Micropterus dolomieui "smallmouth bass," M. salmoides

"largemouth bass" (Perciformes) (F).

Description: Oocysts spherical, 12.0 (11.7-12.5). Wall smooth

and colorless, composed of a single layer::: 0.1. Micropyle, polar

granule, and oocyst residuum absent. Sporocysts ellipsoid, 11.4 by

6.2 (11.0-11.7 by 6.0-6.5), in contact with oocyst wall. Sporocysts arranged three in one plane with fourth overlying. Wall :::

0.2; Stieda body absent. Sporocyst residuum finely granular,

dispersed. Sporozoites vermiform, one end reflexed, 9.1 by 2.1

(8.9-9.3 by 2.0-2.2), (without reflexed portion), arranged head to

tail in sporocyst. Each sporozoite has a spherical refractile body.

Sporulation: About 90% leave the gut semisporulated; the rest

exit fully sporulated.

Geographical location: Ontario, Canada.

Location in host: Anterior intestinal epithelium; feces.

Reference: Molnar and Hanek (1974).

Eimeria moronei Molnar and Fernando, 1974

(Fig. 19)

Host: Morone americana "white perch" (Perciformes) (F).

Description: Oocysts spherical, 7.2-8.0. Wall colorless and of

uniform thickness, composed of a single layer ::: 0.1. Oocyst

residuum absent; a micropyle is not described. Polar granule present, 1.0-2.0. Sporocysts ovoid, 5.7-6.0 by 3.9-4.0, in contact with

6

oocyst wall. Sporocysts arranged three in one plane with fourth

overlying. Wall::: 0.2; Stieda body present, consisting of a distinct

knoblike thickening at tapered end. Sporocyst residuum spherical,

compact, refractile, 1.5-2.0. Sporozoites vermiform, 5.0-5.4 by

1.5-2.0, arranged head to tail in sporocyst. Refractile bodies

absent.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Locatioll in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando (1974).

Eimeria myoxocephali Fitzgerald, 1975

(Fig. 20)

Host: Myoxocephalus polyacanthocephalus "great sculpin" (Perciformes) (M).

Description: Oocysts spherical, 37.2 (34.0-40.0). Wall bilayered

and smooth, ::: 2.0. Micropyle and oocyst residuum absent; a polar

granule is not described or depicted. Sporocysts composed of a

thin membrane stretched tightly around the sporozoites. Sporocyst

residuum present, small, and spherical. Sporozoites elongate, 16.7

by 3.7, twisted around each other and arranged head to tail in

sporocyst membrane. A single refractile body is present in each

sporozoite.

Sporulation: Endogenous.

Geographical location: Puget Sound, Washington, U.S.A.

Location in host: Anterior intestinal mucosa; feces.

Reference: Fitzgerald (1975).

Eimeria ojibwana Molnar and Fernando, 1974

(Fig. 21)

Host: Cottus bairdi "mottled sculpin" (Perciformes) (F).

Description: Oocysts spherical, 10.4-11.0. Wall colorless and of

uniform thickness, composed of a single layer::: 0.1. Polar granule

and oocyst residuum absent; a micropyle is not described. Sporocysts coffee-bean shaped, 9.0-9.2 by 5.0-5.8, in contact with

oocyst wall. Sporocysts usually arranged so that pairs of parallel

sporocysts cross each other. Wall ::: 0.2; a Stieda body is not

described. Sporocyst residuum composed of irregularly sized

granules, dispersed. Sporozoites vermiform, reflexed on one end,

8.4-8.7 by 2.0-2.5 (without reflexed portion), arranged head to

tail in sporocyst. Each sporozoite has a spherical refractile body.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando (1974).
Eimeria osmeri Molnar and Fernando, 1974

(Fig. 22)

Host: Osmerus mordax "rainbow smelt" (Salmoniformes) (F).

Description: Oocysts spherical, 10.4-11.0. Wall colorless and of

uniform thickness, composed of a single layer :::: 0.1. Oocyst

residuum absent; a micropyle is not described. One or two polar

granules present, 1.5-1.7. Sporocysts ovoid, 7.1-7.8 by 4.3-4.7,

with a narrow space separating them from oocyst wall. Stieda

body present, consisting of thickening of sporocyst wall at tapered

end, :::: 0.4 high; rest of sporocyst wall :::: 0.2. Sporocyst residuum

spherical and compact, refractile, 1.3-1.7. Sporozoites vermiform,

reflexed on one end, 6.0-7.1 by 1.3-1.7 (without reflexed portion),

arranged head to tail in sporocyst. Refractile bodies absent.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.

Reference: Molnar and Fernando, 1974.

Eimeria pungitii Molnar and Hanek, 1974

(Fig. 23)

Host: Pungitius pungitius "ninespine stickleback" (Gasterosteiformes) (F).

Description: Oocysts ellipsoid, 12.5 by 9.8 (12.1-13.0 by

8.6-10.4). Wall smooth and colorless, composed of a single layer

:::: 0.1. Micropyle, polar granule, and oocyst residuum absent.

Sporocysts ellipsoid, 10.0 by 3.6 (9.1-10.4 by 3.4-3.9), in contact

with oocyst wall. All four sporocysts arranged lengthwise and

parallel in oocyst. Wall :::: 0.2; Steida body absent. Sporocyst

residuum granular, dispersed, completely filling sporocyst. Sporozoites banana-shaped, 8.7 by 2.3 (8.4-9.1 by 2.1-2.4), arranged

head to tail in sporocyst. Each sporozoite contains a spherical

refractile body.

Sporulation: Most oocysts leave the host unsporulated.

Geographical location: Quebec, Canada.

Location in host: Anterior intestinal epithelium; feces.

Reference: Molnar and Hanek (1974).

Eimeria salvelini Molnar and Hanek, 1974

(Fig. 24)

Host: Salvelinus fontinalis "brook trout" (Salmoniformes) (F).

Description: Oocysts spherical, 12.0 (11.7-12.5). Wall smooth

and colorless, composed of a single layer:::: 0.1. Micropyle, polar

granule, and oocyst residuum absent. Sporocysts ovoid, 9.2 by 5.1

(9.0-9.4 by 5.0-5.3), in contact with oocyst wall. Caplike Stieda

body present at tapered end, 1.1 (0.9-1.3) high; rest of wall :::: 0.2.

Sporocysts arranged three in one plane with fourth overlaying.

Sporocyst residuum spherical and compact, 1.8 (1.5-2.1).

7

Sporozoites vermiform, 6.8 by 1.5 (6.5-7.1 by 1.3-1.7), characteristically interlaced. Refractile bodies absent.

Sporulation: Endogenous.

Geographical location: Quebec, Canada.

Location in host: Anterior intestinal epithelium; feces.

Reference: Molnar and Hanek (1974).

Eimeria squali Fitzgerald, 1975

(Fig. 25)

Host: Squalus acanthias "spiny dogfish shark" (Chondrichthyes) (M).

Description: Oocysts ovoid or ellipsoid, 24.0-36.0 by 20.0-29.0.

Wall brownish lavender, pitted, with the bottoms of the pits light

green or grey. Sporont spherical, 19.9 (16.0-28.0). Micropyle absent; a polar granule is not described. Oocyst residuum:::: 12.4.

Sporocysts ellipsoid, 19.6 by 5.9. Several fibers extend from

pointed ends of sporocysts when forced from oocysts, disappearing shortly after leaving oocyst. A Stieda body is not described.

Sporocyst residuum consisting of several small, spherical granules

in each sporocyst. Sporozoites slender, spindle-shaped, 13.6 by

2.2, slightly twisted in a head to tail arrangement. Each sporozoite

has weakly visible cross striations while still in the sporocyst extending for about one-fifth of its length at the larger end. Refractile body not described but depicted at end opposite to striations in

sporozoite.

Sporulation: Exogenous.

Geographical location: Puget Sound, Washington, U.S.A.

Location in host: Mucosa of spiral valve; feces.

Reference: Fitzgerald (1975).

Eimeria tedlai Molnar and Fernando, 1974

(Fig. 26)

Host: Perca flavescens "yellow perch" (Perciformes) (F).

Description: Oocysts spherical, 9.5-10.0. Wall colorless and of

uniform thickness, consisting of a single layer :::: 0.1. Oocyst

residuum absent; a micropyle is not described. Polar granule present, refractile, 1.0-1.5. Sporocysts ovoid, 8.4-8.7 by 4.5-4.7, in

contact with oocyst wall. Sporocysts usually arranged so that pairs

of parallel sporocysts cross each other. Caplike Stieda body at

tapered end, 1.5-2.0 high; rest of wall :::: 0.3. Sporocyst residuum

spherical or ellipsoid, refractile, 1.5-2.6 by 1.5-2.0. Sporozoites

vermiform, one end reflexed, 6.0-6.5 by 1.6-2.1 (without reflexed

portion), arranged head to tail in sporocyst. Refractile bodies

absent.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Intestinal epithelium; feces.
Reference: Molnar and Fernando (1974).

Eimeria tr-uttae (Leger and Hesse, 1919)

Stankovitch, 1924

(Fig. 27)

Synonym: Goussia truttae Leger and Hesse, 1919.

Hosts: Salvelinus fontinalis "brook trout," Salmo trutta fario

"German brown trout" (Salmoniformes) (F).

Description: Oocysts spherical, 12.8 (12.3-13.0). Wall smooth

and colorless, composed of a single layer:::: 0.1. Micropyle and

oocyst residuum absent. Polar granule present, 0.9 (0.7-1.0).

Sporocysts ovoid, 10.1 by 6.0 (9.3-11.0 by 5.0-6.5), in contact

with oocyst wall. Sporocysts arranged parallel in oocyst. Disclike

Stieda body present at tapered end of sporocyst, 2.7 (2.6-2.8).

Sporocyst residuum spherical and finely granular, 2.8 (2.6-3.0).

Sporozoites vermiform, one end reflexed, 8.5 by 1.6 (7.8-9.0 by

1.3-2.0), (without reflexed portion), arranged head to tail in

sporocyst. Refractile bodies absent.

Sporulation: Endogenous.

Geographical locations: Quebec, Canada; France.

Location in host: Anterior intestinal epithelium; feces.

References: Leger and Hesse (1919); Molnar and Hanek (1974).

Remarks: Description is based predominately on that of Molnar

and Hanek (1974).

In addition to those species listed previously, a number of incidental findings of Eimeria from North American fishes occur in

the literature. Listed below are those which we have located.

Others undoubtedly exist.

Eimeria sp. Davis, 1946

Host: Salvelinus fontinalis "brook trout" (Salmoniformes) (F).

Description: None given.

Sporulation: Endogenous.

Geographical location: Rutland, Vermont, U.S.A.

Location in host: Intestinal epithelium of pyloric caeca; kces.

Reference: Davis (1946).

Remarks: Various developmental stages of an eimerian were

found in the epithelium of the pyloric caeca and, to a lesser ex!ent,

in the epithelium proximal to the caeca. This species may be E.

salvelini or E. truttae, which occur in S. fontinalis in Ontario,

Canada.

Eimeria sp. Fantham and Porter, 1948

Host: Fundulus heteroclitus "mummichog" (Atheriniformes)

(B).

8

Description: None given.

Sporulation: Endogenous (?).

Geographical location: Halifax, Nova Scotia.

Location in host: Unknown.

Reference: Fantham and Porter (1948).

Remarks: This species may be E. funduli (Duszynski e( al.

1979).

Eimeria sp. Fantham and Porter, 1948

Host: Salvelinus fontinalis "brook trout" (Salmoniformes) (F).

Description: None given.

Sporulation: Endogenous.

Geographical location: Quebec, Canada.

Location in host: Intestine.

Reference: Fantham and Porter (1948).

Remarks: This species may be E. salvelini or E. truttae, which

occur in S. fontinalis in Ontario, Canada.

Eimeria sp. Munson, 1974

Host: Liparis atlanticus "seasnail" (Perciformes) (M).

Description: None given.

Sporulation: Endogenous (?).

Geographical location: New Hampshire, U.S.A.

Location in host: Intestinal epithelium; heaviest infections in

posterior intestine.

Reference: Munson (1974).

Remarks: This species was found in 128/128 (100%) of all fish

examined and caused serious histopathological damage. It probably represents an undescribed new species.

Eimeria sp. drias and Noble, 1971

Host: Nezumia bairdi "marlin-spike" or "common grenadier"

(Gadiformes) (M).

Description: Sporocysts averaged 9.5 by 5.5.

Sporulation: Endogenous.

Geographical location: Off coast of Greenland.

Location in host: Stomach epithelium.
Reference: Orias and Noble (1971).

Remarks: This eimerian probably represents an undescribed

new species.

Genus Goussia Labbe. 1896

Oocysts contain four sporocysts, each with two sporozoites;

Stieda body absent; sporocyst wall bivalved.

Goussia caseosa Lorn and DykoV8, 1982

(Fig. 28, sporocyst only)

Host: Macrourus berglax "roughhead grenadier" (Gadiformes)

(M).

Description: Oocysts irregular in shape, 42.0 (40.0-47.0). Wall

thin, membranelike, easily ruptured, and coated with fine

granules. Micropyle and polar granule not mentioned or depicted.

Oocyst residuum absent. Sporocysts ellipsoid, 19.2 by 13.6

(18.0-20.3 by 12.0-15.5), with thick sporocyst wall. Sporocyst

residuum absent. Sporozoites 26.0 by 7.0, arranged head to tail in

sporocyst and partially curled around one another. Anterior ends

of sporozoites with transverse striations, disappearing when

sporozoites excyst. Sporozoite refractile bodies not mentioned but

clearly depicted.

Sporulation: Endogenous.

Geographical location: Grand Banks, Newfoundland.

Location in host: Swim bladder, gas gland, gall bladder, intestinal contents, mesenteric blood vessels.

Reference: Lorn and Dykova (1982a).

Goussia degiustii (Molnar and Fernando, 1974)

DykoV8 and Lorn, 1981

(Fig. 29)

Synonyms: Eimeria spleni DeGiusti and Hnath, 1968; Eimeria

degiusti Molnar and Fernando, 1974.

Hosts: Notropis cornutus "common shiner," Pimephales notatus

"bluntnose minnow," P. promelas "fathead minnow," Campostoma anomalum "central stoneroller" (Cypriniformes) (F).

Description: Oocysts spherical, 19.0-22.0. Wall colorless and of

uniform thickness, consisting of a single layer ::::: 0.1. Oocyst

residuum absent; a micropyle is not described. One or two polar

granules present, 1.0-2.0. Sporocysts ellipsoid, 13.0-14.5 by

6.5-8.5, with a relatively narrow space separating them from

oocyst wall. Wall ::::: 0.2. A thin membranous collar, apparently

visible only with electron microscopy, girdles the sporocyst at the

suture. Sporocyst residuum compact and coarsely granular,

9.0-10.0 by 7.0-8.0. Sporozoites banana-shaped, 10.5-12.0 by

2.5-2.8, arranged head to tail in sporocyst. A spherical refractile

body is present in each sporozoite.

Sporulation: Endogenous.

Geographical location: Ontario, Canada.

Location in host: Spleen.

References: DeGiusti and Hnath (1968); Lorn (1971); Molnar

and Fernando (1974); Dykova and Lorn (1981); Paterson and

Desser (1982).

Remarks: Eimeria spleni was mentioned, but not described, by

DeGiusti and Hnath (1968) and later examined ultrastructurally

by Lorn (1971). However, Molnar and Fernando (1974) described

the morphology of the species and, thus, E. spleni is a nomen

nudum.

Goussia gadi (Fiebiger, 1913) Dykov8 and Lorn, 1981

(Fig. 30)

Synonym: Eimeria gadi Fiebiger, 1913.

Hosts: Gadus (=Melanogrammus) aeglefinus "haddock," G. morrhua "common cod" or "codfish," G. virens "coal-fish," Enchelyopus cimbrius "fourbeard rock ling," Molva vulgaris "ling"

(Gadiformes) (M).

Description: Oocysts spherical, 26.0-28.0, with a smooth thin

wall. Micropyle and polar granule not described. Oocyst residuum

sometimes present. Sporocysts ovoid, 11.0-15.0 by 7.5-10.0, with

apical pore and longitudinal ridges extending length of sporocyst

(not visible with light microscopy). Sporocyst residuum present.

Sporozoites 16.0 by 4.0, and contain one or more refractile

bodies.

Sporulation: Endogenous.

Geographical location: North Atlantic Ocean.

Location in host: Swim bladder.

References: Fiebiger (1913); Odense and Logan (1976); Scott

(1981); Schulman and Shtein (1962); Dykova and Lorn (1981).

Remarks: This description is based on that given by Fiebiger

(1913) and Odense and Logan (1976).

KEY TO EIMERIIDAE OF NORTH AMERICAN FISHES

1a. Sporocysts nonbivalved; with or without Stieda body (Eimeria) 2

1b. Sporocysts bivalved; without Stieda body (Goussia)4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

4 All tetrasporocystic oocysts which have dizoic sporocysts and lack Stieda bodies may eventually prove to be Goussia. The suture is often difficult or impossible to see with light

microscopy. However, both Odense and Logan (1976) and Lorn and Dykovil (1982a) have published photomicrographs which suggest that crushing sporocysts between slide and

cQverglass can separate the valves at the suture; this making electron microscopy unnecessary for diagnosis.

9
2a. Stieda body present; oocysts spherical; oocyst residuum absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2b. Stieda body absent; oocysts vary in shape " 10

3a. Oocysts> 20 lim with wall:::: 1.0 lim thick; sporocysts with sporopodia; sporulation endogenous, in liver of Atheriniformes

............................................................... " E·funduli

3b. Oocysts < 15 lim with wall:::: 1.0 lim; sporocysts without sporopodia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4a. One or more polar granules present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4b. Polar granule absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

5a. Stieda body disc-shaped; sporulation endogenous, in anterior intestinal epithelium of Salmoniformes E. truttae

5b. Stieda body not as above 6

6a. Stieda body caplike at tapered end of sporocyst; sporulation endogenous, in intestinal epithelium of Perciformes E. tedlai

6b. Stieda body a knoblike thickening at tapered end of sporocyst 7

7a. Oocyst small, < 10 lim in greatest diameter; sporulation endogenous, in intestinal epithelium of Perciformes E. moronei

7b. Oocyst> 10 lim in diameter 8

8a. Sporocysts 8-10 lim long and in contact with oocyst wall; sporulation endogenous, in intestinal epithelium of Perciformes

................................................................................................E. glenorensis 5

8b. Sporocysts 8 lim long leaving space separating them from oocyst wall; sporulation endogenous, in intestinal epithelium

of Salmoniformes E. osmeri 5

9a. Stieda body formed by 4 papillae, sporocysts hexagonal in cross-section; sporulation endogenous, in anterior intestinal

epithelium of Anguilliformes E. anguillae

9b. Stieda body caplike, sporocyst round in cross-section; sporulation endogenous, in anterior intestinal epithelium of Salmoniformes E. salvelini

lOa. Oocyst residuum present II

lOb. Oocyst residuum absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

II a. Oocyst spheroid to ovoid, outer wall smooth; sporulation endogenous, oocysts in testes of Clupeiformes E. brevoortiana

11 b. Oocyst ellipsoid to ovoid, outer wall pitted, brown; sporulation exogenous, merogony in spiral valve mucosa of Chondrichthyes E. squali

12a. Oocyst spheroid, > 30 lim in diameter, with thick (3 lim) wall; sporulation endogenous, in anterior intestinal mucosa of

Scorpaeniformes E. myoxocephali

12b. Oocyst spheroid or otherwise, <30 lim in any dimension, with thin (:::: 0.1 lim) wall. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13a. One or more polar granules present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

13b. Polar granule absent. . . . . . . . . . . . . . . . . . . . . . . . .. 17

14a. Oocyst spheroid, < 10 lim in diameter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

14b. Oocyst ellipsoid :....................... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

15a. Sporocysts ovoid, flat on one side; sporozoites without refractile body; sporulation endogenous, in anterior intestinal epithelium of Cypriniformes E. catostomi

15b. Sporocysts ellipsoid, not flat on one side, sporozoites with refractile body; sporulation endogenous, in intestinal epithelium

of Gasterosteiformes E. haneki

16a. Oocyst small, greatest length < 20 lim; sporulation endogenous, in anterior intestinal epithelium of Salmoniformes E. hoffmani

16b. Oocyst large, smallest length> 20 lim; sporulation endogenous, in kidneys and renal parenchyma of Cypriniformes E. freemani

17 a. Sporocyst residuum absent; sporulation exogenous, merogony in anterior intestinal epithelium of Cypriniformes E. aurati

17b. Sporocyst residuum present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18a. Oocyst irregular in shape with wall tightly adhering to sporocysts; sporulation endogenous, in intestinal epithelium of

Scorpaeniformes E. duszynskii

18b. Oocyst spheroid or ellipsoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

19a. Oocyst ellipsoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

19b. Oocyst spheroid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

20a. Oocyst 7.5-9 x 6-7 lim, sporocysts 6-8 x 2-3.5 lim; sporulation endogenous, in anterior intestinal epithelium of Cypriniformes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . E. fernandoae

20b. Oocyst 12-13 x 8-10.5 lim, sporocysts 9-10.5 x 3-4 lim; sporulation mostly exogenous; merogony in anterior intestinal

epithelium of Ga&terosteiformes E. pungitii

21 a. Oocyst smallest diameter> 15 lim, sporulation endogenous in liver of Gasterosteiformes E. gasterostei

21 b. Oocyst largest diameter 15 /-1m; sporocysts elongate; sporulation endogenous in gut epithelium 22

22a. Spo.rocysts appear triangular in cross section, with granular residuum 23

22b. Sporocysts not triangular in cross section. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

23a. Sporocyst residuum dispersed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

23b. Sporocyst residuum compact, coarsely granular; ellipsoid sporocysts always < 9.0 /-1m long; sporulation endogenous in

intestinal epithelium of Siluriformes E. ictaluri6

'Oocysts of these "species" are nearly identical and, in fact, may represent the same species. In their original description, Molnar and Fernando (1974) argued, "It is well

known that Eimeria spp. are highly specific to single or closely related host species" and" ... when morphologically almost indistinguishable or very similar Eimeria spp. occur in

distantly related host species, we consider these parasites as distinct species.... " This assumption may prove invalid with time, but our knowledge of the life cycles of fish

eimerians is too incomplete to refute it at this time.

·Oocysts of these "species" are nearly identical and may represent a single species. See footnote 5.

10
24a. Sporocyst residuum finely granular, sporocysts always> 9.0 ~m long; sporulation endogenous in intestinal epithelium of

Perciformes . E. laureleus6

24b. Sporocyst residuum granules of various sizes; sporulation endogenous, in intestinal epithelium of Scorpaeniformes E. ojibwana6

25a. Both oocyst diameter and sporocyst smallest length> 11.0 ~m; sporulation mostly exogenous, merogony and some sporulation in anterior intestinal epithelium of Perciformes E. micropteri6

25b. Oocyst diameter and sporocyst length not as above . . . . . . . . . . . . . . . 26

26a. Oocyst diameter always> 14.0 ~m; sporocyst length always";;; 10.0 ~m; sporulation endogenous in intestinal epithelium of

Cypriniformes. . . . . . . . . . . . . . . . . . . . . . . . . . E. hybognathi6

26b. Oocyst and sporocyst measurements not as above 27

27a. Sporozoite longer than sporocyst with one end reflexed back upon itself; sporulation endogenous, in intestinal epithelium

of Cypriniformes E. iroquoina6

27b. Sporozoites slightly shorter than length of sporocyst, rarely reflexed; sporulation endogenous and exogenous; merogony in

anterior intestinal epithelium of Cypriniformes E. etheostomae

28a. Oocysts irregular in shape, with thin, membranelike wall; sporulation endogenous in numerous tissues of Gadiformes .... G. caseosa

28b. Oocysts spherical. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

29a. One or more polar granules present; sporulation endogenous, in spleen of Cypriniformes G. degiustii

29b. Polar granule absent; sporulation endogenous, in swim bladder of Gadiformes G. gadi

ACKNOWLEDGMENTS

We wish to express thanks to Dorothy Sanders and Jackie

Taylor for kindly typing the manuscript.

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11

HARDCASTLE, A. B.

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12

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4a

4b

10llm

Figures 1·30.-Composite line drawings of oocysts and/or sporocysts of Eimeriidae of North American fish. Minor discrepancies may exist between drawings and text.

All figures redrawn from previously published drawings or photomicrographs. 1. Eimeria anguillae (from Molnar and Hanek 1974); 2. E. aurmi (from Hoffman 19(5); 3.

E. brevoortiana (from Hardcastle 1944); 4. E. catostomi (from Molnar and Hanek 1974) (a) oocyst, entire view (b) section through oocyst showing position of polar

granule; 5. E. duszynskii (from Conder et al. 1980); 6. E. etheostomae (from Molnar and Hanek 1974); 7. E. fernandoae (from Molnar and Hanek 1974); 8. E. freemani

13
10

(from Molnar and Fernando 1974); 9. E. funduli (from Duszynski el al. 1979; pers. obs.); 10. E. gasterostei (from Schulman and Shleln 1962); 11. E. glenorensis (from

Molnar and Fernando 1974); 12. E. haneki (from Molnar and Fernando 1974); 13. E. hoffmani (from Molnar and Danek 1974); 14. E. hybognalhi (from Molnar and Fer14
19

nando 1974); 15. E. ictaluri (from Molnar and Fernando 1974); 16.~. iroquoina (from Molnar and Fernando 1974); 17. E. laureleus (from Molnar and Fernando 1974);

18. E. micropteri (from Molnar and Fernando 1974); 19. E. moronei (from Molnar & Fernando 1974); 20. E. myoxocephali (from Fitzgerald 1975); 21. E. ojibwana (from

15
10Jun

Molnar and Danek 1974); 22. E. Dsmeri (from Molnar and Fernando 1974); 23. E. pungitii (from Molnar and Danek 1974); 24. E. salvelini (from Molnar and Danek

16
24

. t~/ :."

r..·/

.\ ·(f

25

10Jlm

1974); 25. E. squali (from Fitzgerald 1975); 26. E. tedlai (from Molnar and Fernando 1974); 27. E. Irullae (from Molnar and Danek 1974); 28. Sporocyst of Goussia

17
28

30

10um

caseosa (from Lorn and DykoV8 1982a); 29. G. degu;stii (from Molnar and Fernando 1974); 30. G. gad; (from Fiebiger 1913; Odense and Logan 1976).

18