NOAA Technical Rep_o_rt_N_M_F_S_I_2_1 A_u..gu_s_t1_9_94

Marine Flora and Fauna of the

Eastern United States

Platyhelminthes: Monogenea

Sherman S. Hendrix

u.s. Department of Commerce
u.s. Deparnnent

of Conunerce

Ronald H. Brown

Secretary

National Oceanic and

Atmospheric AdnUnistration

D. James Baker

Under Secretary for Oceans and

Atmosphere

National Marine

Fisheries Service

Rolland A. Schmitten

Assistant Administrator for

Fisheries

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NOAA Technical Report NMFS 121

A Technical Report of the Fishery Bulletin

Marine Flora and Fauna of the

Eastern United States

Platyhelminthes: Monogenea

Sherman S. Hendrix

August 1994

U.S. Department of Commerce

Seattle, Washington
Foreword _

This NOAA Technical Report NMFS is part of the subseries "Marine Flora and

Fauna of the Eastern United States" (formerly "Marine Flora and Fauna of the

Northeastern United States"), which consists of original, illustrated, and modern manuals on the identification, classification, and general biology of the

estuarine and coastal marine plants and animals of the eastern United States.

The manuals are published at irregular intervals on as many taxa of the region

as there are specialists available to collaborate in their preparation. These

manuals are intended for use by students, biologists, biological oceanographers, informed laymen, and others wishing to identify coastal organisms for

this region. They can often serve as guides to additional information about

species or groups.

The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region," edited by R. I. Smith and produced in 1964

under the auspices of the Systematics Ecology Program, Marine Biological

Laboratory, Woods Hole, Massachusetts. Geographic coverage of the "Marine

Flora and Fauna of the Eastern United States" is planned to include organisms

from the headwaters of estuaries seaward to approximately the 200-m depth on

the continental shelf from Maine to Florida, but may vary somewhat with each

major taxon and the interests of collaborators. Whenever possible, representative specimens dealt with in the manuals are deposited in the reference collections of major museums.

The "Marine Flora and Fauna of the Eastern United States" is being prepared

in collaboration with systematic specialists in the United States and abroad.

Each manual is based primarily on recent and ongoing revisionary systematic

research and a fresh examination of the plants and animals. Each manual treats

a separate major taxon and includes an introduction, illustrated glossary, uniform and originally illustrated keys, an annotated checklist (with available

information on distribution, habitat, life history, and related biology), references to the major literature of the group, and a systematic index.

ii
Marine Flora and Fauna of the Eastern United States

Platyhelminthes: Monogenea

SHERMAN S. HENDRIX

Department of Biology

Gettysburg College

Gettysburg, PA 17325

ABSTRACT

This manual includes an introduction to the general biology of the platyhelminth class

Monogenea, an illustrated key to the generic level, an annotated systematic list of 108

species in 75 genera and 28 families, a glossary of terms, a host-parasite list, a selected

bibliography, and a systematic index. The species listed have been reported on fishes taken

in the coastal waters of the northwestern Atlantic Ocean from the U.S.-Canada border to

Beaufort, North Carolina. Many of the species, however, have wider geographic distributions that are not reported herein unless adjacent to the geographic area above. In addition,

a single species of Neodermata, Udonella caligorum, that is found attached to copepods on

fish gills is also listed.

Introduction

All monogeneans are parasitic on or, occasionally, parasitic in marine and freshwater fishes, in the urinary

bladders of some turtles and amphibians, and on the

eye of Hippopotamus. Most species are ectoparasitic on

the gil\ filaments of their fish hosts; some are ectoparasitic on fins, on the body surface, in the nostrils, and in

the buccal cavity; others are endoparasitic in the esophagus, cloaca, urinary tract, and (in at least one juvenile)

the heart. Most monogeneans are highly host specific

(Hargis, 1957b), which aids in the specific identification of worms from a particular host. Many of the

Monogenea reported from marine and estuarine fishes

along the northeastern coast ofthe United States (Maine

to North Carolina) have a more extensive, sometimes

cosmopolitan, geographic distribution than is covered

in this manual. Only a fraction of potential fish hosts

have been reported with these parasites. Because many

host species have not yet been examined for these

helminths, much remains to be done to expand both

geographic ranges and host records for Monogenea.

All known Monogenea are hermaphroditic. Most are

oviparous (Kearn, 1986) with a short-lived free-swimming ciliated larva, the oncomiracidium (Llewellyn,

1963, 1968, and 1972). Oncomiracidia of at least three

marine species are known to seek their specific hosts

primarily by chemotaxis, responding to components in

fish mucus (Kearn, 1967, 1973, and 1974a; Whittington

and Kearn, 1986 and 1990; Whittington, 1987, a and b).

Once attached, larvae shed the ciliated epidermis and,

as juveniles, remain in place or migrate to the normal

site of development, e.g. the ventral surface skin or

gills, where they mature to adults. The "viviparous

Monogenea," such as Gyrodactylus spp., give birth to

non-ciliated larvae, which are transmitted to new hosts

by direct contact, either as larvae or adult worms.

The biology of monogeneans found along the U.S.

Atlantic coast is still poorly known. Most studies on

these worms comprise new species descriptions or surveys that give data on prevalence, intensity ofinfection,

or both (e.g. McMahon, 1963 and 1964; Meyers, 1978),

although a few publications deal with the ciliated oncomiracidial stage (e.g. Kingston et aI., 1969), microecology (e.g. Suydam, 1971), or pathology (e.g. Lawler,

1977; Overstreet, 1978 and 1992).

Monogenea is now generally recognized as a separate

class within the phylum Platyhelminthes, rather than a

subclass ofthe class Trematoda. However, considerable

controversy remains in the recognition of evolutionary

relationships among the higher taxa within the

Monogenea (Bychowsky, 1937 and 1957; Llewellyn, 1970

and 1981; Gusev, 1978; Lebedev, 1979 and 1988; Mamaev

and Lebedev, 1979; Lambert, 1980, a and b; Malmberg,

1982,1986, and 1990; Rohde, 1990;Justine, 1991; Boeger

and Kritsky, 1993). To present the marine monogene-
2 NOAA Technical Report NMFS 121

ans in this manual, I generally follow the system proposed by Llewellyn (1970), as modified by BeverleyBurton (1984) for the orders and families found on

Canadian hosts. The exceptions are that Udonella has

been removed from the Turbellaria, the microbothriids

assigned to the order Microbothriidea Lebedev, 1988,

and the monocotylids assigned to the order Monocoty-

!idea Lebedev, 1988.

Only adult monogeneans are treated in this manual.

The free-swimming oncomiracidial stage is not covered

except to mention, in the systematic section, those species for which a description has been given. Monogenean species from estuarine and continental shelffishes

are included while species from offshore deep-water

fishes are not. The northern extreme ofthe geographic

range covered herein is the U.S.-Canada border; the

southern extreme is Beaufort, North Carolina. Many of

the monogeneans listed below have a wider geographic

distribution. Generally, only those reports from adjacent geographic regions are included; reports from

other parts of the globe are not.

Collection and Examination _

Collecting monogeneans from freshly caught marine

fishes most often involves examining the body surface

and fins before removing gills by the following methods:

1. Place the gills in a separate container of dilute

formalin, e.g. 1 part concentrated (40% formaldehyde

or "100%") formalin to 4,000 parts seawater (Pritchard

and Kruse, 1982), which relaxes and fixes the worms

with a minimum of contraction or distortion.

2. Examine the fins, skin, scales, buccal cavity, nasal

capsules, and cloaca of the fishes for monogeneans.

3. Leave the material in the dilute formalin solution

for approximately one hour, which is sufficient time to

relax most specimens.

4. Shake the container vigorously for about a minute

to dislodge worms.

5. Pour the liquid into a cylinder or other tall container and let it stand for several minutes to allow the

worms to settle.

6. Decant the supernatant fluid and examine the

sediment in a petri dish under a dissecting microscope.

7. Pipet worms into a fixative, such as 5% formalin or

AFA (alcohol-formalin-acetic acid).

An alternative method is to place the heads or whole

fish directly into a fixative, such as 10% formalin or

AFA, at a proportion of 2-3 parts fixative to 1 part gill

material.

Alternative methods for relaxing and isolating worms

are given by Pritchard and Kruse (1982). For example,

freezing the gills, branchial basket, or even whole fishes

for 6-12 hours often aids in preventing mucus production by gill tissues and kills the worms in a relaxed state.

However, for studies involving transmission electron

microscopy, physiology, or behavior, carefully remove

living worms from the host with the aid of a dissecting

microscope. For scanning electron microscopy thoroughly wash worms by vigorous shaking in several

changes of artificial seawater to remove attached mucus

and debris before fixation (Halton, 1974).

After specimens have been in the fixative for 12-24

hours, transfer specimens to vials that contain internal

labels and 70% ethanol for storage. However, specimens may be left in the formalin fixative almost indefinitely. Because of the importance of hamuli and marginal hooklets in taxonomy, small monogeneans such

as Gyrodactylus spp. are usually mounted unstained on

slides in glyceroljelly by using a double coverslip method

(Pritchard and Kruse, 1982). However, another staining method uses Gormori's trichrome solution with

good results (Kritsky et al., 1978).

Treat larger monogeneans in a manner similar to the

whole mount preparation techniques employed for digenetic trematodes. After fixing, store the worms in

70% ethanol until stained. Most staining procedures

use either alcoholic carmine or aqueous hematoxylin.

Several general parasitology laboratory manuals (e.g.

Pritchard and Kruse, 1982; Meyer et al., 1992) provide

detailed accounts of the fixation, staining, dehydration,

clearing, and mounting techniques employed to study

these organisms. Cooper (1988) described the preparation ofserial sections of platyhelminth parasites, which

are useful in tracing the location of ducts and other

structures.

Several structures are important for the identification of Monogenea. Most important for identification

of monogenean genera is the posterior attachment organ, or haptor, and its associated hard (sclerotized)

structures (Figs. 1-4). The shape and nature of the

anterior attachment structures, reproductive system,

and digestive system are also important in keying out

these worms (Figs. 1 and 2). The anterior attachment

organ may comprise a pair of concave disklike structures, a pair of buccal suckers, head organs (paired,

glandular duct openings), or a single, weak, oral sucker.

The number and placement of the testes, presence or

absence and shape and number of spines within the

male copulatory complex, shape and position of the

ovary and uterus, and position of the vagina(e) and

genital pore are all useful diagnostic characters. Eggs,

when present in the uterus or ootype, can also aid

identification. The intestine usually consists of a pair of

straight or highly branched ceca that end blindly or are

confluent at the posterior end ofthe body. However, in

some of the larger species the intestinal ceca may be

obscured by extensive vitellaria. In some taxa the shape

of the pharynx is of taxonomic value.
Hendrix: Monogenea of the Eastern United States 3

mouth

oral sucker

pharynx·

genital atrium

uterus

vas deferens

vaginal pore

ootype with glands

oviduct

vagina

vitelline duct

distal part of ovary

ovary

intestinal cecum

testis

vitellaria

loculus

septum

hamulus

Figure 1

Dasybalolrema dasybalis (family Monocotylidae), generalized composite anatomy, ventral

view. Scale: 1 mm.
4 NOAA Technical Report NMFS 121

Figure 2

Pauciconfilntla pogoniae (family Microcotylidae), generalized

composite anatomy, ventral view. Scale: 2 mm.

•••• • •

·.·•.

• •

••

·•

·.·.

··./

• I .' • I ••j

·, • I "I

..::1

..:-1

• I I','..:;~'+f+-++--!------ •• ej

..

•• 0

•

o

..

..·

••·

·:.·

.. ..

..

..

buccal sucker

pharynx

esophagus

genital atrium

uterus

vitelline duct

ovary

oviduct

ootype

testis

intestinal cecum

vitellaria

clamp

haptor

Monogeneans included in the key of this manual are

designated by a specific name when only one member

of the genus is known from the coastal waters of the

western Atlantic from the U.S.-Canada border to Beaufort, North Carolina. Where more than one representative of a genus has been reported, the key is to genus

and the reader should consult the systematic account

and the host-parasite list for more information at the

species level. Drawings included in the key were made

with the aid of a microprojector. The majority of illustrations are from specimens borrowed from the U.S.

National Museum Helminth Collection, Biosystematics

Laboratory, U.S. Department of Agriculture-Agricultural Research Service (USDA-ARS), Beltsville, MD

20705, and were usually the holotype or a paratype of

the species.

Glossary _

Anchors. See hamuli.

Anterior adhesive area. Anterior attachment organs,

which are usually sessile or disklike and produce a

sticky adhesive substance that allows the worm to

cling to the host. (See Fig. 3, A and B.)

Bifurcation. A branching, such as the intestine at the

terminus of the esophagus, divided into two parts.

Buccal suckers. Paired muscular suckers opening laterally into the buccal cavity ofsome polyopisthocotyleans. (See Fig. 2.)

Clamps. Haptoral sclerotized structures that are found

in many polyopisthocotyleans and consist of a framework of sclerites, capable of grasping host gill tissue

between a pair of valves. The size, shape, arrangement, number of clamp sclerites, and number of

clamps are of major taxonomic importance in differentiating taxa. (See Fig. 4, C and D.)

Dorsal marginal spines. Small spines ofvarious shapes

arranged in rows along the dorsal body margin in

some capsalids. (See Figs. 12B and 13B.)

Genital atrium. A cavity, often muscular and containing distinctive spines, which receives the terminal

ducts of both the male and female reproductive systems. (See Figs. 1 and 2.)

Genital corona. Clusters or circles of spines within or

at the opening of the genital atrium. (See Figs. 47B

and 71B.)

Genito-intestinal canal. A small duct, found only in

polyopisthocotyleans, connecting the oviduct to the

right intestinal cecum.

Hamuli. The paired sclerotized hooklike structures of

the haptor. The hamuli are movable and attach the

parasite to the host. Much ofthe American literature

refers to these structures as anchors. (See Figs. 1 and

3, C and D.)
Hendrix: Monogenea of the Eastern United States 5

A B

duct

opening

-l-....;.,.,.;,.---"""*----- pharynx ------+---+,:

adhesive

gland

c 0

marginal

hook

accessory

sclerite

transverse

bar

hamulus

Figure 3

Undivided disklike haptors and glandular anterior adhesive areas: (A) glandular anterior

adhesive area; (B) anterior adhesive area with three pairs of head organs; (C) haptor with two

pairs of hamuli and a pair of accessory sclerites; (D) haptor with two pairs of hamuli, two

transverse bars, and marginal hooks.

Haptor. The posterior attachment organ which usually

carries additional components such as marginal hooks,

hamuli, accessory sclerites, armed suckers, or clamps.

The structure and its hard parts are ofmajor taxonomic

importance in differentiating the orders, families, and

genera. (See Figs. 1,2,3, C and D, 4, A-D.)

Haptoral appendix. A muscular extension of the

haptor having two terminal suckers and usually hamuli, characteristic of hexostomatids. (See Fig. 4A.)

Head organ. Groups of anterolateral postpharyngeal

glands that open via the head organs and produce a

sticky secretion that allows the anterior end of a

worm to adhere temporarily to the host while feeding. (See Fig. 3, A and B.)

Intestinal ceca. The intestine, which can be a single

(=cecum) unbranched tube ending blindly (see Fig.

8), two (=ceca) unbranched or highly branched tubes

that may end blindly (see Fig. 9), or be posteriorly

confluent (see Fig. 16). Ceca are sometimes obscured

by extensive vitellaria (see Fig. 21).

Loculi. Haptoral depressions, usually limited by septa,

found in the capsalids and monocotylids. (See

Fig.!.)

Male copulatory complex (apparatus). An often muscular complex that may be armed with a sclerotized

stylet or other accessory structures. (See Fig. 82D.)

Marginal hooks. Radially arranged pairs ofsmall hooks

on the margin of the larval haptor, which persist in

the adult haptor of many taxa, such as the gyrodactylids and dactylogyrids. (See Fig. 3D.)

Oncomiracidium. The ciliated larval stage, which

hatches from the egg, typical of the Monogenea.

Opisthaptor. See haptor.

Oral sucker. A muscular sucker surrounding the

mouth ofsome Monogenea. (See Fig.!.)

Pedunculate. Descriptive of a structure borne on a

stalk. (See Fig. 52.)

Peduncular bar. A sclerotized transverse bar, which is

usually pitted and found immediately anterior to the

haptor in some gyrodactylids. (See Fig. 34.)
6 NOAA Technical Report NMFS 121

B

appendix

intestinal

cecum

sucker-----v

sclerite

hamulus ------~=_7

c ~~.... . ... ::: e.::. .... .. .... :.. -... ..... .:-:..:-.-:. ... :..-. .. .:-.--":--------- .... . .. .. ... . .. . . ... .... .' . .'... clamp

Figure 4

Representative polyopisthocotylean haptors: (A) haptor with appendix and three pairs ofpedunculate

suckers, each containing a sclerite; (B) haptor with four pairs of sessile suckers, each containing

sclerites; (C) asymmetric haptor with multiple clamps; (D) symmetric haptor with multiple clamps.

Pharynx. A muscular pumping organ lying between

the mouth and esophagus that aids in ingestion of

food. (See Figs. 1 and 11, A and B.)

Preoral suckers. A pair of weakly muscular disklike

structuresjust anterior and lateral to the mouth. See

anterior adhesive organs above. (See Fig. 12A.)

Prohaptor. An obsolete term for the unarmed holdfast organ (anterior adhesive area) at the anterior

end of the body bearing suckers and glands or both.

The nature of this organ is of taxonomic value. (See

Fig. 3, A and B.)

Pseudohaptor. A functional posterior organ characteristic of Pseudacanthocotyla, which is not derived

from the larval haptor but which develops just anterior to it. (See Fig. 5.)

Pseudosucker. A weakly muscular, funnel-like structure

surrounding the mouth. It may contain glandular openings around the margin as in Dasybatotrema. (See Fig. 24A.)
Ridge sclerites. Smallsclerotized structuresfound in the

rims ofloculi in some monocotylids. (See Fig. 26C.)

Sclerites. Sclerotized supporting structures associated

with hamuli, clamps, or suckers of the haptor (see

Fig. 3C and 4, A and B), vaginae, or male copulatory

complex.

Septa. Ridges that divide the haptor of capsalid and

monocotylid monogeneans into loculi. (See Fig. 1.)

Squamodisc. A circular or oval structure associated with

the haptor and consisting of concentric rows of small

spines characteristic of diplectanids. (See Fig. 368.)

Suckers. Muscular attachment devices found primaHendrix: Monogenea of the Eastern United States 7

rily on the haptor of polyopisthocotyleans. (See Fig.

4, A and 8.)

Terminal lappet. A sclerite-bearing flap of tissue at

the posterior end of the haptor ofsome polyopisthocotyleans. (See Fig. 65C.)

Transverse supporting bar. Transverse sclerotized bars

associated or articulating with hamuli in many monogeneans, e.g. Dactylogyrida. (See Fig. 3D.)

Vitellaria. Usually extensive glands that produce nutrients for the development of the oncomiracidium

and proteins for the formation of eggshells. (See

Figs. 1 and 2.)
8 NOAA Technical Report NMFS 121

Key to the Monogenea Parasitic on Marine Fishes

of the Eastern United States _

1 Haptor well developed, undivided and disklike, with or without hamuli; haptoral suckers and

clamps absent; anterior adhesive area usually with glands opening via head organs, bilobed sessile,

or disklike areas (see Fig. 3, A and B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

1 Haptor divided, usually with 4 or more pairs of suckers or clamps, pedunculate or sessile, hamuli

present or absent; anterior region containing buccal suckers or weak oral sucker (see Figs. 2 and

4, A-D.) (Order Polyopisthocotylida) 39

2(1) Posterior attachment organ a pseudohaptor with numerous radiating rows of spines, larval haptor

with 16 marginal hooks, undeveloped, at posterior margin of pseudohaptor. Parasitic on skates

(Rajidae). . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Acanthocotylidae) Pseudacanthocotyla verrilli (Fig. 5)

2(1) Posterior attachment organ a haptor lacking radial rows ofspines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3

3(2) Haptor distinctly muscular, disklike, with or without 1 or 2 pairs of hamuli and anterior sclerotized

accessory sclerites and lacking transverse supporting bars; anterior adhesive area disklike or

glandular (see Fig. 6) 4

Figure 5

Pseudacanthocotyla verrilli, ventral view.

Scale: 1 mm. Drawn from USNM 7175.
Hendrix: Monogenea of the Eastern United States 9

3(2) Haptor a membranous disk, with 1 or 2 pairs of hamuli, each pair supported by a transverse bar;

anterior adhesive area glandular (see Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 27

4(3) Intestine with unbranched cecum. Usually parasites of ectoparasitic copepods on fish gills, rarely

attached to gill tissue (Class Udonellidea, FamilyUdonellidae) Udanella caligarum (Fig. 8)

4(3) Intestine with 1 or 2 ceca, unbranched or diverticulate. Parasites of fishes and not crustacea 5

5(4) Haptor disklike, with or without septa, hamuli usually present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6

5(4) Haptor small and muscular, lacking septa and hamuli. Parasitic on requiem sharks (Carcharhinidae).

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Microbothriidae) 25

6(5) Intestine with 2 ceca with numerous diverticula. Usually parasitic on teleosts and, occasionally,

elasmobranchs. (Family Capsalidae) 7

Figure 6

Muscular haptor with two pairs of hamuli and a pair of accessory sclerites.

Figure 7

Haptor with two pairs of hamuli, each supported by a transverse

bar, and marginal hooks.

Figure 8

Udonella caligorum, ventral view.

Scale: 0.5 mm. Mter Price (1938b).
10 NOAA Technical Report NMFS 121

6(5) Intestine with 1 or 2 unbranched ceca. Parasitic on elasmobranchs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

7(6) Testes 2 8

7(6) Testes more than 2 9

8(7) Haptor aseptate, with papillae in posterior half; anterior adhesive area sessile. Parasitic on the

roughtail stingray, Dasyatis centroura, and Atlantic halibut, Hippoglossus hippoglossus . .. Enrobdella spp. (Fig. 9)

]

]

B

c

Figure 9

Entobdella hippoglossi: (A)

ventral view. Scale: 2 mm;

(B) accessory sclerite. Scale:

0.25 mm; (C) anterior hamulus. Scale: 0.25 mm; (D)

posterior hamulus. Scale:

0.2 mm. Drawn from USNM

8149.

o ...

. .. .. .

o 0 0 Q 0

,j :,r,c,,"o' "-' C 0 (]

'00000°°0:)0 a

0 000 °0 C) () 0

.0 0000°

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° 0

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°00°0°0°~(]..o.oooo~~~.. o. 0o .. 0/ ., 0 0 0 0 (]:) .' .... ,

0 00 °0 0 '

...,..:.' ' ,,-:,'"
8(7)

Hendrix: Monogenea of the Eastern United States 11

Haptor aseptate, lacking papillae; anterior adhesive area disklike. Parasitic on the cownose ray,

Rhinoptera bonasus Benedeniella posterocolpa (Fig. 10)

9(7)

9(7)

Pharynx constricted near middle (see Fig. IIA).

Pharynx not constricted (see Fig. 11B) .

8

c

o ]

10

14

Figure 10

Benedeniella posterocolpa: (A) ventral view. Scale: 2 mm; (B) accessory sclerite. Scale: 0.5

mm; (C) anterior hamulus. Scale: 0.5 mm; (D) posterior hamulus. Scale: 0.2 mm. Drawn

from USNM 38146.

A B

Figure 11

Pharynx shapes: (A) constricted; (B) not constricted.
12 NOAA Technical Report NMFS 121

10(9)

10(9)

Testes extend laterally to intestinal diverticula.

Testes confined to intracecal area; parasitic on tunas (Scombridae) ....

11

13

11(10) Ratio of disklike anterior adhesive area diameter to haptor diameter is 1:2; one pair of accessory

sclerites present in septate haptor. Parasitic on Istiophoridae and Scombridae.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Tristomella spp. (Fig. 12)

.. . . . . .. ... . . . .. ...

.. :..

B

c

Figure 12

Tristomella laevis: (A) ventral view. Scale: 3 mm; (B) dorsal marginal spines; (C) hamulus. Scale: 0.55 mm. Drawn from

USNM 18874.
Hendrix: Monogenea of the Eastern United States 13

11(10) Ratio of diameter ofsuckerlike anterior adhesive area to haptor diameter less than 1:3; hamuli and

sclerites absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12(11) Ratio of diameter of anterior adhesive area to haptor diameter approximately 1:4. Parasitic on

ocean sunfish, Mola mola. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Capsala martinierei (Fig. 13)

.c

',0

A 8

Figure 13

Capsala martinierei: (A) ventral view. Scale: 10 mm; (B) dorsal marginal spines. Drawn from USNM 41146.
14 NOAA Technical Report NMFS 121

12(11) Ratio of diameter of anterior adhesive area to haptor diameter from 1:5 to 1:6. Parasitic on ocean

sunfish Tricotyla molae (Fig. 14)

B

Figure 14

Tricotyla molae: (A) ventral view. Scale: 3 mm; (B) dorsal marginal spines. Drawn from USNM 8152.
Hendrix: Monogenea of the Eastern United States 15

13(10) Parasitic in nasal capsules of tunas (Scombridae) Nasicola klawei (Fig. 15)

B

Figure 15

Nasicola klawei: (A) ventral view. Scale: 2 mm; (B) dorsal marginal spines. Drawn from USNM 59865.
16 NOAA Technical Report NMFS 121

13(10) Parasitic on gills of little tunny, Euthynnus alletteratus Caballerocotyla manteri (Fig. 16)

14(9)

14(9)

Haptor aseptate; anterior adhesive area with 2 shallow grooves. Parasitic on sturgeons, Acipenserspp.

Nitzschia spp. (Fig. 17)

Haptor septate; anterior adhesive area suckerlike or disklike. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

..

... . .. . .... .. .. . .. . ......

Figure 16

Caballerocotyla manteri: ventral view. Scale: 0.5 mm. Drawn from

USNM 37228.

Figure 17

Nitzschia sturionis: ventral view. Scale: 3 mm.

Drawn from USNM 73135.
Hendrix: Monogenea of the Eastern United States 17

15(14) Haptor with 7 marginal and 1 central loculi; dorsal marginal spines serrated or bifid. Parasitic on

swordfish, Xiphias gladius, and smooth hammerhead, Sphyrna zygaena. . . . . . . . . . .. Tristoma spp. (Fig. 18)

B

1J

Figure 18

Tristoma coccineum: (A) ventral view. Scale: 2 mm; (B) dorsal marginal spines. Drawn from USNM 35645.
18 NOAA Technical Report NMFS 121

15(14) Haptor with 9 marginal and 1 central loculi; dorsal marginal spines wider than long, with

numerous small cusps. Parasitic on white marlin, Tetrapturus albidus. Capsaloides spp. (Fig. 19)

16(6)

16(6)

Testes 2 or more, tandem 23

Testis 1. Parasitic on elasmobranchs (Family Monocotylidae) 17

B

c

"\,

]

Figure 19

Capsaloides comutus: (A) ventral view. Scale: 1 mm; (B) dorsal marginal spine; (C)

hamulus. Scale: 0.01 mm. Drawn from USNM 35136.
Hendrix: Monogenea of the Eastern United States 19

17(16) Haptor septate, with more than 20 loculi. Parasitic on lamniform sharks... Cathariotrema selachii (Fig. 20)

17(16) Haptor septate, with 20 or fewer loculi. Parasitic on rays (Dasyatidae) and skates (Rajidae) 18

18(17) Haptor with 18 or 20 loculi 19

18(17) Haptor with fewer than 12 loculi. 20

19(18) Haptor with 20 loculi, hamuli absent. Parasitic on skates, Raja spp Empruthotrema raiae (Fig. 21)

A 8

Figure 20

Cathariotrema selachii: (A) ventral view. Scale: 1 mm; (B) hamulus. Scale: 0.1

mm. Drawn from USNM 35674.

Figure 21

Empruthotrema raiae. ventral view. Scale:

0.3 mm. Drawn from USNM 35160.
20 NOAA Technical Report NMFS 121

19(18) Haptor with 18 loculi and 1 pair oflarge distally recurved hamuli. Parasitic on the roughtail stingray

and little skate, Raja ennacea Thaumatocotyle dasybatis (Fig. 22)

]

Figure 22

Thaumatocotyle dasybatis: (A) ventral view. Scale: 0.5 mm; (B) hamulus.

Scale: 0.1 mm. Drawn from USNM 35660.
Hendrix: Monogenea of the Eastern United States 21

20(18) Haptor with 1 central and 10 peripheral loculi, the most posterior pair with 12 muscular papillae on

dorsal surface. Parasitic on the spotted eagle ray, Aetobatis narinari PapillicotyleJloridana (Fig. 23)

20(18) Haptor with fewer than 11 loculi, papillae absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21

8

Figure 23

Papillicotylefloridana: (A) ventral view. Scale: 0.5 mm; (B) hamulus. Scale: 0.025

mm. Drawn from USNM 49447.
22 NOAA Technical Report NMFS 121

21(20) Haptor with 1 central and 7 peripheral loculi, with numerous marginal papillae on rim of haptor,

ridge sclerites or sinuous sclerotized ridge absent. Parasitic on the roughtail stingray.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Dasybatotrema dasybatis (Fig. 24)

21(20) Haptor with 1 central and 8 peripheral loculi, marginal papillae absent, ridge sclerites or sinuous

sclerotized ridge present. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 22

A B

Figure 24

Dasybatotrema dasybatis: (A) ventral view. Scale: 1 mm; (B) hamulus.

Scale: 0.1 mm. Drawn from USNM 35656.
Hendrix: Monogenea of the Eastern United States 23

22(21) Haptor with sinuous sclerotized ridge on septa. Parasitic on the roughtail stingray and spiny dogfish,

Squalus acanthias Heterocotyle minima (Fig. 25)

8

c

]

Figure 25

Heterocotyle minima: (A) ventral view. Scale: 0.2 mm; (B) penis stylet.

Scale: 0.025 mm; (C) hamulus. Scale: 0.05 mm. Drawn from USNM

35651.
24 NOAA Technical Report NMFS 121

22(21) Haptor with numerous ridge sclerites on septa and rim. Parasitic on stingrays, Dasyatis spp.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Monocotyle spp. (Fig. 26)

B

c

Figure 26

Monocotylediademalis: (A) ventral view. Scale: 0.5 mm; (B) hamulus. Scale:

0.05 mm; (C) ridge sc1erites. Drawn from USNM 49349.
Hendrix: Monogenea of the Eastern United States 25

23(16) Haptor septate; anterior adhesive area glandular with numerous ducts. Parasitic on remoras

(Echeneidae) Dionchusspp. (Fig. 27)

23(16) Haptor lacking septa; anterior adhesive area with few gland ducts. Parasitic on elasmobranchs.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Loimoidae) 24

Figure 27

Dionchus agassizi: (A) ventral view. Scale: 0.5 mm; (B) hamulus.

Scale: 0.05 mm. Drawn from USNM 35676.
26 NOAA Technical Report NMFS 121

24(23) Haptor with 14 muscular peduncles, each contammg a marginal hook, cuticular ridges absent;

anterior adhesive area with 4 pairs of head organs. Parasitic on the bluntnose stingray, Dasyatis say.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Loimopapillosum dasyatis (Fig. 28)

24(23) Haptor lacking peduncles, cuticular ridges present on dorsal surface; anterior adhesive area

consisting of 2 or 4 small preoral suckers in a transverse row. Parasitic on requiem sharks

(Carcharhinidae) Loimos spp. (Fig. 29)

Figure 28

Loimopapillosum dasyatis: ventral view.

Scale: 0.5 mm. Drawn from USNM

38145.

A 8

Figure 29

Loimos salpinggoides: (A) ventral view. Scale: 0.5

mm; (B) hamulus. Drawn from USNM 35675.
Hendrix: Monogenea of the Eastern United States 27

25(5) Testis 1, male copulatory apparatus unarmed. Parasitic on spiny dogfish and dusky sharks, Carcharhinus

obseurns. Microbothrium apiculatum (Fig. 30)

25(5) Testes 2 or more, male copulatory apparatus armed with spines 26

Figure 30

Microbothrium apiculatum: ventral

view. Scale: 1 mm. Drawn from

USNM 35684.
28 NOAA Technical Report NMFS 121

26(25) Testes 2, opposite, male copulatory apparatus with 2 rows of about 25 long riblike spines in the

thicker anteroventral wall and 1 row of about 30 short straight spines on the posterodorsal wall.

Parasitic on requiem sharks (Carcharhinidae). . . . . . . . . . . . . . . . . . . . . . . . .. Dermophthirius spp. (Fig. 31)

Figure 31

Dermophthirius carcharhini: ventral view. Scale: 0.5 mm. Drawn

from USNM 35687.
Hendrix: Monogenea of the Eastern United States 29

26(25) Testes several, male copulatory apparatus with a row of large simple recurved spines and several

rows of small straight spines on its ventral surface. Parasitic on lemon shark, Negaprion brevirostris.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neodennophthirius harkemai (Fig. 32)

Figure 32

Neodennophthirius harkemai: ventral view. Scale: 1 mm. Drawn

from USNM 37751.
30 NOAA Technical Report NMFS 121

27(3) Viviparous (embryo may be visible in uterus); haptor with 16 marginal hooks and 1 pair of hamuli;

eyespots absent; vitellaria poorly developed. Parasitic on killifishes (Cyprinodontidae) and sticklebacks (Gasterosteidae). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Gyrodactylidae) 28

27(3) Oviparous; haptor with 14 marginal hooks and 2 pairs of hamuli; eyespots present; vitellaria

extensive. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 30

28(27) Haptor with deep transverse bar absent, lateral winglike processes present, pitted peduncular bar

presentjust anterior to haptor. Parasitic on Fundulus spp Swingleus sp. (Fig. 33)

Figure 33

Swingleus polyclilhroides: ventral

view. Scale: 0.1 mm. Drawn from

USNM 72511.
Hendrix: Monogenea of the Eastern United States 31

28(27) Haptor with deep transverse bar present, lateral winglike processes absent, peduncular bar present

or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 29

29(28) Peduncular bar large, present on ventral surface just anterior to haptor. Parasitic on cyprinodonts.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Fundulotrema prolongis (Fig. 34)

29(28) Peduncular bar absent. Parasitic on cyprinodonts and sticklebacks. . . . . .. Gyrodactylus stephanus (Fig. 35)

30(27) Haptor with 3 transverse bars; posterior of body with spines. Parasitic on drums (Sciaenidae).

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Diplectanidae) 31

Figure 35

Gyrodactylus stephanus:

ventral view. Scale: 0.1

rom. Drawn from USNM

49332.

-4-1-1+1-------- embryo

~~~j)-------_ peduncular

bar

Figure 34

Fundulotrema prolongis: ventral view. Scale: 0.1 mm. Drawn

from USNM 49331.
32 NOAA Technical Report NMFS 121

30(27) Haptor with 0, 1, or 2 transverse bars; posterior of body lacking spines. . . . . . . . . . . . . . . . . . . . . . . . . . .. 32

31(30) Haptor with dorsal and ventral squamodisks composed of scalelike spines; rows of accessory

hooklike spines on posterior of body absent. Parasitic on spotted seatrout, Cynoscion nebulosus.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Diplectanum bilobatum (Fig. 36)

31(30) Haptor lacking squamodisks; bilateral patches or rows of accessory hooklike spines present on

posterior body. Parasitic on sciaenids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Rhamnocercus spp. (Fig. 37)

B

Figure 36

Diplectanum miobatum: (A) ventral view. Scale: 0.2 mm; (B) ventral

squamodisk. Scale: 0.05 mm. Drawn from USNM 39364.

Figure 37

Rhamnocercus bairdiella: ventral view.

Scale: 0.2 mm. Drawn from USNM

49345.
Hendrix: Monogenea of the Eastern United States 33

32(30) Haptor bilobed, with peduncles, transverse bars absent. Parasitic on Atlantic torpedo, Torpedo

nobiliana.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Amphibdella jlavolineata (Fig. 38)

32(30)

33(32)

Haptor with 1 or 2 transverse bars.

Haptor with 1 transverse bar. .

33

34

33(32) Haptor with 2 transverse bars. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 35

B

Figure 38

Amphibdella flavolineata: (A) ventral view. Scale: 1 mm; (B) hamulus.

Drawn from USNM 35159.
34

34(33)

NOAA Technical Report NMFS 121

Haptor broadly bilobed; eyespots present. Parasitic on smooth flounder, Pleuronectes putnami.

Protancyroeephaloides lietpsettae (Fig. 39)

B

c

o

]

]

Figure 39

Protancyrocephaloides liopsettae: (A) ventral view. Scale: 0.5 mm; (B) transverse bar; (C) ventral hamulus; (D) dorsal hamulus. B-D scales: 0.05 mm.

Drawn from USNM 74609.
Hendrix: Monogenea of the Eastern United States 35

34(33) Haptor entire; eyespots absent. Parasitic on spiny dogfish and Atlantic torpedo.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Amphibdelloides maccallumi (Fig. 40)

35(33) Haptor with radially arranged row of troughlike sclerotized supporting structures. Parasitic on

windowpane, Scophthalmus aquosus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Bothitrema bothi (Fig. 41)

35(33) Haptor lacking radially arranged sclerotized supporting structures. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 36

Figure 40

Amphibdelloides maccallumi: (A) ventral view. Scale: 0.5 mm; (B)

ventral bar; (C) ventral hamulus; (D) dorsal hamulus. B-D

scales: 0.05 mm. Drawn from USNM 35700.

Figure 41

Bothitrema bothi: ventral view.

Scale: 0.5 mm. Drawn from

specimens collected from

windowpane, Scophthalmus

aquosus, off NewJersey.

]

]

(((f

c

B~

I I

o

b

·.I·::·:TlIII~~

t

~.:~

~~

&i'

::.

... .'.. ;&.

::.~ "'. ::)j.

~#,I ;e.

~.

~~ii ...•.... ........ :::/ . ,.. .

.'-"

i::~'

..\1.= .:.t .. .... ~::\... \.. ~. . . ..... -:..": .:. :;:~..: .: .....•~. :: .' ., .:..;", -. .'... '. ". " ·~·.i:\I~ 'fl'. Of. ':1 :': "~

A
36 NOAA Technical Report NMFS 121

36(35) Intestinal ceca not confluent posteriorly; body much broader than haptor; male copulatory apparatus contains a thin fishhook-shaped sc1erotized penis with a thin elongate V-shaped accessory piece.

Parasitic on Atlantic needlefish, Strongylura marina. . . . . . . . . . . . . . . . Ancyrocephalus parvus (Fig. 42)

36(35) Intestinal ceca confluent posteriorly; body about as broad as haptor. 37

Figure 42

Ancyrocephalus parous: (A) ventral view. Scale: 0.2 mm; (B) hamuli and transverse bars. Scale:

0.025 mm. Drawn from USNM 8143.
Hendrix: Monogenea of the Eastern United States 37

37(36) Haptor dorsal transverse bar with knobbed lateral extremities; penis sclerotized, tubular, straight,

with spiral thread and V-shaped accessory piece. Parasitic on white perch, Marone americana.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Onchocleidus nactus (Fig. 43)

37(36) Haptor dorsal transverse bar without lateral knobs; penis tube curved, without spiral thread. . . . . . . . .. 38

E

o

c

A B

Figure 43

Onchocleidus nactus: (A) ventral view. Scale: 0.05 mm; (B) dorsal hamulus; (C) ventral hamulus;

(D) dorsal bar; (E) ventral bar. B-D scales: 0.02 mm. Drawn from USNM 73744.
38 NOAA Technical Report NMFS 121

38(37) Haptor ventral transverse bar bifid laterally, lacking prominent sculpturing. Parasitic on Atlantic

spadefish, Chaetodipterusfaber Pseudohaliotrema longiphallus (Fig. 44)

A B

c

o

Figure 44

Pseudohaliotrema longiphallus: (A) ventral view. Scale: 0.2 mm; (B) ventral hamulus; (C) ventral transverse bar; (D) dorsal transverse bar. B-D

scales: 0.025 mm. Drawn from USNM 36959.
Hendrix: Monogenea of the Eastern United States 39

38(37) Haptor ventral transverse bar rounded laterally, with prominent medial sculpturing. Parasitic on

striped mullet, Mugil cephalus. Haliotrema vanbenedeni (Fig. 45)

39(1) Haptor with 4 or more clamps, posterior haptoral appendix present or absent; anterior adhesive

area a pair of buccal suckers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 44

39(1) Haptor with 3 pairs ofsuckers, each having a curved hooklike sclerite; posterior haptoral appendix

present; anterior adhesive area an oral sucker or shallow muscular grooves. . . . . . . . . . . . . . . . . . . . . . .. 40

A

B

Figure 45

Haliotrema vanbenedeni: (A) dorsal view. Scale: 0.2 mm; (B) hamuli and transverse bars. Scale: 0.05

mm. Drawn from USNM 49342.
40 NOAA Technical Report NMFS 121

40(39) Haptoral appendix armed with 3 pairs of sclerites and 1 pair of small suckers; mouth flanked by 1

pair of ventral muscular depressions; 2 pairs of eyespots present. Parasitic on shortnose sturgeon,

Acipenser brevirostrum Diclybothrium annatum (Fig. 46)

40(39) Haptoral appendix with 1 pair of small hamuli and 1 pair of distinctly muscular suckers; anterior

adhesive area an oral sucker; eyespots absent. Parasitic on elasmobranchs.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Hexabothriidae) 41

8

\

\

Figure 46

Diclybothrium armatum: (A) ventral view. Scale: 2 mm; (B) haptoral appendix.

Scale: 0.2 mm. Drawn from USNM 73136.
Hendrix: Monogenea of the Eastern United States 41

41(40) Male copulatory complex armed with small curved spines; eggs with filamentous appendage at

abopercular pole. Parasitic on smooth dogfish, Mustelus canis.. . . . . . . . . . .. Hexabothrium musteli (Fig. 47)

41(40) Male copulatory complex unarmed; eggs with or without 2 polar appendages 42

B

c

Figure 47

Hexabothrium musteli: (A) ventral view. Scale: 0.5 mm; (B) cirrus. Scale: 0.05 mm; (C) sucker

sclerite. Scale: 0.1 mm. Drawn from USNM 8132.
42 NOAA Technical Report NMFS 121

42(41) Vaginae merge to form a single duct before entering vitelline reseIVoir; ovary proximally lobate,

descending and ascending branches sinuous; eggs with ridges, lacking elongate polar appendages.

Parasitic on the barndoor skate, Raja laevis Rajonchocotyle laevis (Fig. 48)

42(41) Vaginae do not merge, enter transverse vitelline duct separately; ovary proximally branched; eggs

lacking ridges, with 2 elongate polar appendages. Parasitic on sharks or spiny dogfish. . . . . . . . . . . . 43

A 8

c

]

Figure 48

Rajonchocotyle laevis: (A) ventral view. Scale: 2 mm; (B) sclerite. Scale: 0.5 mm; (C) egg. Scale 0.1 mm. Drawn from

USNM 36721.
Hendrix: Monogenea of the Eastern United States 43

43(42) Descending branch of ovary straight; ootype smooth. Parasitic on requiem sharks (Carcharhinidae)

and hammerhead sharks (Sphyrnidae) Erpocotylespp. (Fig. 49)

8

Figure 49

Erpocotyle sphymae: (A) ventral view. Scale: 1 mm; (B) sucker with sclerite. Scale: 0.2 mm. Drawn

from USNM 8136.
44 NOAA Technical Report NMFS 121

43(42) Descending branch of ovary sinuous; ootype with longitudinal row of large cells giving ribbed

appearance. Parasitic on spiny dogfish.. . . . . . . . . . . . . . . . . . . . . . Squaloochocotyle spp. (Fig. 50)

A

]

Figure 50

Squalonchocotyle squali: (A) ventral view. Scale: 1 mm; (B) sclerite. Scale: 0.1 mm.

Drawn from USNM 8133.
44(39)

Hendrix: Monogenea of the Eastern United States 45

Haptor with 4 clamps, terminal lappet broad. Parasitic on jacks (Caranx spp.).

Protomicrocotyle mirahilis (Fig. 51)

44(39)

45(44)

Haptor with more than 4 clamps.

Haptor with 5 to 8 clamps .

45

46

45(44) Haptor with more than 8 clamps 63

o o •

•• 0

.0 . .. e••••

.. .. o

Figure 51

Protomicrocotyle mirabilis: ventral view. Scale:

0.5 mm. Drawn from USNM 37110.
46 NOAA Technical Report NMFS 121

46(45) Clamps with a patch of approximately 30 spines in the outer anterior quadrant, 8 sclerites on each

asymmetric clamp. Parasitic on pollock, Pollachius virens Diclidophora denticulata (Fig. 52)

46(45) Clamps without spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 47

47(46) Clamps approximately equal in size, with muscular sucker in one quadrant 48

47(46) Clamps unequal in size, without muscular sucker. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Figure 52

Diclidophora denticulata: (A) ventral view. Scale: 2 mm; (B) clamp. Scale: 0.4 mm. Drawn from

USNM 63394.
Hendrix: Monogenea of the Eastern United States 47

48(47) Body constricted to form prehaptoral peduncle; vitelline follicles absent in peduncle. Parasitic on

summer flounder, Paralichthys dentatus, and seatrout, Cynoscion spp Neoheterobothrium spp. (Fig. 53)

48(47) Body not constricted to form prehaptoral peduncle; vitelline follicles extend to or into haptor.. . . . . .. 49

Figure 53

Neoheterobothrium affine: (A) ventral view. Scale: 2 mm; (B) clamp. Scale: 0.2 mm. Drawn from USNM 4676.
48 NOAA Technical Report NMFS 121

49(48) Haptor with 3 pairs of pedunculate clamps and 1 pair ofsmall sessile clamps at the posterior of the

haptoral appendix. Parasitic on silver perch, Bairdiella chrysoura, and white perch. . .. Pedocotyle spp. (Fig. 54)

49(48) Haptor with 4 pairs of pedunculate clamps; haptoral appendix and sessile clamps absent. 50

Figure 54

Pedocotyle morone: (A) ventral view. Scale: 2 mm; (B) pedunculate clamp.

Scale: 0.2 mm. Drawn from USNM 35594.
Hendrix: Monogenea of the Eastern United States 49

50(49) Testes postovarian; seminal receptacle preovarian; vitelline follicles extend to but do not enter

haptor. Parasitic on pigfish, Orthopristis chrysoptera, and northern kingfish, Menticirrhus saxatilis.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Choricotyle spp. (Fig. 55)

Figure 55

Choricotyle aspinachorda, ventral view. Scale: 2

mm. Drawn from USNM 38151.
50 NOAA Technical Report NMFS 121

50(49) Testes and seminal receptacle both postovarian; vitelline follicles extend into the haptor. Parasitic

on searobins, Prionotus spp Orbocotyle prionoti (Fig. 56)

51(47) Clamps pedunculate, with 8 sclerites . 52

51(47) Clamps pedunculate or sessile, fewer than 8 sclerites 53

A

•••

....

..I.....I~tt.,J-• . ·.t~ ..:.·'1 ••'• • • ••,. ." l~' • ...'I,; .:.~ \~.', ••~4' :"I,'j::.' ::.: '/-,', ~::.,.1·,,: '. . ~. " '.:'J.".:.: :':,j-:" ..: . :..... ,.~,.~ ~:-..:(:~; .. ..~.'\.:~.. ..::~r~~·· ••.~,.~... .'7.,-,.•.•• ::.:.~l..'· :::'4:":{. ,',.., ,- : ::... ..: . •••• • 1, ',_. •

.: t.·:. . Q "r:~~" .

", . - . .. . . . ':':::" .; :'~:" :::~ ':'.:: :'\',:::

::......:.;.: .:1::':. ...:=.. :.:.~<:.>,: :::;••••: .'. .... ... . ." '-. '..' :.:. .'., '.

',':.'::: .:::::.

:.::..... "':.~::

.:.:..... :.:'.~ .:.::..•. ....', .......,'... '..:..

·:~1!:.:,i!:f~~~J ..: .'. ';:" ..... ·i..···.::.:::•• :• .. ......... . ..:::~~~.:.::~:: . ....... ...: .:. ...:..:.:: . ... . .. ... . ... ..

B

Figure 56

Orbocotyle prionoti: (A) ventral view. Scale: 0.5 mm; (B) pedunculate clamp. Scale: 0.1 mm. Drawn from

specimens collected from northern searobins, Prionotus carolinus, in the New York Bight.
Hendrix: Monogenea of the Eastern United States 51

52(51) Clamps progressively larger towards posterior of haptor. Parasitic on hakes (Gadidae) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diclidophoroides maccallumi (Fig. 57)

:1.~:. -.1 .. · ,-. .t~· .. .' .,,: . .\ \ . •el'. ' • .,'.'" . ~\ .. '. ':e. '~I~'" ,. I. • .,:•.. ' •• .':~'~ '. .', ... ,"~:~ ',-: .,".", '.. ." '. . . ••1. '.1 ": •••••• • • ' •• " •• , • I ••

.. ..." . · ". . . . . .'... : , ' , ... .-.' ..' I • .' ••: .' ·'.' '. " .' .. ... • '., 0 . 6) • ' ••

"~' ..0 0 . 0 ..' '. I. I, • •• 0 "·oOoC::·· • • •• ' GO .. 0:' I. • .' •• CD'" .:...00 0 0 ..... ". • 0 CJ 0'.. . ., ·',·0 0 o' .. I • "•• • 0 O. e. • ,', 0·;·.·· • ·.0 0 • ····OOO·~·· .' " . 0 ••• ' :.' ',0 G.· •• : . • •• : .'.:.' • •• 1 ••

.1. . .. " . ..

Figure 57

Diclidophoroides maccallumi: ventral view. Scale:

2 mm. Drawn from USNM 35106.
52 NOAA Technical Report NMFS 121

52(51) Clamps progressively smaller towards posterior of haptor. Parasitic on Atlantic croaker, Micropogonias

undulatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Absonifibula bychowskyi (Fig. 58)

53(51) Suckerlike muscular clamps with 3 sclerites, the middle one X-shaped. Parasitic on scombrids.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Fatnily Hexostomatidae) 54

53(51) Clamps with more than 3 sclerites, none X-shaped 55

A

..... • 0·

o

'..

• 0

..

..

00 •

o 0

'. c

Figure 58

Absonifibula bychowskyi: (A) ventral view. Scale: 0.3 mm; (B) clamp; (C) cirrus spines.

Drawn from USNM 73245.
Hendrix: Monogenea of the Eastern United States 53

54(53) Haptor with 4 pairs of sessile clamps in 2 longitudinal rows, the posterior pair slightly smaller than

the others. Parasitic on little tunny. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Neohexostoma euthynni (Fig. 59)

Figure 59

Neohexostoma euthynni: (A) ventral view. Scale: 1 mm; (B) sessile clamp.

Scale: 0.2 mm. Drawn from USNM 74151.
54 NOAA Technical Report NMFS 121

54(53) Haptor with 4 pairs ofsessile clamps in a transverse row along posterior margin, median pair about

one-half the size of the others. Parasitic on Atlantic bonito, Sarda sarda HexostQ11W lintoni (Fig. 60)

55(53) Clamps sessile, anterior pair much larger than posterior 3 pairs, terminal lappet with 1 pair of

hamuli and 2 pairs of marginal hooks of which 1 pair is enlarged. Parasitic on hakes (Merluccius

spp.) Anthocotyle merluccii (Fig. 61)

55(53) Clamps sessile, anterior pair not larger than posterior 3 pairs " 56

B

Figure 60

Hexostoma lintoni: ventral view. Scale: 2 mm.

Drawn from USNM 6676.

Figure 61

Anthocotylemerlurcii: (A) ventral view. Scale: 2 mm; (B) large clamp. Scale: 1 mm.

Drawn from USNM 35607.
Hendrix: Monogenea of the Eastern United States 55

56(55) Haptor divided by a deep cleft into 2 narrow lobes, each lobe with 4 clamps; 1 pair ofsmall hamuli

at the base of the cleft. Parasitic on Florida pompano, Trachinotus carolinus.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bicotylophora trachinoti (Fig. 62)

56(55) Haptor not divided into 2 narrow lobes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 57

57(56) Clamps approximately spherical, with 2 inverted V-shaped sclerites forming the anterior and

posterior edges ofthe clamp opening; hamuli at the posterior margin of haptor; genital corona with

2 types of hooks. (Family Mazocraeidae) 58

Figure 62

Bicotylophora trachinoti: ventral view. Scale: 1 mm. Drawn

from USNM 35611.
56 NOAA Technical Report NMFS 121

57(56) Clamps elongated and usually modified in shape, V-shaped sclerites separated medially; 2 pairs of

marginal hooks present at posterior margin of haptor; genital corona with a circle of curved hooks.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Macrovalvitrematidae) 61

58(57) Haptor with 4 clamps on one side and 1 small clamp on the other. Parasitic on chub mackerel,

Scomberjaponicus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Grubea cochlear (Fig. 63)

58(57) Haptor with 4 clamps on each side. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 59

A

...

...-.'

o •

00 •

o o.••_

·.. ·.

o • ·.

o ·0 .

·.'

·

o'·-•- ....

·. ·'. :..

·.

B

Figure 63

Grubea cochlear. (A) ventral view. Scale: 2 mm; (B) clamp. Scale: 0.2 mm. Drawn from

USNM 8160.
Hendrix: Monogenea of the Eastern United States 57

59(58) Clamps on short, well-separated peduncules along the lateral margins of the body distributed

anteriorly to the level of the gonads. Parasitic on herrings (Clupeidae). Mazocraeoides spp. (Fig. 64)

59(58) Clamps confined to a distinct haptor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 60

Figure 64

Mazocraeoides georgei: ventral view. Scale: 0.5 mm.

Drawn from USNM 35627.
58 NOAA Technical Report NMFS 121

60(59) Clamps pedunculate; haptoral lappet divided, with a pair of hamuli; vitellaria extend into haptor.

Parasitic on Atlantic menhaden, Brevoortia tyrannus. . . . . . . . . . . . . . . . . . .. aupeocotyle brevoortia (Fig. 65)

c

Figure 65

Clupeocotyle brevoortia: (A) ventral view. Scale: 1 mm; (B) clamp. Scale: 0.05 mm; (C) haptoral

lappets. Scale: 0.1 mm. Drawn from USNM 37492.
Hendrix: Monogenea of the Eastern United States 59

60(59) Clamps sessile; haptoral lappet not divided, with a pair of hamuli; vitellaria extend to anterior

margin of haptor. Parasitic on Atlantic mackerel, Scomber scombrus. . . . . . . . . . . . .. Kuhnia scombri (Fig. 66)

61(57) Clamps dissimilar in shape, anterior 3 pairs of clamps an elongate "firetong" shape, posterior pair

oblong; lateral margin of body behind the genital corona pleated; testis single. Parasitic on silver

perch Hargisia bairdiella (Fig. 67)

61(57) All clamps similar in shape; lateral body pleats absent; testes numerous. . . . . . . . . . . . . . . . . . . . . . . . . .. 62

Figure 66

Kuhnia scombri: (A) ventral view. Scale: 2 mm; (B) clamp. Scale:

0.05 mm. Drawn from USNM 35620.

Figure 67

H argisia bairdiella: ven tral

view. Scale: 0.1 mm. After

Hargis, 1956a.

A 8
60 NOAA Technical Report NMFS 121

62(61) Clamp sclerites with toothlike serrations, anterior pair of clamps largest; genital corona with a circle

of 6 to 7 small curved hooks. Parasitic on pigfish. . . . . . . . . . . . . . . . . . . . . . . .. Pseudotagia cupida (Fig. 68)

62(61) Clamps sclerites lack toothlike serrations, posterior pair of clamps largest; genital corona with 10 to

12 curved hooks. Parasitic on Atlantic croaker. Macrovalvitrematoides micropogoni (Fig. 69)

63(45) Haptor elongate, clamps in lateral rows on a distinct haptor or on side of body. . . . . . . . . . . . . . . . . . . .. 67

63(45) Haptor truncate or fishtail shaped, clamps in a row along posterior margin of haptor. 64

8

Figure 68

Pseudotagia cupida: ventral

view. Scale: 0.3 mm. Drawn

after Hargis, 1956a.

Figure 69

Macrovalvitrematoides micropogoni: (A) ventral view with clamps in lateral view. Scale: 0.2 mm;

(B) clamp. Scale: 0.05 mm. Drawn from specimens collected from Atlantic croaker, Micropogonias

undulatus, off Beaufort, North Carolina.
Hendrix: Monogenea of the Eastern United States 61

64(63) Haptor fishtail shaped, clamps of two different shapes, elongate "firetong" and ovoid microcotylid,

on opposite lobes of haptor, hamuli absent. Parasitic on Florida pompano.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. (Family Pyragraphoridae) Pyragraphorus pyragraphorus (Fig. 70)

64(63) Haptor truncate, clamps of one shape along posterior margin of haptor, hamuli present 65

Figure 70

Pyragraphorus pyragraphorus: (A) ventral view. Scale: 1 mm; (B) clamps. Scale: 0.1 mm. Drawn

from USNM 36559.
62 NOAA Technical Report NMFS 121

65(64) Haptor with approximately 37 clamps with accessory sclerites and multiple ribs (gastrocotylid type),

hamuli at terminus of haptor; genital atrium armed with a circle of 12 spines. Parasitic on Spanish

mackerels, Scomberomorus maculatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Pseudaxine mexicana (Fig. 71)

65(64) Clamps lacking accessory sclerites and ribs (microcotylid type), hamuli at midpoint in clamp row;

genital atrium armed or unarmed (Family Axinidae) 66

8

]

Figure 7I

Pseudaxine mexicana: (A) ventral view. Scale: 0.5 mm; (B) genital atrium.

Scale: 0.025 mm. Drawn from USNM 9167.
Hendrix: Monogenea of the Eastern United States 63

66(65) Haptor with approximately 46 clamps; genital atrium armed with 2 rows of lateral and 1 row of

medial spines; male copulatory organ armed with circle of 12 spines; ovary V-shaped. Parasitic on

silverstripe halfbeaks, Hyporhamphus unifasciatus. . . . . . . . . . . . . . . . . . . . . . . . . .. Axine hyporhampi (Fig. 72)

B

Figure 72

Axine hyporhampi: (A) ventral view. Scale: 0.5 mm; (B) genital atrium and cirrus spines. Drawn from USNM

36550.
64 NOAA Technical Report NMFS 121

66(65) Haptor with 42 to 82 clamps; genital atrium and male copulatory organ unarmed; ovary J-shaped.

Parasitic on Atlantic needlefish Nudaciraxine gracilis (Fig. 73)

67(63) Clamps with accessory sclerites and multiple ribs (gastrocotylid type). 68

67(63) Clamps lack accessory sclerites and multiple ribs (microcotylid type). 71

68(67) Clamps confined to haptor proper. . 69

68(67) Clamps along lateral body margins.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

69(68) Haptor with about 80 somewhat asymmetric clamps in two equal rows; vitellaria not extending into

haptor; male copulatory organ with numerous short spines. Parasitic on bluefish, Pomatomus saltatrix,

striped bass, Marone saxatilis, and mackerels, Scomberomorus spp Gotocotyla acanthophallus (Fig. 74)

Figure 74

Gotocotyla acanthophallus: (A) ventral view. Scale: I mm; (B) hamulus. Scale: 0.05 mm; (C) cirrus.

Drawn from USNM 36558.

~J

c

B

Figure 73

Nudaciraxine gracilis: ventral view.

Scale: 1 mm. Drawn from USNM

37723.
Hendrix: Monogenea of the Eastern United States 65

69(68) Haptor with about 135 clamps in two unequal rows, a short row of 50 and a long row of 85 opposite;

vitellaria extending into haptor; male copulatory organ with numerous elongate spines. Parasitic on

mackerels. Scomberocotyle scomberomori (Fig. 75)

B

Figure 75

Scomberocotyle scomberomori: (A) ventral view. Scale: 1 mm; (B) cirrus and genital atrium. Scale:

0.1 mm. Drawn from USNM 37494.
66 NOAA Technical Report NMFS 121

70(68) Haptor footlike with about 40 clamps; testes few, large; vitellaria confined to ventral half of haptoral

region. Parasitic on mackerels. Thoracocotyle crocea (Fig. 76)

B

Figure 76

Thoracocotyle crocea: (A) ventral view. Scale: 1 mm; (B) clamp. Scale: 0.05 mm. Drawn from

USNM 35588.
Hendrix: Monogenea of the Eastern United States 67

70(68) Haptor not footlike, with about 200 clamps in roughly equal rows; testes numerous, small; vitellaria

not so confined. Parasitic on wahoo, Acanthocybium solandri. Neothoracocotyle acanthocybii (Fig. 77)

71(67) Clamps in distinctly unequal rows (Family Heteraxinidae) 72

A B

Figure 77

Neothoracocotyle acanthocybii: (A) ventral view. Scale: 2 mm; (B) genital atrium. Scale: 0.1 mm. Drawn from USNM 38132.
68 NOAA Technical Report NMFS 121

71(67) Clamps in approximately equal rows (Family Microcotylidae) 73

72(71) Clamps asymmetrical, unequal in size and shape, clamps in the long row larger, about 36-55 in

number, clamps of the short row 11-17 in number; genital atrium and male copulatory organ

armed with more than 30 spines. Parasitic on jacks, Caranx spp. . . . . . . . . . . . . . .. Cemocotyle spp. (Fig. 78)

c

8

Figure 78

Cemocotyle noveboracensis: (A) ventral view. Scale: 2 mm; (B) clamp. Scale: 0.05 mm; (C)

genital atrium and cirrus. Scale: 0.1 mm. Drawn from USNM 37738.
Hendrix: Monogenea of the Eastern United States 69

72(71) Clamps symmetric and approximately equal in size and shape, with 25-35 in the long row and 18-21

in the short row; genital atrium armed with fewer than 20 spines; male copulatory organ unarmed.

Parasitic on spot, Leiostomus xanthurus Heteraxinoides xanthophilis (Fig. 79)

73(71) Genital atrium unarmed. 74

73(71) Genital atrium armed with spines 75

A

Figure 79

Heteraxinoides xanthophilis: (A) ventral view. Scale: 0.5 mm; (B) clamp.

Scale: 0.025 mm; (C) genital atrium spines. Scale: 0.05 mm. Drawn from

USNM 38158.
70 NOAA Technical Report NMFS 121

74(73) Clamps in double rows on each side of haptor; male copulatory organ unarmed. Parasitic on black

drum, Pogonias cromis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Pauciconfibula pogoniae (Fig. 80)

74(73) Clamps in a single row on each side of haptor; male copulatory organ armed with spines. Parasitic

on striped bass. Gamacallum macroura (Fig. 81)

8

c

A

.. .. .

·.

.'.' .. . ..•. .. " ·.. '.. .. " .... ·.'. · . · . · . .' .'. . . '. ' : ...' ' . .' '... ., .' . .. .. .... . '.. '..

.'" •••'•• ··rt~~2~J!J'IIV/·J;l •• • 't'

.. .. .., . :. ." ...... . .. ..

....':' .. '

.. . .' "."c..:;l:..;:,.~ti2~

••• :•• '.;'6·.~.~~~~,

" . ," " .'..... ~: .. .. .,' .' :'. ·'... • ..'. \Ah(,)"y,'lj:;!

. . ·. ., .

·.. .'

/ .'. '..:. ,':: " • •••..<:...,.!,j""'~ .' '. .. · ' :,.:: .' '

·.. .. ..' " t,

·.·.. ·.

Figure 81

Gamacallum macroura: (A) ventral view. Scale: 2 mm; (B) clamp; (C) egg. Drawn

from USNM 36525.

Figure 80

Pauciconfibula pogoniae: ventral view. Scale:

2 mm. Drawn from USNM 35090.
Hendrix: Monogenea of the Eastern United States 71

75(74) Genital atrium with 6 to 7 lateral muscular suckerlike pockets, armed with 3 rows ofspines. Parasitic

on seatrout, Cynoscion spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Cynoscionicola spp. (Fig. 82)

75(74) Genital atrium lacking lateral pockets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 76

A B

c

o

Figure 82

Cynoscionicola pseudohet(ffacantha: (A) ventral view. Scale: 1 mm; (B) clamp;

(C) egg; (D) genital atrium complex. Scale: 0.1 mm. Drawn from USNM

38251.
72 NOAA Technical Report NMFS 121

76(75) Vitellaria extend to posterior end of body proper; genital atrium usually armed with a circle or patch

ofspines. Parasitic on perciform fishes. . Microcotyle spp. (Fig. 83)

B

Figure 83

Microcotyle pornatorni: (A) ventral view. Scale: 0.5 mm; (8) clamp; (C) genital atrium

spines. Scale: 0.05 mm. Drawn from specimens collected from bluefish, Pornatornus

saltatrix, off Beaufort, North Carolina.
Hendrix: Monogenea of the Eastern United States 73

76(75) Vitellaria do not reach end of body; genital atrium armed with 15 to 20 pairs ofspines in 3 groups.

Parasitic on striped mullet Metamicrocotyla macracantha (Fig. 84)

A

o 0

00

00 .0.

0 0 0

....;

.0

0 0

.0o

00

.. ;" O.

o

B

c

Figure 84

Metamicrocotyla macracantha: (A) ventral view. Scale: 2 mm; (B) clamp; (C) genital atrium spines.

Drawn from USNM 38253.
74 NOAA Technical Report NMFS 121

Annotated Systematic List _

Numerous authors have contributed to reports of the

monogeneans, as well as the one udonellid discussed

below, from fish hosts of the estuarine and marine

coastal waters from the U.S.-Canada border to Beaufort, North Carolina. All known records for this region

are given for each of the 108 species. In addition,

annotated references to their biology and morphology

are included. No attempt is made here to give individual global records for those species with cosmopolitan distributions or to cite reports of their occurrence

in distant geographic locations. The taxonomic system

followed in this manual is generally a modification of

the scheme of Llewellyn (1970) as used by BeverleyBurton (1984) for Canadian monogeneans. Superfamilies, families, genera, and species are arranged alphabetically within each of the orders of the class. Several

general references that may prove useful in the study of

Monogenea include Hargis (1957a), Koratha (1955a),

Lebedev (1979), Margolis and Arthur (1979), Price

(1939b), Rohde (1978b, 1979, 1981, and 1986), and

Schell (1985). For older literature and synonyms, see

Baer and Euzet (1961), Hargis et al. (1969, 1970, 1971,

1972, and 1982), Sproston (1946), orYamaguti (1963).

Scientific as well as common names of fishes are included when known and follow Special Publication No. 20

of the American Fisheries Society (Robins et aI., 1991).

Class UDONELLIDEA Ivanov, 1952

Order UDONELLIDA Ivanov and Mamkaev, 1973

Family UDONELLIDAE Taschenberg, 1879

Udonella caligorumJohnston, 1835

Synonyms: Nitzschia papillosa Linton, 1898; Lintonia

papillosa (Linton, 1898) Monticelli, 1904; Calinella

myliobati Guberlet, 1936; Udonella socialis Linton, 1910;

and several others.

Records: Atlantic cod, Gadus morhua, Woods Hole

(Linton, 1898), Georges Bank (Linton, 1940); lumpfish,

Cyclopterus lumpus, and little tunny, Euthynnus alletteratus,

off Chesapeake Bay (Zwerner and Lawler, 1972).

Remarks: Udonella caligorum is found as a hyperparasite on copepods, such as Caligus spp., themselves parasitic on the gills of a variety of fish hosts. For example,

this species has been reported from the parasitic copepod Caligus praetextus in the mouth and skin of red

drums, Sciaenops ocellatus, in Mississippi Sound and at

Palacois, Texas (Overstreet, 1983a and 1983b), and on

the isopod Livoneca vulgaris off San Francisco, California (Crane, 1972). Udonella caligorum is distributed along

both the Atlantic and Pacific coasts of North and South

America as well as off the coasts of Europe, Australia,

and New Zealand. Halton and Jennings (1965) provided some information on its feeding and digestive

processes; Kabata (1973) and Villalba (1985) gave the

distribution and probable feeding habits on copepod

hosts in Scottish and Chilean waters, respectively. Schell

(1972) described early development.Justine et al. (1985)

briefly described sperm ultrastructure; Rohde et al.

(1989) examined the fine structure ofsperm, sense receptors, flame bulbs, and tegument as a basis for phylogenetic

position. Price (1938b) and Nichols (1975) described the

morphology and possible taxonomic relationships. Udonefla

caligorum is included in this manual because of its

cooccurrence with monogeneans on fish gills.

Class MONOGENEA Carus, 1863

Order MICROBOTHRIIDEA Lebedev, 1988

The microbothriids as well as the "monocotylids" (families Monocotylidae and Loimoidae) are assigned to

orders following the scheme of Lebedev (1988) and

Boeger and Kritsky (1993). Earlier, Llewellyn (1970)

and Beverley-Burton (1984) mentioned the uncertain

relationships of the families with other monogeneans.

Lawler (1981) gave an extensive review of the zoogeography and host specificity of members of the superfamily Capsaloidea using Price's (1936) scheme, which includes the present orders Microbothriidea, Monocotylidea, and portions of Dactylogyrida.

Family MICROBOTHRIIDAE Price, 1936

Dermophthirius carcharhini MacCallum, 1926a

Synonyms: None.

Records: On the skin and nasal cavities of what is

probably the dusky shark, Carcharhinus obscurus (reported by MacCallum [1926a] as Careharhinus commersonii), Woods Hole, Massachusetts. The exact species

represented by this host name is questionable (Lawler,

1981).

Remarks: Also reported from the skin of the blacktip

shark, Carcharhinus limbatus, off Grand Isle, Louisiana,

and Mississippi (Thatcher, 1959). Benz (1987) suggested

that the specimens identified by Thatcher (1959) may

be D. penneri. Cheung and Ruggieri (1983) believed

that specimens reported as D. carcharhini from sharks

off Senegal by Euzet and Maillard (1967) may be D.

nigrellii. It was redescribed by Price (1938b), but Cheung

and Ruggieri (1983) pointed out a misinterpretation of

the haptor structure by Price. Rand et al. (1986) described

its attachment to the Galapagos shark, C. galapagensis.

Dermophthirius penneri Benz, 1987

Synonyms: None.
Records: On the body surface of the blacktip shark,

NewJersey (Benz, 1987).

Remarks: Benz (1987) also reported it from the Gulf

of Mexico, off Florida.

Microbothrium apiculatum Olsson, 1869

Synonyms: Dermophagus squali MacCallum, 1926b, and

Philura orata MacCallum, 1926a.

Records: On the skin in the cloacal region of the

spiny dogfish, Squalus acanthias, Chesapeake Bay

(Zwerner and Lawler, 1972), and Woods Hole, Massachusetts (MacCallum, 1926, a and b); probably the dusky

shark, Carcharhinus obscurus (reported as Carcharhinus

commersoniz), Woods Hole (MacCallum, 1926, a and b).

Remarks: Other geographic records include Canada

(Beverley-Burton, 1984) and the eastern Atlantic. Price

(1938b) redescribed the species and (1963) reviewed

the family and gave a key to the species of Microbothrium.

Neodermophthirius harkemai Price, 1963

Synonyms: Cadenatia polytestis Euzet and Maillard,

1967.

Record: On the gills of the lemon shark, Negaprion

brevirostris, North Carolina (Price, 1963).

Remarks: Also reported off Senegal by Euzet and

Maillard (1967).

Order MONOCOTYLIDEA Lebedev, 1988

Family LOIMOIDAE Bychowsky, 1957

Loimopapillosum dasyatis Hargis, 1955e

Synonyms: None.

Record: On the gills ofthe bluntnose stingray, Dasyatis

say, Chesapeake Bay (McMahon, 1963).

Remarks: Also reported from stingrays collected at

Alligator Harbor, Florida (Hargis, 1955e).

Loimos salpinggoides MacCallum, 1917

Synonyms: None.

Records: On the gills ofthe dusky shark, Woods Hole

(MacCallum, 1917).

Remarks: Price (1938b) redescribed the species, and

Manter (1944) supplemented its description.

Loimos scoliodoni (Manter, 1938) Manter, 1944

Synonyms: Tricotyle scoliodoni Manter, 1938.

Records: On the gills ofthe Atlantic sharpnose shark,

Rhizoprionodon terraenovae, Beaufort, North Carolina

(Manter, 1938).

Remarks: Also found on sharks from Alligator Harbor, Florida (Hargis, 1955e), and Texas by Koratha

(1955b), who also redescribed it.

Hendrix: Monogenea of the Eastern United States 75

Family MONOCOTYLIDAE Taschenberg, 1879

Cathariotrema selachii (MacCallum, 1916b)Johnston and

Tiegs, 1922

Synonyms: Monocotyle selachii MacCallum, 1916b, and

Paramonocotyle selachii (MacCallum, 1916b)Johnston, 1934.

Records: In the nasal cavities of the thresher shark,

Alopias vulpinus; dusky shark; and the smooth hammerhead, Sphyrnazygaena (MacCallum, 1916b); "shark" (reported as Carcharias commersonii by Price, 1938b), all at

Woods Hole, Massachusetts.

Remarks: Redescribed by Price (1938b).

Dasybatotrema dasybatis (MacCallum, 1916b) Price,

1938b

Synonyms: Monocotyledasybatis MacCallum, 1916b, and

Monocotyloides dasybatis (MacCallum, 1916b) Johnston,

1934.

Records: On the gills ofthe roughtail stingray, Dasyatis

centroura, Woods Hole, Massachusetts (MacCallum,

1916b).

Remarks: Price (l938b) redescribed the species. Euzet

and Maillard (1967) gave a detailed description of the

haptor and revised the generic diagnosis based on specimens from Dasyatis marmorata taken off Senegal, Africa.

Young (1967) revised the generic diagnosis.

Empruthotrema raiae (MacCallum, 1916b)Johnston and

Tiegs, 1922

Synonyms: Acanthocotyle raiae MacCallum, 1916b.

Records: In the nasal fossae of the little skate, Raja

erinacea, and winter skate, R. ocellata, Woods Hole, Massachusetts (MacCallum, 1916b); gills of the clearnose

skate, R. eglanteria. Chesapeake Bay (McMahon, 1963).

Remarks: Also reported from Alligator Harbor,

Florida (Hargis, 1955e). Redcscribed by Price (1938b)

and Kearn (1976), who also described the oncomiracidium and gave an emended generic diagnosis

based on specimens from eastern Atlantic rajids.

Whittington and Kearn (1992) also emended the generic diagnosis.

Heterocotyle minima (MacCallum, 1916b) Price, 1938b

Synonyms: Monocotyle dasybatis minimus MacCallum,

1916b; Trionchus dasybatis MacCallum, 1916b; Monocotyle

minima (MacCallum, 1916b) Johnston and Tiegs, 1922;

and Monocotyloides minimus (MacCallum, 1916b)

Johnston, 1934. Heterocotyle minimus may be a synonym

of H. (Monocotyle) pastinacae Scott, 1904, a species insufficiently described at this time.

Records: On the gills of the roughtail stingray and

spiny dogfish, Woods Hole, Massachusetts (MacCallum,

1916b).

Remarks: Redescribed by Price (1938b). Young (1967)

reviewed the genus and produced a key to species.
76 NOAA Technical Report NMFS 121

Monocotyle diademalis Hargis, 1955d

Synonyms: Heterocotyloides diademalis (Hargis, 1955d)

Yamaguti, 1963.

Records: On the gills of the bluntnose stingray and

southern stingray, Dasyatis americana, Chesapeake Bay

(McMahon, 1963).

Remarks: Also known from dasyatids collected at Alligator Harbor, Florida (Hargis, 1955d). Timofeeva

(1984) emended the generic diagnosis and produced a

key to the species of Monocotyle. Measures et al. (1990)

emended the subfamily and generic diagnoses and provided a phylogenetic analysis of the genus.

Monocotyle pricei Pearse, 1949

Synonyms: Heterocotyloides pricei (Pearse, 1949)

Yamaguti, 1963.

Records: On the gills ofsouthern and bluntnose stingrays, Chesapeake Bay (McMahon, 1963). Also reported

from the gills ofsheepshead, Archosargus probatocephalus,

collected at Beaufort, North Carolina (Pearse, 1949).

Hargis (1955d) considered this an accidental or abnormal host.

Remarks: Redescribed by Hargis (1955d) from Alligator Harbor, Florida, material. Kingston et al. (1969)

described the oncomiracidium.

Papillicotyle floridana (Pratt, 1910) Young, 1967

Synonyms: Monocotyle jlmidana Pratt, 1910; Heterocotyle

jlmidana (Pratt, 1910) Price, 1938b; Heterocotyle aetobatis

Hargis, 1955d; and Alloheterocotyla aetobatisYamaguti, 1968.

Records: On the gills of the spotted eagle ray, Aetobatis

narinari, Beaufort, North Carolina (Pearse, 1949).

Remarks: Reported from the Tortugas, Florida (Pratt,

1910), the Gulf coast of Florida (Hargis, 1955d), and

Hawaii (Yamaguti, 1968). Redescribed by Price (1938b).

In addition, Young (1967) noted that this species has

10, not 8, radial septa.

Thaumatocotyle dasybatis (MacCallum, 1916b) Price,

1938b

Synonyms: Merizocotyle dasybatis MacCallum, 1916b, and

Pseudomerizocotyledasybatis (MacCallum, 1916b) Kay, 1942.

Records: In the nasal fossae (MacCallum, 1916b)

and on the gills of the roughtail stingray and the little

skate, Woods Hole, Massachusetts (Price, 1938b).

Remarks: Redescribed by Price (1938b). This may be

a synonym of T. concinna Scott, 1904 (see Brinkmann,

1940; Lawler, 1981).

Order DACTYLOGYRIDA Bychowsky, 1937

Family ACANTHOCOTYLIDAE Price, 1936

Pseudacanthocotyla verrilli (Goto, 1899) Yamaguti, 1963

Synonyms: Acanthocotyle verrilli Goto, 1899, and A.

borealis Brinkmann, 1940.

Records: On the skin of the little skate, Maine

(Manter, 1925 and 1926), and Cape Cod, Massachusetts (Goto, 1899); and the thorny skate, R. radiata,

Cape Cod (Goto, 1899).

Remarks: Also reported from Canada, Greenland,

Iceland, Norway, and the Bering Sea. Redescribed by

Price (1938b). Using histochemical techniques, Joffe

and Kotikova (1988) demonstrated cholinesterase in

the nervous system.

Family AMPHIBDELLIDAE Bychowsky, 1957

Amphibdella flavolineata MacCallum, 1916b

Synonyms: None.

Records: On the primary gill mucosa of the Atlantic

torpedo, Torpedo nobiliana, Woods Hole, Massachusetts

(MacCallum, 1916b), and Menemsha Bight near Woods

Hole (Linton, 1940); "stingray,"Woods Hole (Price, 1937).

Remarks: Redescribed by Price (1937). This species

was also reported from the Irish Sea, off southern England, and the Mediterranean. A detailed account of its

biology, morphology, and taxonomy was given by

Llewellyn (1960). Juveniles are found in the heart ventricle of the host, and Lyons (1971) described epidermal ultrastructure of these juveniles as well as the gilldwelling adults.

Amphibdelloides maccallumi Uohnston and Tiegs, 1922)

Price, 1937

Synonyms: A mphibdelloides torpedinis Parona and

Perugia, 1890b, not Chatin, 1874; A. torpedinis MacCallum, 1916b; and Amphibdella maccallumiJohnston and

Tiegs, 1922.

Records: On the secondary gill lamellae of the spiny

dogfish, Woods Hole, Massachusetts (Price, 1937); and

Atlantic torpedo, Woods Hole (MacCallum, 1916b;

Price, 1937).

Remarks: Records of the species beyond the range of

this manual include the Irish Sea, Irish Atlantic Slope,

off Plymouth, England, Mediterranean Sea, off California, and New Zealand. Redescribed by Alexander (1954),

Llewellyn (1960), and Dillon and Hargis (1965a).

Llewellyn (1960) studied its morphology and biology

and provided a list of amphibdellids from electric rays

(Torpedinidae). According to Llewellyn (1960), the oncomiracidium described by Euzet (1957) was believed to be

that of either this species or A. vallei. Lyons (1966) provided histochemical data on hamuli and marginal hooks.

FamilyANCYROCEPHALIDAE Bychowskyand Nagibina,

1978

Ancyrocephalus parous Linton, 1940

Synonyms: None.
Records: On the gills of the Atlantic needlefish,

Strongylura marina, Woods Hole, Massachusetts (Linton,

1940), and Chesapeake Bay (Kingston et aI., 1969).

Remarks: Also reported from Alligator Harbor,

Florida (Hargis, 1955b), off Alabama by Williams and

Rogers (1972), and on redfin needlefish, S. notata, in

Biscayne Bay, Florida (Skinner, 1978). The species was

redescribed by Williams and Rogers (1972), and

Kingston et al. (1969) described the oncomiracidium.

Family BOTHITREMATIDAE Bychowsky, 1957

Bothitrema bothi (MacCallum, 1913c) Price, 1936

Synonyms: Acanthocotyle bothi MacCal1um, 1913c.

Records: On the gills ofthe windowpane, Scophthalmus

aquosus, Woods Hole, Massachusetts (MacCal1um, 1913c,

1916a, and 1917), and NewJersey (Meyers, 1978). Also

occasionally found in nares of NewJersey windowpanes

(this publication).

Remarks: Price (1937) redescribed this species.

Family CAPSALIDAE Baird, 1853

Lawler (1981) gave an extensive review of the zoogeography and host specificity of members of the superfamily Capsaloidea using the scheme of Price (1936).

Stunkard (1962) as well as Wheeler and Beverley-Burton (1987) pointed out the problems in clearly delineating genera within the subfamily Capsalinae.

Benedeniella posterocolpa (Hargis, 1955e) Yamaguti, 1963

Synonyms: Benedenia posterocolpa Hargis, 1955e.

Records: On the ventral surface skin of the cownose

ray, Rhinoptera bonasus, Chesapeake Bay (McMahon, 1963).

Remarks: Also reported from this host in Tampa Bay,

Florida (Hargis, 1955e).

Caballerocotyla manteri (Price, 1951) Price, 1960

Synonyms: Capsala manteri Price, 1951.

Records: On the gills of the little tunny, Chesapeake

Bay (Zwerner and Lawler, 1972).

Remarks: Price (1951) reported this species off the

Tortugas, Florida. Mamaev (1968) described the subspecies, C. m. affinis from South China Sea scombrids.

Price (1960), Stunkard (1962), Wagner and Carter

(1967), and Bussieras (1972) gave keys to the species.

Justine et al. (1985) described spermatozoan ultrastructure; and Justine and Mattei (1987) described spermatogenesis. Yamaguti (1968) considered Caballerocotyla

to be a subgenus of Capsala.

Capsala martinierei Bose, 1811

Synonyms: None.

Records: On the skin of ocean sunfish, Mola mola,

Hendrix: Monogenea of the Eastern United States 77

Atlantic coast from Newfoundland (Threlfall, 1967)

and Nova Scotia, Canada (Logan and Odense, 1974).

Remarks: This species has also been reported from

the eastern Atlantic Ocean, Mediterranean Sea, and

the Pacific coast of North America and Chile. Price

(1939a) redescribed the adult. Logan and Odense

(1974) studied the skin pathology caused by this worm,

and Kearn (1963) described the oncomiracidium. There

is a great deal of confusion in the literature over the

identification of this species as well as its separation

from Tricotyla molae and Tristoma coccineum. This species

has not yet been reported from this manual's range,

but it is included because of the cosmopolitan distribution ofits host and the proximity ofthe Canadian records.

Capsaloides cornutus (Verrill, 1875) Price, 1939a

Synonyms: Tristoma cornutum Verrill, 1875; Capsala

cornuta (Verrill, 1875) Johnston, 1929; and Capsaloides

cornutum (Verrill, 1875) Price, 1938a.

Records: On the gills of white marlin, Tetrapturus

albidus, Woods Hole, Massachusetts, and the Block Island region, Rhode Island (Verrill, 1875; Price, 1939a).

Remarks: Price (1960) gave a key to the species.

Capsaloides magnaspinosus Price, 1939a

Synonyms: None.

Records: In the nares of white marlin, Woods Hole,

Massachusetts (Price, 1939a).

Remarks: The description is based on three specimens collected by MacCal1um.

Entobdella bumpusii (Linton, 1900)Johnston, 1929

Synonyms: Epibdella bumpusii Linton, 1900; Epibdella

(Phylline) bumpusii (Linton, 1900) Monticelli, 1902;

Phylline bumpusii (Linton, 1900) Linstow, 1903; Phyllonella

bumpusii (Linton, 1900) MacCallum, 1927; and Entobdella

(Parepibdella) bumpusii (Linton, 1900) Johnston, 1929.

Records: On the skin and gills of the roughtail stingray, Woods Hole, Massachusetts (Linton, 1900).

Remarks: Redescribed by Price (1939a). Linton

(1908) described the genitalia and egg formation. Crane

(1972) gave a key to the species. Lyons (1966) reported

histochemical tests on hamuli and marginal hooks.

Klassen et al. (1989) reviewed and revised the genus,

giving an emended diagnosis and key to species.

Entobdella hippoglossi (Milller, 1776) Blainville, 1818

Synonyms: Phylonella hippoglossi (Muller, 1776) Coto,

1899; Epibdella (Phylline) hippoglossi (Muller, 1776)

Mon ticelli, 1902; Entobdella (Entobdella) hippoglossi

(Muller, 1776) Johnston, 1929; Epibdella bumpusii of

Canavan, 1934; Entobdella In'attstroemi Brinkmann, 1952;

E. curvunca Ronald, 1957; E. rosaceus Crane, 1972; E.

steingroeveri (Cohn, 1916) Johnston, 1929; E. squamula

(Heath, 1902) Johnston, 1929.
78 NOAA Technical Report NMFS 121

Records: On the skin ofAtlantic halibut, Hippoglossus

hippoglossus, from Woods Hole, Massachusetts, and Swans

Island, Maine (Price, 1939a).

Remarks: Also reported from this host offNova Scotia,

Greenland, Iceland, northern Europe, Alaska, the

Bering Sea, and the Barents Sea; arrowtooth flounder,

Atheresthes stomias, Bering Sea; Pacific halibut, H.

stenolepis, eastern Pacific and Bering Sea. Reported by

Meserve (1938) on the bullseye puffer, Sphoeroides annulatus. This record is considered erroneous by modern workers (Lawler, 1981). Halton andJennings (1965)

reported some aspects of nutrition, and Arme (1977)

reported on the amino acid composition. Kearn (1974a)

gave information on larval hatching and (1974b) described the oncomiracidium. Klassen et al. (1989) provided comparative morphometric data.

Nasicola klawei (Stunkard, 1962) Yamaguti, 1968

Synonyms: Tristoma sp. of Rossignol and Repelin,

1962, and Caballerocotyla klawei Stunkard, 1962.

Records: In the nasal capsules of yellowfin tuna.

Thunnus albacares, off New York (Bane, 1969).

Remarks: Additional reports from T. albacares include

Puerto Rico, Bahamas, Venezuela, Gulf of Guinea, eastern Atlantic Ocean, and the Pacific Ocean offnorthefll

Peru and Hawaii (reported as Neothnnnus macropterus by

Stunkard [1962]); blackfin tuna, T. atlanticus, Puerto

Rico; and tuna, Hawaii. Bane (1969) gave comparative

measurements of worms from the two Atlantic hosts.

Nitzschia sturionis (Abildgaard, 1794) Krl"yer, 1852

Synonyms: Phylline sturionis (Abildgaard. 1794)

Monticelli, 1908.

Records: On the gills of the Atlantic sturgeon,

Acipenser oxyrhynchus, New Brunswick, Canada (Appy

and Dadswell, 1978).

Remarks: This species is included because the record

is near the geographic range of this manual. Also, N.

superba may be a synonym of N. sturionis (Bychowsky,

1957). It has been reported from the gills of sturgeons

from northern and eastern Europe. Timofeeva (1983)

studied the nervous system. Gusev and Timofeeva (1986)

described ciliary cells and chaetotaxy of the oncomiracidium, andJoffee et al. (1987) gave a description

of the adult pharynx.

Nitzschia superba MacCallum, 1921

Synonyms: Nitzschia elegans of Verrill (1875) amI of

Linton (1898) and N. elongataofLinton (1901).

Records: On the gills and inner surface of the operculum of shortnose sturgeon, Acipenser brevirostrum, Woods

Hole, Massachusetts; and Atlantic sturgeon, Woods Hole

(Linton, 1898).

Remarks: Redescribed by Price (1939a). Bychowsky

(1957) considered this to be a synonym of N. sturionis.

However, more work is needed to clarity the validity of

this species.

Tricotyla molae (Blanchard, 1847) Guiart, 1938

Synonyms: Capsala molae (Blanchard, 1847)Johnston,

1929; Tricotyla cutanea var. mediterranea or T. c. microcotyla

ofGuiart (1938); and TristomumrudolphianumofDiesing

(1850) and of Linton (1898 and 1900).

Records: On the skin of the ocean sunfish, Woods

Hole, Massachusetts (Linton, 1898 and 1940), NewJersey (Leidy, 1890; Price, 1962c), and off Delaware (this

publication) .

Remarks: Also reported from the eastern Atlantic

Ocean and Mediterranean Sea. Because of their similarity, past workers have probably confused Tricotyla

molae with Capsala martinierei.

Tristoma coccineum Cuvier, 1817

Synonyms: None.

Records: On the gills ofsmooth hammerhead, Woods

Hole, Massachusetts; and swordfish, Xiphias gladius,

Woods Hole (Price, 1939a) and northwest Atlantic (Ill'S,

1971) .

Remarks: Iles (] 971) detailed differences between T.

wccineum and T. integrum from swordfish captured offshore along the middle Atlantic coast. This species is

alsu known from the eastern Atlantic Ocean and Mediterranean Sea.

Tristoma integrum Diesing, 1850

Synonyms: Tristomum rotundum Goto, ]894, and T.

coccineumCuvier, ofLinton (]898, 1900, 1901,and ]940).

Records: On the gills of swordfish from the northwest Atlantic Ocean (Iles, 1971) and Woods Hole, Massachusetts (Price, 1939a).

Remarks: None.

Tristomella laeuis (Verrill, 1875) Guiart, 1938

Synonyms: Tristoma laeve Verrill, 1875; Tristomum

histiophori Bell, 1891; Tristomum laeve var. armata Goto,

1899; and Tristomella laeve (Verrill, 1875)johnston, 1929.

Records: In the mouth of white marlin, Block Island,

Rhode Island (Verrill, 1875); gills of swordfish, Woods

Hole and Block Island region (Linton, 1940).

Remarks: Also reported off Puerto Rico by Dyer et al.

(1992), from the eastern and southern Atlan tic Ocean

(Pritchard, 1961), and off India on billfishes (Istiophoridae and Xiphiidae). Price (1938a) redescribed

the species from Brazilian dolphin, Coryphaena hippurus.

Tristomella lintoni (Price, 1939a) Price, 1960

Synonyms: Tristoma laeve of Lin ton (1898 and 1901)

and Capsala lintoni Price, 1939a.

Records: On the gills of skipjack tuna, Katsuwonus
pelamis, near Martha's Vineyard, Massachusetts (Price,

1939a).

Remarks: The species description is based upon a

single immature specimen.

Tristomella unchidiocotyle (Setti, 1899) Guiart, 1938

Synonyms: Tristomum onchidiocotyle Setti, 1899; Capsala

onchidiocotyle (Setti, 1898) Johnston, 1929; Capsala

onchidiocotyle(Setti, 1899) Price, 1938a; and C. maccallumi

Price, 1939a.

Records: On the gills of the little tunny, Woods Hole

region (Price, 1939a).

Remarks: Also reported from tunas in the Mediterranean and off the coast of Angola.

Family DACTYLOGYRIDAE Bychowsky, 1933

Haliotrema vanbenedeni (Parona and Perugia, 1890a)

Young, 1968

Synonyms: Tetronchus Van Benedenii Parona and

Perugia, 1890a; Ancyrocephalus vanbenedenii (Parona and

Perugia, 1890a) Johnston and Tiegs, 1922; Haplocleidus

vanbenedeni (Parona and Perugia, 1890a) Palombi, 1949;

Pseudohaliotrema mugilinus Hargis, 1955c; and Haliotrema

mugilinus (Hargis, 1955c) Yamaguti, 1963.

Records: On the gills of the striped mullet, Mugil

cephalus, Chesapeake Bay (Zwerner and Lawler, 1972).

Remarks: Also reported from mullets (Mugilidae) off

Georgia (Rawson, 1976), Biscayne Bay (Skinner, 1975)

and Alligator Harbor, Florida, Puerto Rico, and in the

North, Mediterranean, and Black Seas. Gusev (1955) redescribed the species; Mizelle and Price (1964) gave a key

to genera in the subfamily Ancyrocephalinae; and Young

(1968) revised the genus. Garcia and Williams (1985)

gave some temporal data on this species from Puerto Rico.

Onchocleidus nactus (Mayes andJohnson, 1975) Wheeler

and Beverley-Burton, 1989

Synonyms: Urocleidus nactus Mayes andJohnson, 1975,

and Pterocleidus nactus (Mayes and Johnson, 1975)

Beverley-Burton et al., 1986.

Records: On the gills ofwhite perch, Morone americana,

estuarine Abermarle Sound, North Carolina (Mayes

andJohnson, 1975), and the lower Hudson River, New

York (Liquori and Insler, 1985).

Remarks: Wheeler and Beverley-Burton (1989)

supplemented the species description. Liquori and

Insler (1985) gave data on seasonal fluctuations of 0.

nactus populations in the Hudson River.

Protancyrocephaloides liopsettae Burn, 1978

Synonyms: None.

Records: On the gills ofsmooth flounder, Pleuronectes

putnami, Great Bay, New Hampshire (Burn, 1978).

Hendrix: Monogenea of the Eastern United States 79

Remarks: Burn (1980) gave further information on

the seasonal distribution of this species.

Pseudohaliotrema langiphallus (MacCallum, 1915) Young,

1967

Synonyms: Diplectanum longiphallus MacCallum, 1915;

Ancyrocephalus longiphallus (MacCallum, 1915)Johnston

and Tiegs, 1922; A. chaetodipteri Pearse, 1949; and

Tetrancistrum longiphallus (MacCallum, 1915) Price, 1937.

Records: On the gills of the Atlantic spadefish, Chaetodipterus faber, Chesapeake Bay (Zwerner and Lawler,

1972), and Beaufort, North Carolina (Pearse, 1949).

Remarks: Also reported from Biscayne Bay (Skinner,

1978) and Alligator Harbor, Florida.

Family DIONCIDAE Bychowsky, 1959

Diunchus agassizi Goto, 1899

Synonyms: None.

Records: On the gills of the sharksucker, Echeneis

naucrates, Chesapeake Bay (Zwerner and Lawler, 1972);

spearfish remora, Remora brachyptera, Newport, Rhode

Island (Goto, 1899); remora, R remora, Woods Hole,

Massachusetts (Linton, 1940).

Remarks: Also reported from echeneids in the Mediterranean Sea, and the Indian and Pacific Oceans. Redescribed by Price (1938b).

Diunchus remorae (MacCallum, 1916b) Price, 1938b

Synonyms: Acanthodiscus remorae MacCallum, 1916b,

and Dionchotrema remorae (MacCallum, 1916b)Johnston

and Tiegs, 1922.

Records: On the gills ofthe sharksucker, Chesapeake

Bay (Zwerner and Lawler, 1972).

Remarks: Also reported from Alligator Harbor,

Florida, the West Indies, eastern Mediterranean Sea,

Great Barrier Reef, and the Madras coast of India.

Justine and Mattei (1986) described ultrastructural observations on spermatozoa and (1987) some stages of

fertilization. Timofeeva (1988) described genital structures in Dionchus, using D. nagibinae, and reported the

presence ofspermatophores. Whittington (1990) detailed

the attachment of egg bundles to the gills of the remora.

Family DIPLECTANIDAE Bychowsky, 1957

Diplectanum bilobatum Hargis, 1955c

Synonyms: None.

Records: On the gills of spotted seatrout, Cynoscion

nebulosus, Chesapeake Bay (Zwerner and Lawler, 1972).

Remarks: Also reported from Alligator Harbor,

Florida, Mississippi, and Louisiana.
80 NOAA Technical Report NMFS 121

Rhamnocercus bairdiella Hargis, 1955c

Synonyms: None.

Records: On the gills of silver perch, Bairdiella

chrysoura, Chesapeake Bay (Kingston et aI., 1969).

Remarks: Originally described from Alligator Harbor, Florida. Kingston et al. (1969) described the oncomiracidium.

Rhamnocercus stichospinus Seamster and Monaco, 1956

Synonyms: None.

Records: On the gills ofsouthern kingfish, Menticirrhus

americanus, Chesapeake Bay (Zwerner and Lawler, 1972).

Remarks: Also reported from gulfkingfish, M. littoralis,

and Atlantic croaker, Micropogonias undulatus, from

Texas (Seamster and Monaco, 1956).

Order GYRODACTYLIDA Bychowsky, 1937

Family GYRODACTYLIDAE Cobbold, 1864

Kritsky and Thatcher (1977) provided a key to the

subfamilies and genera of these viviparous forms from

North American hosts.

Fundulotrema prolongis (Hargis, 1955a) Kritsky and

Thatcher, 1977

Synonyms: Gyrodactylus sp. of Linton (1940) and G.

prolongis Hargis, 1955a.

Records: On the skin of the sheepshead minnow,

Cyprinodon variegatus, Chesapeake Bay (Zwerner and

Lawler, 1972); skin ofmummichog, Fundulus heteroclitus,

Chesapeake Bay (Dillon, 1966), Bridgeport, Connecticut (Barkman and James, 1979), and Woods Hole

(Linton, 1940); skin ofstriped killifish,F. majalis, Chesapeake Bay (Dillon, 1966).

Remarks: Also reported on various killifishes from

the Gulf of Mexico to Canada. Rcdescribed by Williams

and Rogers (1971). Cone and Odense (1988) emended

the generic diagnosis and provided scanning electron

microscope (SEM) micrographs.

Gyrodactylus stephanus Mueller, 1937

Synonyms: None.

Records: On the skin of the sheepshead minnow,

Chesapeake Bay (Zwerner and Lawler, 1972); skin of

mummichog, Chesapeake Bay (Dillon, 1966), and

Bridgeport, Connecticut (Barkman and James, 1979);

skin ofstriped killifish, Chesapeake Bay (Dillon, 1966).

Remarks: Also reported from gulf killifish, Fundulw

grandis, banded killifish, F. diaphanus, and ninespine

stickleback, Pungitius pungitius. Mizelle and Kritsky

(1967) produced a key to North American species of

Gyrodactylus.

Swingleus sp. of Billeter (1974)

Synonyms: None.

Records: On the skin of the mummichog, Freeport,

New York (Billeter, 1974); striped killifish, Northport,

New York (Billeter, 1974), and the Patuxent River of

the Chesapeake Bay (Billeterl).

Remarks: Billeter (1974) had too few specimens for a

species description, but he did state that the marginal

hooks differed in size and shape from those of S.

polyclithroides Rogers, 1969.

Order POLYOPISTHOCOTYLIDA (Odhner, 1912)

emend. Beverley-Burton, 1984

Superfamily DICLYBOTHROIDEA Bychowsky, 1957

Family DICLYBOTHRIIDAE Bychowsky and Gusev, 1950

Diclybothrium armatum Leuckart, 1835

Synonyms: None.

Records: On the gills of shortnose sturgeon, New

Brunswick, Canada (Appy and Dadswell, 1978).

Remarks: This species is included because the record

is near the geographic range of this manual. This species is also found on lake sturgeon, Acipenserfulvescens,

in the Great Lakes and St. Lawrence River as well as on

Acipenserspp. in Europe and Asia. Wright and Dechtiar

(1974) showed some SEM features of surface structures. Gusev and Slusarev (1986) detailed oncomiracidial

ciliary cells and chaetotaxy.

SuperfamilyHEXABOTHRIOIDEABeverley-Burton, 1984

Family HEXABOTHRIIDAE Price, 1942

Euzet and Maillard (1974) reviewed the historical, systematic, and phylogenetic aspects of this family. Boeger

and Kritsky (1989) discussed its phylogeny and coevolution with hosts and revised the family based on cladistic

analysis.

Erpocotyle maccallumi (Price, 1942) Yamaguti, 1963

Synonyms: Squalonchocotyle canis MacCallum, 1931,

and Neoerpocotyle maccallumi Price, 1942.

Records: On the gills of the blacktip shark, Woods

Hole, Massachusetts (MacCallum, 1931).

Remarks: Price (1942) redescribed the species. It has

also been reported from Paragaleus gruveli (Carcharhinidae) off Senegal, Africa (Euzet and Maillard, 1967).

Erpocotyle macrohystera Price, 1942

Synonyms: Squalonchocotyle vulgaris MacCallum, 1931,

not Cerfontaine, 1899.

I Billeter, P., Department of Biology, Charles County Community

College, LaPlata, MD 20646. Personal commun., August 1980.
Records: On the gills of the sandbar shark, Carcharhinus plumbeus, Woods Hole (MacCallum, 1931).

Remarks: Price (1942) redescribed the species.

Erpocotyle mavori (Linton, 1940) Yamaguti, 1963

Synonyms: Onchocotyle mavori Linton, 1940, and

Neoerpocotyle mavori (Linton, 1940) Price, 1942.

Records: Found on the bottom of an aquarium containing white perch at Woods Hole (Linton, 1940).

Remarks: Boeger and Kritsky (1989) considered this

species incertae sedis, belonging to an as yet undefined

genus. Because hexabothriids are parasites of elasmobranchs, the normal host is probably a shark (Boeger

and Kritsky, 1989).

Erpocotyle microstoma (Brooks, 1934) Yamaguti, 1963

Synonyms: Neoerpocotyle microstoma Price, 1942.

Records: On the gills of the smooth hammerhead,

North Carolina (Brooks, 1934).

Remarks: Also reported from the great hammerhead,

Sphyrna mokarran, in the Pacific Ocean near the Panama

Canal and redescribed by Caballero et a1. (1956) from

this material.

Erpocotyle sphyrnae (MacCallum, 1931) Price, 1942

Synonyms: Squalonchocotyle sphyrnae MacCallum, 1931.

Records: On the gills of the smooth hammerhead,

Woods Hole, Massachusetts (MacCallum, 1931).

Remarks: Also reported from S. zygaena and S. diplana

offSenegal (Euzet and Maillard, 1967) and from S. lewini,

offHawaii (Yamaguti, 1968). Redescribed by Price (1942).

Hexabothrium musteli (MacCallum, 1931) Price, 1942

Synonyms: Acanthonchocotyle musteli MacCallum, 1931,

and Onchocotyle musteli (MacCallum, 1931) Dollfus, 1937.

Records: On the gills of the smooth dogfish, Mustelus

canis, Woods Hole, Massachusetts (MacCallum, 1931).

Remarks: Euzet and Maillard (1967) reported it from

Senegal and Lion Gulf, Mediterranean Sea. Lopez-Roman and De Armas (1987) provided morphological

and SEM micrographs ofthis species from hosts around

the Canary Islands. It was redescribed by Price (1942)

and Maillard (1972). Boeger and Kritsky (1989) considered this species incertae sedis.

Rajonchocotyle laevis Price, 1942

Synonyms: None.

Records: On the gills of the barndoor skate, Raja

laevis, Woods Hole (Price, 1942).

Remarks: This species was described from a single

specimen.

Squalonchocotyle abbreviata (Olsson, 1876) Cerfontaine,

1899

Synonyms: Onchocotyle abbreviata Olsson, 1876, and

Erpocotyle abbreviata Price, 1942.

Hendrix: Monogenea of the Eastern United States 81

Records: On the gills of spiny dogfish, Atlantic coast

of Canada (Threlfall, 1969).

Remarks: I include this species because the record is

near the geographic range of the manual. It is also known

from the Irish Sea; Norway; Peter the Great Bay, Russia;

Roscoff, France; the Atlantic Ocean; and the North Sea.

Squalonchocotyle squali MacCallum, 1931

Synonyms: Squalonchocotyle acanthi MacCallum, 1931,

and Erpocotyle squali Price, 1942.

Records: On the gills of the spiny dogfish, Woods

Hole, Massachusetts (MacCallum, 1931).

Remarks: Redescribed by Price (1942). Also reported

from New Zealand and the Black Sea.

Superfamily MAZOCRAEOIDEA (Bychowsky, 1957)

emend. Beverley-Burton, 1984

Family ANTHOCOTYLIDAE Bychowsky, 1957

Anthocotyle merluccii van Beneden and Hesse, 1863

Synonyms: Anthocotyle merluccii americanus MacCallum,

1916b, and A. americanus (MacCallum, 1916b) Price,

1943b.

Records: On the gills ofsilver hake, Merluccius bilinearis,

New Jersey (Meyers, 1978), Woods Hole, Massachusetts

(MacCallum, 1916b); offshore hake, M. albidus, offChesapeake Bay and NewJersey (new host record).

Remarks: Also reported from silver hake in Canada

and from hakes (Gadidae) in the eastern, southeastern,

and southwestern Atlantic Ocean; Mediterranean Sea;

and the eastern Pacific Ocean off Eureka, California;

the Strait of Georgia, Canada; and Peru. Redescribed

by Price (1943b). Llewellyn (1956) described the

microecology as well as adhesive attitude, and in 1963

he described larval development. Lyons (1966) studied

histochemistry of the sclerites.

Family AXINIDAE Unnithan, 1957

Axine hyporhampi Price, 1962a

Synonyms: Axinoides hyporhampi of Kingston et a1.

(1969) and of Zwerner and Lawler (1972).

Records: On the gills of the silverstripe halfbeak,

Hyporhamphus unifasciatus, Chesapeake Bay (Kingston

et aI., 1969) and Woods Hole, Massachusetts (Price,

1962a).

Remarks: None.

Nudaciraxine gracilis (Linton, 1940) Price, 1962a

Synonyms: Axine gracilis Linton, 1940, and Axinoides

gracilis (Linton, 1940) Sproston, 1946.

Records: On the gills of Atlantic needlefish, Chesapeake Bay (McMahon, 1964; Kingston et aI., 1969) and

Woods Hole, Massachusetts (Linton, 1940).
82 NOAA Technical Report NMFS 121

Remarks: Hargis (l956c) reported this species at Alligator Harbor, Florida. Redescribed by Price (1962a).

Kingston et al. (1969) described the oncomiracidium.

Family DICUDOPHORIDAE Fuhrmann, 1928

Absonifibula bychowskyi Lawler and Overstreet, 1976

Synonyms: None.

Records: On gills of the Atlantic croaker, Chesapeake

Bay and Pamlico Sound, North Carolina (Thoney, 1991).

Remarks: Originally described from this host in Mississippi Sound.

Choricotyle aspinachorda Hargis, 1955h

Possible synonym: Diclidophora sp. of Linton (1905).

Records: On the gills of the pigfish, Orthopristis

chrysoptera, Chesapeake Bay (Kingston et al., 1969) and

Beaufort, North Carolina (Linton, 1905).

Remarks: Hargis (1955h) described the species from

Alligator Harbor, Florida, specimens. It has been reported from O. ruber in Venezuela (Bashirullah and

Rado, 1987). According to Hargis (1959) this species

may be a synonym of C. caudalis Koratha, 1955b.

Choricotyle louisianensis Hargis, 1955h

Synonyms: None.

Records: On the gills of the northern kingfish, Menticirrhus saxatilis, off Chesapeake Bay (Zwerner and

Lawler, 1972).

Remarks: The type host is the southern kingfish, M.

americanus, from Louisiana. It was also reported from

Puntarenas, Costa Rica.

Diclidophora denticulata (Olsson, 1876) Price, 1943a

Synonyms: Octobothrium denticulatum Olsson, 1876; Diclidophora carbonarii Cerfontaine, 1895; and Dactylocotyle

denticulata (Olsson, 1876) Cerfontaine, 1895.

Records: On the gills of the pollock, Pollachius virens,

Woods Hole, Massachussets (Linton, 1900).

Remarks: Also reported from Canada, Ireland, Scotland, England (but see Llewellyn et al., 1980), the North

Sea, France, the Barents Sea, and the Mediterranean

Sea. Redescribed by Price (1943a). Its life cycle and

biology were described by Frankland (1955). Sproston

(1945b) and Llewellyn (1958) described the gill clamping mechanism. Egg hatching rhythm was studied by

MacDonald (1975). Vitelline cell histochemistry was

reported by Halton et al. (1974) and amino acid composition by Arme (1977). Llewellyn and Tully (1969)

assessed speciation in diclidophorans and their gadid hosts.

Diclidophoroides maccallumi Price, 1943a

Synonyms: Dactylocotylephycidis ofSrnfford (1904); Dactylocotyle minor of Manter (1926); Diclidophora maccallumi

(Price, 1943a) Sproston, 1946; and Choricotyle merlangi

(MacCallum, 1917) Llewellyn, 1941.

Records: On the gills of red hake, Urophycis chuss,

Mount Desert Island, Maine (Manter, 1925 and 1926),

NewJersey (Meyers, 1978); spotted hake, U. regia, Chesapeake Bay (Kingston et al., 1969) and North Carolina

(Suydam, 1971); on silver hake, Woods Hole, Massachusetts (MacCallum, 1917).

Remarks: Redescribed by Price (1943a) from U. chuss

and by Rubec (1991) from Canadian U. chesteri. Suydam

(1971) described the microecology of this species on

host gills.

Neoheterobothrium affine (Linton, 1898) Price, 1943a

Synonyms: Octoplectanum affine Linton, 1898; Diclidophora affinis (Linton, 1898) Linton, 1901; Choricotyle

affine (Linton, 1898) Llewellyn, 1941, not Heterobothrium

affine of Nagibina, 1953.

Records: On the gills and in mouth ofsummer flounder, Paralichthys dentatus, Chesapeake Bay (Jansen and

Burreson, 1990), NewJersey (Meyers, 1978), and Woods

Hole, Massachusetts (Linton, 1898).

Remarks: Also reported from the mouth ofthe southern flounder, P. lethostigma, Louisiana. Redescribed by

Price (1943a). Mamaev (1987) redefined the genus.

Neoheterobothrium cynoscioni (MacCallum, 1917) Price,

1943a

Synonyms: Diclidophora cynoscioni MacCallum, 1917;

Choricotyle cynoscioni (MacCallum, 1917) Llewellyn, 1941;

and C. reynoldsi Frayne, 1943.

Records: On the gills of weakfish, Cynoscion regalis,

Chesapeake Bay (Frayne, 1943; Kingston et al., 1969),

Woods Hole, Massachusetts (MacCallum, 1917); spotted seatrout, Chesapeake Bay (Frayne, 1943).

Remarks: Also reported from silver seatrout, C. nothus,

in Florida and Louisiana: spotted seatrout and sand

seatrout, C. arenarius, mouth, gills, and skin, Mississippi; and grunts (Haemulidae), Venezuela. Redescribed

by Price (1943a) and Frayne (1943).

Orbocotyle prionoti (MacCallum, 1917) Euzet and Suriano,

1975

Synonyms: Diclidophora prionoti MacCallum, 1917;

Cyclocotyla prionoti (MacCallum, 1917) Price, 1943a; and

Choricotyle prionoti (MacCallum, 1917) Llewellyn, 1941.

Records: On the gills ofthe northern searobin, Prionotus carolinus, Woods Hole, Massachusetts (MacCallum,

1917); striped searobin, P. evolans, off Chesapeake Bay

(Zwerner and Lawler, 1972).

Remarks: Hargis (1955h) reported this species from

the leopard searobin, P. scitulus, in Florida and Louisiana. Redescribed by Price (1943a).
Pedocotyle minima Hargis, 1955h

Synonyms: None.

Records: On the gills ofsilver perch, Chesapeake Bay

(Kingston et al., 1969).

Remarks: The species was described from Alligator

Harbor, Florida, hosts. Kingston et al. (1969) described

the oncomiracidium.

Pedocotyle morone (MacCallum, 1913c) MacCallum,

1913d

Synonyms: Podocotyle morone MacCallum, 1913c.

Records: On the gills of white perch, New York City

fish market (MacCallum, 1913c).

Remarks: Hargis (1955h) questioned the validity of

this host record. The species was redescribed by Price

(1943a) .

Family DISCOCOTYLIDAE Price, 1936

Bicotylophora trachinoti (MacCallum, 1921) Price, 1936

Synonyms: Dactylocotyle trachinoti MacCallum, 1921.

Records: On the gills ofFlorida pompano, Trachinotus

carolinus, Chesapeake Bay (McMahon, 1963).

Remarks: Also reported from Alligator Harbor,

Florida, as well as Mississippi, Mexico, Venezuela, and

Uruguay; on permit, T. falcatus, Mexico and the Ivory

Coast; and on the gafftopsail pompano, T. rhodopus,

Mexico. Redescribed by McMahon (1963), Caballero

and Bravo-Hollis (1965), as well as Nasir and Fuentes

Zambrano (1983). Lawler (1977) mentioned its role as

a pest in aquaculture. Kohn et al. (1992) provided

measurements and SEM micrographs of this species

from Brazilian T. carolinus.

Family GASTROCOTYLIDAE Price, 1943a

Gotocotyla acanthophallus (MacCallum and MacCallum,

1913) Yamaguti, 1963

Synonyms: Microcotyle acanthophallus MacCallum and

MacCallum, 1913, and Lithidocotyle acanthophallus

(MacCallum and MacCallum, 1913) Sproston, 1946,

and of Hargis (1956d).

Records: On the gills of bluefish, Pomatomus saltatrix,

Chesapeake Bay (McMahon, 1964; Kingston etal., 1969);

king mackerel, Scomberomorus cavalla, Chesapeake Bay

(Zwerner and Lawler, 1972); Spanish mackerel, S.

maculatus, Chesapeake Bay (McMahon, 1964); and

striped bass, Morone saxatilis, Atlantic Ocean (New York

City fish market) (MacCallum and MacCallum, 1913).

Remarks: Also reported from mackerels (Scombridae)

in Florida, Louisiana, and Mexico. Hargis (1956c) considered the striped bass to be an accidental host.

Hendrix: Monogenea of the Eastern United States 83

Pseudaxine mexicana Meserve, 1938

Probable synonym: Pseudaxine texana Koratha, 1955b.

Records: On the gills of Spanish mackerel, Chesapeake Bay (McMahon, 1964).

Remarks: Also reported from mackerels in Florida,

Louisiana, Texas, and Mexico. Redescribed by BravoHollis (1953). Dillon and Hargis (1965b) emended the

generic diagnosis. Lebedev (1986) removed this species to the monotypic genus Mexicotyle. Bravo-Hollis

(1989) gave new Mexican collecting localities and compared the gastrocotylid species of the Atlantic and Pacific Oceans off Mexico.

Family HETERAXINIDAE Price, 1962b

Mamaev (1990) provided a systematic list of members

of this and related families under the suborder Microcotylinae.

Cemocotyle carangis (MacCallum, 1913c) Sproston, 1946

Synonyms: Microcotyle carangis MacCallum, 1913c, and

Cotocotyla carangis (MacCallum, 1913c) Meserve, 1938.

Records: On the gills of blue runner, Caranx crysos,

Woods Hole, Massachusetts (MacCallum, 1913c)

Remarks: Also reported at Alligator Harbor, Florida,

and Veracruz, Mexico. Redescribed by Price (1962b).

Bravo-Hollis and Salgado-Maldonado (1983) provided

new morphological data on specimens collected from

Mexican C. crysos.

Cemocotylenoveborancensis (MacCallum, 1918) Sproston,

1946

Synonyms: Axine carangis MacCallum, 1918, A.

(Heteraxine) carangis (MacCallum, 1918) Yamaguti, 1938,

and Heteraxine carangis (MacCallum, 1918) Yamaguti,

1938.

Records: On the gills of crevalle jack, Caranx hippos,

New York region (MacCallum, 1918).

Remarks: Also reported from Alligator Harbor,

Florida, and Campeche, Mexico. Redescribed by Price

(1962b). Bravo-Hollis and Salgado-Maldonado (1983)

gave new morphological data on specimens from Mexican C. hippos.

Heteraxinoides xanthophilis (Hargis, 1956c) Yamaguti,

1963

Synonyms: Heteraxine xanthophilis Hargis, 1956c, and

Heteraxinoides xanthophiloides Price, 1962b.

Records: On the gills of spot, Leiostomus xanthurus,

Chesapeake Bay (Kingston et al., 1969; Thoney, 1988, a

and b, 1991), Pamlico Sound, North Carolina (Thoney,

1991), and Woods Hole, Massachusetts (Price, 1962b).
84 NOAA Technical Report NMFS 121

Remarks: Also reported from Florida (Hargis, 1956c)

and Texas (Joy, 1976). Kingston etal. (1969) described

the oncomiracidium. Thoney (1988a) showed its developmental variation on hosts of different sizes and

(1988b) studied egg and oncomiracidial structures.

Thoney (1991) discussed population dynamics of this

and other parasites on juvenile spot.

Family HEXOSTOMATIDAE Price, 1936

Hexostoma lintoni Price, 1961a

Synonyms: Hexacotyle thynni of Linton (1901).

Records: In the mouth ofAtlantic bonito, Sarda sarda,

Woods Hole, Massachusetts (Linton, 1901 and 1940).

Remarks: Price (1961a) redescribed this species from

a single specimen.

Neohexostoma euthynni (Meserve, 1938) Price, 1961a

Synonyms: Hexostoma euthynni Meserve, 1938, and H.

macracanthum Fujii, 1944.

Records: On the gills oflittle tunny, Chesapeake Bay

(Zwerner and Lawler, 1972).

Remarks: Also reported from scombrids at the

Tortugas, Florida; Baja California; Galapagos Islands;

Great Barrier Reef, Australia; and the South China Sea.

Redescribed by Millemann (1956). Rohde (1978a) considered this genus to be synonymous with Hexostoma.

Rohde (1980) described some aspects of its cecal ultrastructure.

Family MACROVALVITREMATIDAE Yamaguti, 1963

Hargisia bairdiella (Hargis, 1956a) Yamaguti, 1963

Synonyms: Tagia bairdiella Hargis, 1956a.

Records: On the gills ofsilver perch, Chesapeake Bay

(McMahon, 1963; Kingston et aI., 1969).

Remarks: First described from Alligator Harbor,

Florida, by Hargis (1956a).

Macrovalvitrematoides micropogoni (Pearse, 1949)

Yamaguti, 1963

Synonyms: Tagia micropogoni Pearse, 1949.

Records: On the gills of Atlantic croaker, Chesapeake Bay (Kingston et aI., 1969) and Beaufort, North

Carolina (Pearse, 1949).

Remarks: Also reported from Florida, Mississippi,

and Texas (Joy and Price, 1976). Redescribed by Hargis

(l956a). Kingston et al. (1969) described the

oncomiracidium. Bravo-Hollis (1981) erected the subfamily Macrovalvitrematinae containing this genus.

Pseudotagia cupida (Hargis, 1956a) Yamaguti, 1963

Synonyms: Tagia cupida Hargis, 1956a.

Records: On the gills of pigfish, Chesapeake Bay

(Kingston eta!., 1969), offNorth Carolina (Suydam, 1971).

Remarks: Hargis (1956a) described the species from

Alligator Harbor, Florida. The species was redescribed

by Kohn et al. (1984) from bluestriped grunts, Haemulon

sciurus, and Kohn et al. (1992) gave a new host record

for Brazil.

Family MAZOCRAEIDAE Price, 1936

Mamaev (1982a) reviewed the systematics of several

genera within the family.

Oupeocotyle brevoortia Hargis, 1955f

Synonyms: Dactylocotyle sp. of Linton (1905); Diclidophora lintoni Koratha, 1955b; Clupeocotyle lintoni (Koratha,

1955b) Hargis, 1959; and probably Diclodophora sp. [sic]

of Westman and Nigrelli, 1955.

Records: On the gills ofAtlantic menhaden, Brevoortia

tyrannus, Beaufort, North Carolina (Linton, 1905),

Chesapeake Bay (McMahon, 1963), New Jersey, and

Long Island (Westman and Nigrelli, 1955).

Remarks: Also reported from gulf menhaden, B.

patronus, in Florida and Texas. Redescribed by McMahon

(1963) .

Grubea cochlear Diesing, 1858

Synonyms: Pleurocotyle scombri Linton, 1940, and Orubea

pneumatophori Price, 1961 b.

Records: On the gills of the chub mackerel, Scomber

japonicus, Woods Hole, Massachusetts (Linton, 1940).

Remarks: Also reported from mackerels from Brazil,

the Mediterranean Sea, the Portuguese coast, and the

Patagonian shelf of the southwest Atlantic Ocean.

Wagner (1975) provided some comparative measurements for species in this genus. Mamaev (1982b) redefined the genus and redescribed the species. Rohde

(l987a) summarized global locality data and redescribed

the species.

Kuhnia scombri (Kuhn, 1829) Sproston, 1945a

Synonyms: Octostoma scombri Kuhn, 1829.

Records: On the gills of Atlantic mackerel, Scomber

scombrus, Chesapeake Bay (Price, 1961b); Newport,

Rhode Island (Goto, 1899); off Cape Hatteras, North

Carolina (Romuk-Wodoracki, 1988); and Woods Hole,

Massachusetts (Price, 1961b).

Remarks: Also reported on mackerels from Canada,

the eastern and southwestern Atlantic Ocean, Mediterranean Sea, eastern and southwestern Pacific Ocean,

and Hawaii. This is a cosmopolitan species on mackerels. Rohde (1989) summarized the known geographical

distribution of Kuhnia spp. from Scomber spp. The species was redescribed by Price (1961 b) and by Nasir and
Fuentes Zambrano (1983). Sproston (1945a) and

Llewellyn (1957) described the anatomy of the clamping mechanism on host gills; Llewellyn (1956), the gill

microecology and (1963) larval development; and Euzet

(1957), the oncomiracidium. Finlayson (1982) provided

detailed information on the reproductive processes of

this species. Rohde (1991) and Rohde and Watson

(1985) studied geographic variation in morphology and

microhabitat in Kuhnia. Rohde (1987b) studied sclerite

morphology. Mamaev and Parukhin (1986) revised the

genus composition and provided a table of comparative

characteristics of Kuhnia spp.

Mazocraeoides georgei Price, 1936

Synonyms: None.

Records: On the gills of blueback herring, Alosa

aestivalis, Chesapeake Bay (Zwerner and Lawler, 1972);

hickory shad, A. mediocris, Woods Hole, Massachusetts

(Linton, 1940); alewife, A. pseudoharengus, Chesapeake

Bay (Zwerner and Lawler, 1972), New Jersey (Meyers,

1978), and Woods Hole, Massachusetts (Linton, 1940);

and Atlantic menhaden, Chesapeake Bay (McMahon,

1963).

Remarks: Hargis (1955g) and McMahon (1963) redescribed the species (but see below). Zwerner and

Lawler (1972) suggested that the oncomiracidium described by Kingston et al. (1969) was probably that ofthis

species. Kohn and Portes Santos (1988) provided measurements and illustrations from Brazilian clupeid fishes.

Mazocraeoides hargisi Price, 1961b

Possible synonyms: Mazocraeoides georgei of Hargis

(1955g) and McMahon (1963).

Records: On the gills of Atlantic menhaden, Chesapeake Bay (Kingston et aI., 1969).

Remarks: Kingston et aI. (1969) described the oncomiracidium. Zwerner and Lawler (1972) questioned

this record and suggested that it was M. georgei. Kohn

and Portes Santos (1988) considered this to be a synonym of M. georgei.

Mazocraeoides olentangiensis Sroufe, 1958

Synonyms: Mazocraeoides similis Price, 1958.

Records: On the gills of gizzard shad, Dorosoma cepedianum, Chesapeake Bay (Zwerner and Lawler, 1972).

Remarks: Originally described from shad in the

Olentangy River, Ohio (Sroufe, 1958). It is also known

from Lake Erie; Norris and Reelfoot Lakes, Tennessee;

and the Tennessee River, Alabama. Wright and Dechtier

(1974) described light and SEM features of surface

structures. Kohn and Portes Santos (1988) synonymized

this with M. georgei. This is primarily a freshwater species

that is also found in slightly brackish estuarine waters.

Hendrix: Monogenea of the Eastern United States 85

Family MICROCOTYLIDAE Taschenberg, 1879

Mamaev (1986) presented a briefsummary of the taxonomic system for the family.

Cynoscionicola heteracantha (Manter, 1938) Price, 1962b

Synonyms: Microcotyle heteracantha Manter, 1938, and

Microcotyle sp. of Linton (1905 and 1940).

Records: On the gills of spotted seatrout, Chesapeake Bay (Zwerner and Lawler, 1972), and Beaufort,

North Carolina (Manter, 1938); weakfish, Woods Hole,

Massachusetts (Linton, 1940), Chesapeake Bay

(Kingston et aI., 1969), and Beaufort, North Carolina

(Linton, 1905).

Remarks: Also reported from spotted seatrout collected at Alligator Harbor, Florida (Hargis, 1956b). Kingston et aI. (1969) described the oncomiracidium. Lambert and Euzet (1979) reviewed the genus.

Cynoscionicola pseudoheteracantha (Hargis, 1956b) Price,

1962b

Synonyms: Microcotyle pseudoheteracantha Hargis, 1956b.

Records: On the gills of silver seatrout, Chesapeake

Bay (Zwerner and Lawler, 1972).

Remarks: Originally described from Louisiana hosts.

Gamacallum macroura (MacCallum and MacCallum,

1913) Unnithan, 1971

Synonyms: Microcotyle macroura MacCallum and

MacCallum, 1913.

Records: On the gills ofstriped bass, Chesapeake Bay

(Zwerner and Lawler, 1972), Atlantic coast of the U.S.

(MacCallum and MacCallum, 1913).

Remarks: None.

Metamicrocotyla macracantha (Alexander, 1954) Koratha,

1955b

Synonyms: Microcotyle macracantha Alexander, 1954.

Records: On the gills of striped mullet, Chesapeake

Bay (Zwerner and Lawler, 1972).

Remarks: Also reported from Georgia (Rawson, 1976);

Biscayne Bay, Florida (Skinner, 1975); U.S. Gulf of

Mexico coast; Puerto Rico; and the Pacific Ocean in the

Gulf of California, Mexico, and Peru. Redescribed by

Hargis (1956b) and also by Kohn et al. (1994), who

redescribed the species from Brazilian Mugilliza. Garcia

and Williams (1985) gave some temporal data on this

species from Puerto Rico.

Microcotyle archosargi MacCallum, 1931

Synonyms: None.

Records: On the gills of sheepshead from the New

York City fish market (MacCallum, 1931).

Remarks: Also reported from Alligator Harbor,

Florida (Hargis, 1956b).
86 NOAA Technical Report NMFS 121

Microcotyle centr()j»"istis MacCallum, 1915

Synonyms: None.

Records: On the gills of black sea bass, Centropristis

striata, New York City fish market (MacCallum, 1915).

Remarks: None.

Microcotyle eueides MacCallum and MacCallum, 1913

Synonyms: None.

Records: On the gills ofstriped bass along the northwestern Atlantic coast (MacCallum and MacCallum,

1913).

Remarks: None.

Microcotyle hiatulae Goto, 1894

Synonyms: Microcotyle furcata Linton, 1940.

Records: On the gills of tautog, Tautoga onitis, Newport, Rhode Island (GolO, 1899), Woods Hole (Linton,

1940), and the mouth ofChesapeake Bay (Thoney and

Munroe, 1987).

Remarks: Thoney and Munroe (1987) redescribed

the species and gave comments on its postlarval development.

Microcotyle langicauda Goto, 1899

Synonyms: Cynoscionicola longicauda (Goto, 1899)

Yamaguti, 1963.

Records: On the gills of weakfish, 'ewport, Rhude

Island (Goto, 1899).

Remarks: None.

Microcotyle otrynteri Pearse, 1949

Synonyms: None.

Records: On the gills of longspinc porgy, Stenotomus

caprinus, Beaufort, North Carolina (Pearse, 1949).

Remarks: The species is poorly described and needs

further investigation.

Microcotyle peprili Pearse, 1949

Synonyms: None.

Records: On the gills ofharvestfish, Peprilus alepidotus,

Chesapeake Bay (McMahon, 1964; Kingston et aI., 1969).

and Beaufort, North Carolina (Pearse, 1949).

Remarks: Redescribed by McMahon (1964).

Microcotyle pomatomi Goto, 1899

Synonyms: Microcotyle australiensis MacCallum, 1921;

M. debueni Mane-Garzon, 1959; and M. temnodontis

Sandars, 1945.

Records: On the gills of bluefish, Beaufort (Linton,

1905; Pearse, 1949) and Hatteras, North Carolina

(Anderson, 1970); Chesapeake Bay (McMahon, 1964);

New Jersey (Meyers, 1978); Newport, Rhode Island

(Goto, 1899); and Woods Hole, Massachusetts (Linton,

1901 and 1940).

Remarks: Although widely distributed along the Atlantic and Gulf coasts of the United States, M. pomatomi

has also been reported from the Portuguese coast; Walvis

Bay, Namibia; the Black Sea; and Australia. It was redescribed by Koratha (1955b), McMahon (1964), and

Williams (1991) who also gave comparative measurements from various authors and discussed the taxonomy

of this species. Anderson (1970) reported the geographic range of this species along the U.S. Atlantic

coast.

Microcotyle poranoti MacCallum, 1915

Synonyms: None.

Records: On the gills ofbutterfish, Peprilus triacanthus,

Chesapeake Bay (McMahon, 1964; Kingston et aI., 1969),

New Jersey (Meyers, 1978), and Woods Hole, Massachusetts (MacCallum, 1915, Linton, 1940).

Remarks: Also reported from butterfish in Canada.

Redescribed by McMahon (1964). Kingston et a1. (1969)

described the oncomiracidium.

Microcotyle stenotomi Goto, 1899

Synonyms: None.

Records: On the gills of scup, Stenotomus chrysops,

Woods Hole, Massachusetts (Linton, 1940); Newport,

Rhode Island (Goto, 1899); Chesapeake Bay (McMahon,

1964); and off orth Carolina (Suydam, 1971).

Remarks: MacCallum (1913a) briefly described fertilization and egg laying. Suydam (1971) described the

distribution of worms on gill arches.

Paucicanfibula poganiae (MacCallum, 1913c) Chisholm,

Beverley-Burton, and McAlpine, 1991

Synonyms: Microcotyle pogoniae MacCallum, 1913c;

llspinatrium pogoniae (MacCallum, 1913c) Yamaguti,

1963; and Pseudoaspinatrium pogoniae (MacCallum,

1913c) Mamaev, 1986.

Records: On the gills of black drum, Pogonias cromis,

New York region (MacCallum, 1913c).

Remarks: Also reported from Alligator Harbor,

Florida, by Hargis (1956b). Chisholm et a1. (1991)

emended the generic diagnosis.

Family NEOTHORACOCOTYLIDAE Lebedev, 1969

Neothoracocotyle acanthocybii (Meserve, 1938) Hargis,

1956d

Synonyms: Gotocotyla acanthocybii Meserve, 1938.

Records: On the gills ofwahoo, Acanthocybium solandri,

offshore from Chesapeake Bay (Zwerner and Lawler,

1972) .

Remarks: Neothoracocotyle acanthocybii was described

from the Galapagos Islands and has been reported

from Hawaii; off Noumea, New Caledonia; and from
the Great Barrier Reef. Yamaguti (1968) redescribed

the species.

Scomberocotyle scomberomori (Koratha, 1955b) Hargis,

1956d

Synonyms: Microcotyle scomberomori Koratha, 1955b.

Reports: On the gills of king mackerel (Zwerner and

Lawler, 1972) and Spanish mackerel, Chesapeake Bay

(McMahon, 1964).

Remarks: Also reported from the Gulf of Mexico off

Florida and Texas.

Thoracocotyle crocea MacCallum, 1913b

Synonyms: ThoracocotyleparadoxicaMeserve, 1938, and

possibly of Pearse, 1949.

Records: On the gills ofSpanish mackerel, New York

fish market (MacCallum, 1913b), Chesapeake Bay

(McMahon, 1964; Kingston et al., 1969); and king mackerel, Beaufort, North Carolina (Pearse, 1949).

Remarks: Also reported from Florida and Mexico.

Redescribed by McMahon (1964).

Family PROTOMICROCOTYLIDAE Poche, 1926

Protomicrocotyle mirabilis (MacCallum, 1918) Johnston

and Tiegs, 1922

Synonyms: Acanthodiscus mirabilis MacCallum, 1918.

Records: On the gills of crevalle jack, Chesapeake

Bay (Zwerner and Lawler, 1972).

Remarks: Also reported on crevalle jack from Alligator Harbor, Florida; Texas; and the Ivory Coast; and

from the horse-eyejack, Caranx latus, Mexico. Redescribed

by Koratha (1955b) and by Caballero and Bravo-Hollis

(1965). Bashirullah and Rodriguez (1992) analyzed the

distribution of P. mirabilis and other monogenean species on the gill arches of Venezuelan C. hippos.

Family PYRAGRAPHORIDAE Yamaguti, 1963

Pyragraphorus pyragraphorus (MacCallum and

MacCallum, 1913) Sproston, 1946

Synonyms: Microcotyle pyragraphorus MacCallum and

MacCallum, 1913.

Records: On the gills of Florida pompano, Atlantic

coast ofthe United States (MacCallum and MacCallum,

1913).

Remarks: Also reported from Baja California, Mexico

(Bravo-Hollis, 1978), and the Great Barrier Reef, Australia. Redescribed by Hargis (1956b). Bravo-Hollis

(1984) gave some comparative measurements.

Hendrix: Monogenea of the Eastern United States 87

Host-Monogenea List _

This host-Monogenea list includes only those species

on fishes occurring along the western Atlantic coast

from the U.S.-Canada border to Beaufort, North Carolina. Hosts from more distant localities are not listed.

Older host names mentioned in older Monogenea reports have been updated and are those given in the

American Fisheries Society Special Publication No. 20

(Robins et al.,1991). Fish families and parasites are

listed alphabetically within each order offishes.

Class ElASMOBRANCHIOMORPHI

Order LAMNIFORMES

Family Alopiidae - thresher sharks

Alopias vulpinus - thresher shark

Cathariotrema selachii

Family Carcharhinidae - requiem sharks

Carcharhinus limbatus - blacktip shark

Dermophthirius penneri

Erpocotyle maccallumi

Carcharhinus obscurus - dusky shark

Cathariotrema selachii

Dermophthirius carcharhini

Loimos salpinggoides

Microbothrium apiculatum

Carcharhinus plumbeus - sandbar shark

Erpocotyle macrohystera

Mustelus canis - smooth dogfish

Hexabothrium musteli

Negaprion brevirostris- lemon shark

Neodermophthirius harkemai

Rhizoprionodon terraenovae - Atlantic sharpnose shark

Loimos scoliodoni

Family Sphymidae - hammerhead sharks

Sphyrna zygaena - smooth hammerhead shark

Cathariotrema selachii

Erpocotyle microstoma

Erpocotyle sphymae

Tristoma coccineum

Order SQUALIFORMES

Family Squalidae - dogfish sharks

Squalus acanthias - spiny dogfish

Amphibdelloides maccallumi

Heterocotyle minima

Microbothrium apiculatum

Squalonchocotyle abbreviata

Squalonchocotyle squali
88 NOAA Technical Report NMFS 121

Order RAJIFORMES

Family Dasyatidae - stingrays

Dasyatis americana - southern stingray

Monocotyle diademalis

Monocotyle pricei

Dasyatis centroura - roughtail stingray

Dasybatotrema dasybatis

Entobdella lrumpusii

Heterocotyle minima

Thaumatocotyle dasybatis

Dasyatis say - bluntnose stingray

Loimopapillosum dasyatis

Monoeotyle diademalis

Monocotyle prieei

Family Myliobatidae - eagle rays

Aetobatis narinari - spotted eagle ray

PapillicotyleJloridana

Rhinoptera bonasus - cownose ray

Benedeniella posterocolpa

Family Rajidae - skates

Raja eglanteria - clearnose skate

Empruthotrema raiae

Raja erinacea - little skate

Empruthotrema raiae

Pseudacanthocotyla verrilli

Thaumatocotyle dasybatis

Raja laevis - barndoor skate

Rajonchocotyle laevis

Raja ocellata - winter skate

Empruthotrema raiae

Raja radiata - thorny skate

Pseudacanthocotyla verrilli

Family Torpedinidae - electric rays

Torpedo nobiliana - Atlantic torpedo

A mphibdella Jlavolineata

Amphibdelloides maecallumi

Class OSTEICHTHYES

Order ACIPENSERIFORMES

Family Acipenseridae - sturgeons

Acipenser brevirostrum - shortnose sturgeon

Diclybothrium armatum

Nitzschia superba

Acipenser oxyrhynchus - Atlantic sturgeon

Nitzschia sturionis

Nitzschia superba

Order CLUPEIFORMES

Family Clupeidae - herrings

Alosa aestivalis - blueback herring

Ma:oeraeoides georgei

Alosa mediocris - hickory shad

Mazoeraeoides georgei

Alosa pseudoharengus - alewife

Mazoeraeoides georgei

Brevoortia tyrannus - Atlantic menhaden

Clupeocotyle brevoortia

Mazoeraeoides georgei

Mazocraeoides hargisi

Dorosoma cepedianum - gizzard shad

Mazocraeoides olentangiensis

Order GADIFORMES

Family Gadidae - cods

Gadus morhua - Atlantic cod

Udonella caligorum

Merluccius albidus - offshore hake

Anthocotyle merluccii

Merluccius bilinearis - silver hake

Anthocotyle merluccii

Diclidophoroides maccallumi

Pollachius virens - pollock

Diclidophora dentieulata

Urophycis chuss - red hake

Diclidophoroides maecallumi

Urophycis regia - spotted hake

Diclidophoroides maecallumi
Order ATHERINIFORMES

Family Belonidae - needlefishes

Strongylura marina - Atlantic needlefish

Ancyrocephalus parous

Nudaciraxine gracilis

Family Cyprinodontidae - killifishes

Cyprinodon variegatus - sheepshead minnow

Fundulotrema prolongis

Gyrodactylus stephanus

Fundulus heteroclitus - mummichog

Fundulotrema prolongis

Gyrodactylus stephanus

SWingleus sp.

Fundulus majalis - striped killifish

Fundulotrema prolongis

Gyrodactylus stephanus

Swingleus sp.

Family Exocoetidae - flyingfishes

Hyporhamphus unifasciatus - silverstripe halfbeak

Axine hyporhampi

Order SCORPAENIFORMES

Family Cydopteridae - snailfishes

Cyclopterus lumpus - lumpfish

Udonella caligorum

Family Triglidae - searobins

Prionotus carolinus - northern searobin

Orbocotyle prionoti

Prionotus evolans - striped searobin

Orbocotyle prionoti

Order PERCIFORMES

Family Carangidae - jacks

Caranx crysos - blue runner

Cemocotyle carangis

Caranx hippos - crevaIle jack

Cemocotyle noveborancensis

Protomicrocotyle mirabilis

Trachinotus carolinus - Florida pompano

Bicotylophora trachinoti

Pyragraphorus pyragraphorus

Hendrix: Monogenea of the Eastern United States 89

Family Echeneidae - remoras

Echeneis naucrates - sharksucker

Dionchus agassizi

Dionchus remorae

Remora brachyptera - spearfish remora

Dionchus agassizi

Remora remora - remora

Dionchus agassizi

Family Ephippidae - spadefishes

Chaetodiptentsfaber - Atlantic spadefish

Pseudohaliotrema longiphallus

Family Haemulidae - grunts

Orthopristis chrysoptera - pigfish

Choricotyle aspinachorda

Pseudotagia cupida

Family Istiophoridae - billfishes

Tetrapturus albidus - white marlin

Capsaloides comutus

Capsaloides magnaspinosus

Tristomella laevis

Family Labridae - wrasses

Tautoga onitis - tautog

Microcotyle hiatulae

Family Mugilidae - mullets

Mugil cephalus - striped mullet

Haliotrema vanbenedeni

Metamicrocotyla macracantha

Family Percichthyidae - temperate basses

Morone americana - white perch

Erpocotyle mavori ?

Onchocleidus nactus

Pedocotyle morone

Morone saxatilis - striped bass

Gamacallum macroura

Gotocotyla acanthophallus

Microcotyle eueides

Family Pomatomidae - bluefishes

Pomatomus saltatrix - bluefish

Gotocotyla acanthophallus

Microcotyle pomatomi

Family Sciaenidae - drums

Bairdiella chrysoura - silver perch

Hargisia bairdiella

Pedocotyle minima

Rhamnocercus bairdiella
90 NOAA Technical Report NMFS 121

Cynoscion nebulosus - spotted seatrout

Cynoscionicola heteracantha

Diplectanum bilobatum

Neoheterobothrium cynoscioni

Cynoscion nothus - silver seatrout

Cynoscionicola pseudoheteracantha

Cynoscion regalis - weakfish

Cynoscionicola heteracantha

Microcotyle longicauda

Neoheterobothrium cynoscioni

Leiostomus xanthurus - spot

Heteraxinoides xanthophilis

Menticirrhus americanus - southern kingfish

Rhamnocercus stichospinus

Menticirrhus saxatilis - northern kingfish

Choricotyle louisianensis

Micropogonias undulatus - Atlantic croaker

Absonijibula lrychowskyi

Macrovalvitrematoides micropogoni

Pogonias cromis - black drum

Pauciconfibula pogoniae

Family Scombridae - mackerels

Acanthocybium solandri - wahoo

Neothoracocotyle acanthocybii

Euthynnus alletteratus - little tunny

Caballerocotyla manteri

Neohexostoma euthynni

Tristomella onchidiocotyle

Udonella caligorum

Katsuwonus pelamis - skipjack tuna

Tristomella lintoni

Sarda sarda - Atlantic bonito

Hexostoma lintoni

Scomberjaponicus - chub mackerel

Grubea cochlear

Scomber scombrus - Atlantic mackerel

Kuhnia scombri

Scomberomorus cavalla - king mackerel

Gotocotyla acanthophallus

Scomberocotyle scomberomori

Thoracocotyle crocea

Scomberomorus maculatus - Spanish mackerel

Gotocotyla acanthophallus

Pseudaxine mexicana

Scomberocotyle scomberomori

Thoracocotyle crocea

Thunnus albacares - yellowfin tuna

Nasicola klawei

Family Serranidae - sea basses

Centropristis striata - black sea bass

Microcotyle centropristis

Family Sparidae - porgies

Archosargus probatocephalus - sheepshead

Microcotyle archosargi

Monocotyle pricei ?

Stenotomus caprinus - longspine porgy

Microcotyle otrynteri

Stenotomus chrysops - scup

Microcotyle stenotomi

Family Stromateidae - butterfishes

Peprilus alepidotus - harvestfish

Microcotyle peprili

Peprilus triacanthus - butterfish

Microcotyle poronoti

Family Xiphiidae - swordfishes

Xiphias gladius - swordfish

Tristoma coccineum

Tristoma integrum

Tristomella laevis

Order PLEURONECTIFORMES

Family Bothidae - lefteye flounders

Paralichthys dentatus - summer flounder

Neoheterobothrium affine

Scophthalmus aquosus - windowpane

Bothitrema bothi

Family Pleuronectidae - righteye flounders

Hippoglossus hippoglossus - Atlantic halibut

Entobdella hippoglossi

Pleuronectes putnami - smooth flounder

Protancyrocephaloides liopsettae
Order TETRAODONTIFORMES

Family Molidae - molas

Mola mola - ocean sunfish

Capsala martinierei

Tricotyla molae

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Systematic Index

Absonifibula bychowskyi 52, 82, 90

Acanthocotyle 75, 76, 77

borealis 76

bothi 77

raiae 75

verrilli . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 76
100 NOAA Technical Report NMFS 121 _

Acanthocotylidae . . . . . . . . . . . . . . . . . . . . . . . .. 8, 76

Acanthocybium solandri 67,86,90

Acanthodiscus 79, 87

mirabilis 87

remorae 79

Acanthonchocotyle musteli 81

Acipenser 16, 40, 78, 80, 88

brevirostrum 40, 78, 80, 88

fulvescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

oxyrhynchus 78, 80

Acipenseridae . . . . . . . . . . . . . . . . . . . . . . . . . .. 80, 88

Acipenseriformes . . . . . . . . . . . . . . . . . . . . . . . . . .. 88

Aetobatis narinari 21, 76, 88

Alloheterocotyla aetobatis 76

Alopias vulpinus 75, 87

Alopiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 87

Alosa 85, 88

aestivalis 85, 88

mediocris 85, 88

pseudoharengus 85,88

Amphibdella 33, 76, 88

flavolineata 33, 76, 88

maccallumi 76

Amphibdellidae . . . . . . . . . . . . . . . . . . . . . . . . . . .. 76

Amphibdelloides 35, 76, 87

maccallumi 35, 76, 87

torpedinis 76

vallei 76

Ancyrocephalidae. . . . . . . . . . . . . . . . . . . . . . . . . .. 76

Ancyrocephalus 36, 76, 79, 89

chaetodipteri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

longiphallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79

parvus 36, 76, 89

vanbenedenii . . . . . . . . . . . . . . . . . . . . 79

Anthocotyle 54,81,88

americanus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

merluccii 54,81,88

merluccii americanus 81

Anthocotylidae 81

Archosargus probatocephalus 76, 85, 90

Aspinatrium pogoniae 86

Atheresthes stomias 78

Atheriniformes . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89

Axine 63,81,89

carangis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

gracilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81

hyporhampi 63,81, 89

Axinidae 62,81

Axinoides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

gracilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

hyporhampi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

Bairdiella chrysoura . . . . . . . . .. 48, 59, 80, 83, 84, 89

Belonidae 89

Benedenia posterocolpa . . . . . . . . . . . . . . . . . . . . . . .. 77

Benedeniella posterocolpa . . . . . . . . . . . . . . . . 11, 77, 88

Bicotylophora trachinoti 55, 83, 89

Bothidae 77,89

Bothitrema bothi 35, 77, 89

Bothitrematidae . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

Brevoortia 58, 84, 85, 88

patronus 58, 84, 85, 88

tyrannus 58, 84, 85, 88

Caballerocotyla 16, 77, 78, 90

klawei 78

manteri. . . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 77, 90

manteri affinis . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

Cadenatia polytestis . . . . . . . . . . . . . . . . . . . . . . . . . . 75

Caligus 74

praetextus 74

sp 74

Calinella myliobati 74

Capsala 13,77,78,79,91

cornuta 77

lintoni 78

maccallumi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

manteri 77

martinierei 13,77,78,91

molae 78

onchidiocotyle . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79

Capsalidae 9, 77

Capsalinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

Capsaloidea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

Capsaloides 18, 77, 89

cornutum 77

rornutus 18, 77, 89

magnaspinosus 77,89

Carangidae . . . . . . . . . . . . . . . . . . . . . . . . .. 83, 87, 89

Caranx 45,68,83,87,89

crysos 83, 89

hippos 83,87,89

latus 87

Carcharhinidae 9,26,28,43,80,87

Carcharhinus 27,74,75,80,87

commersonii 74, 75

galapagensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 74

limbatus 74, 75, 87

obscurus 27,74,75,80,88

plumbeus 81, 87

Carcharias commersonii 75

Cathariotrema selachii 19, 75, 87

Cemocotyle 68, 83, 89

carangis 83, 89

noveborancensis 68, 83, 89

Centropristis striata 86, 90

Chaetodipterus faber 38, 79, 89

Choricotyle 49, 82, 89, 90

affine 82

aspinachorda 49, 82, 89

caudalis 82

cynoscioni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 82
______________________ Hendrix: Monogenea of the Eastern United States 101

louisianensis 82, 90

merlangi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

prionoti 82

reynoldsi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

Clupeidae 57,85,88

Clupeiformes 88

Clupeocotyle 58, 84, 88

brevoortia 58, 84, 88

lintoni 58, 84, 88

Coryphaena hippurus . . . . . . . . . . . . . . . . . . . . . . . .. 78

Cyclocotyla prionoti . . . . . . . . . . . . . . . . . . . . . . . . . . 82

Cyclopteridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89

Cyclopterus lumpus 74,89

Cynoscion 32,47,71,79,82,85,86,90

arenarius 82

nebulosus 32, 79, 82, 85, 90

nothus 82, 85, 90

regalis 82, 85, 86, 90

Cynoscionicola 71, 85, 90

heteracantha 85, 90

longicauda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

pseudoheteracantha. . . . . . . . . . . . . . . . . . . 71, 85, 90

Cyprinodon variegatus 80, 89

Cyprinodontidae 30,31,80,89

Dactylocotyle 82, 83, 84

denticulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

mznor 82

phycidis 82

sp 84

trachinoti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

Dactylogyrida . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 76

Dactylogyridae 79

Dasyatidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19, 88

Dasyatis. . . . . .. 10, 20, 22, 23, 24, 26, 75, 76, 77, 88

americana 76, 88

untroura . . . . . . . . .. 10, 20, 22, 23, 75, 76, 77, 88

marmorata 75

say 26, 75, 76, 88

Dasybatotrema dasybatis 22, 75

Dermophagus squali 75

Dermophthirius 28, 74, 87

carcharhini 28, 74, 87

nigrellii 74

penneri 74

Diclidophora 46,82,84,88

affinis 82

carbonarii 82

cynoscioni. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

denticulata 46, 82, 88

lintoni 84

maccallumi 82

prionoti 82

Diclidophoridae . . . . . . . . . . . . . . . . . . . . . . . . . . .. 82

Diclidophoroides 51, 82

maccallumi 51,82

Diclodophora sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 84

Diclybothriidae . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 80

Diclybothrium armatum 40,80, 88

Diclybothroidea . . . . . . . . . . . . . . . . . . . . . . . . . . .. 80

Dionchotrema remorae . . . . . . . . . . . . . . . . . . . . . . . . 79

Dionchus 25, 79,89

agassizi 25, 79, 89

nagibinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79

remorae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79, 89

Dioncidae 79

Diplectanidae 31,79

Diplectanum 32, 79, 90

bilobatum 32, 79, 90

longiphallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

Discocotylidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 83

Dorosoma cepedianum 85, 88

Echeneidae 25, 79, 89

Echeneis naucrates 79, 89

Elasmobranchiomorphi 87

Empruthotrema raiae 19, 75, 88

Entobdella 10,77,88

brattstroemi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

bumpusii 77,88

curvunca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

hippoglossi . . . . . . . . . . . . . . . . . . . . . . . . . la, 77, 90

rosaceus 77

squamula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

steingroeveri 77

Ephippidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89

Epibdella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

bumpusii 77

hippoglossi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

Erpocotyle 43,80,81,87

abbreviata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81

maccallumi 80, 87

macrohystera 80, 87

mavori 81,89

microstoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81

sphyrnae 43,81

squali 81

Euthynnus alletteratus . . . .. 16,53,74,77,79,84,90

Exocoetidae 89

Fundulotrema prolongis 31, 80, 89

Fundulus 30, 80, 89

diaphanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

grandis 80

heteroclitus 80, 89

majalis 80, 89

Gadidae 51,81,88

Gadiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 88

Gadus morhua 74, 88

Gamacallum macroura 70, 85, 89

Gasterosteidae 30,31,80

Gastrocotylidae . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 83

Gotocotyla 64, 83, 86, 90
102 NOAA Technical Report NMFS 121 _

acanthocybii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

acanthophallus 64, 83, 90

carangis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

Grubea 56, 84, 90

cochlear 56, 84, 90

pneumatophori . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Gyrodactylida . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 80

Gyrodactylidae . . . . . . . . . . . . . . . . . . . . . . . . .. 30, 80

Gyrodactylus 31,80,89

prolongis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

stephanus 31,80,89

Haemulidae. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 82, 89

Haemulon sciurus 84

Haliotrema 39, 79, 89

mugilinus 79

vanbenedeni 39, 79, 89

Haplocleidus vanbenedeni 79

Hargisia bairdiella 59, 84, 89

Heteraxine 83

carangis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

xanthophilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 83

Heteraxinidae. . . . . . . . . . . . . . . . . . . . . . . . . .. 67, 83

Heteraxinoides 69, 83, 90

xanthophilis 69, 83, 90

xanthophiloides 83

Heterobothrium affine 82

Heterocotyle 23, 75, 76, 87, 88

aetobatis 76

floridana 76

minima 23.75,87.88

pastinacae 75

Heterocotyloides 76

diademalis 76

pricei 76

Hexabothriidae 40. 80

Hexabothrioidea . . . . . . . . . . . . . . . . . . . . . . . . . .. 80

Hexabothrium musteli 41.81,87

Hexacotyle thynni . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Hexostoma 54, 84, 90

euthynni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 84

lintoni 54, 84, 90

macracanthum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Hexostomatidae . . . . . . . . . . . . . . . . . . . . . . . .. 52, 84

Hippoglossus 10,78,90

hippoglossus 10, 78, 90

stenolepis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 78

Hyporhamphus unifasciatus 63. 81. 89

Istiophoridae 12, 78. 89

Katsuwonus pelamis. . . . . . . . . . . . . . . . . . . . . . . 78, 90

Kuhnia scombri 59, 84, 90

Labridae 89

Lamniformes 19,87

Leiostomus xanthurus 69, 83, 90

Lintonia papillosa 74

Lithidocotyle acanthophallus . . . . . . . . . . . . . . . . 83

Livoneca vulgaris 74

Loimoidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25, 75

Loimopapillosum dasyatis 26, 75, 88

Loimos 26, 75

salpinggoides 26, 75, 87

sroliodoni 75, 87

Macrovalvitrematidae 56,84

Marrovalvitrematoides micropogoni 60, 84, 90

Mazocraeidae . . . . . . . . . . . . . . . . . . . . . . . . . .. 55, 84

Mazocraeoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81

Mazocraeoides 57,85, 88

georgei 57,85,88

hargisi 85, 88

olentangiensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

similis 85

Menticirrhus 49, 80, 82, 90

americanus 80, 82, 90

littoralis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

saxatilis 49, 82, 90

Merizorotyle dasybatis. . . . . . . . . . . . . . . . . . . . . . . .. 76

Merluccius 54,81,88

albidus 81, 88

bilinearis 81, 88

Mptamirrocotyla macracantha 73, 89

Mexicotyle . . . . . . . . . . . . . . . . . . . . . . . . 83

Microbothriidae 9, 74

Microbothriidea . . . . . . . . . . . . . . . . . . . . . . . . . . .. 74

Mirrobothrium apiculatum 27, 75, 87

Mirrocotyle 72,83,85,89

acanthophallus. . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

arrhosargi 85, 89

australiensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

carangis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

cPntropristis 86, 89

debueni 86

eueides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 89

furrata 86

heteracantha. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 85

hiatulae 86, 89

longicauda 86, 89

macracantha. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

macroura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

otrynteri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 89

peprili 86, 89

pogoniap. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

pomatomi 72,86, 89

poronoti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 89

pseudohetpracantha. . . . . . . . . . . . . . . . . . . . . . . . . 85

pyragraphorus 87

scomberomori 87

stenotomi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 89

temnodontis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

Microcotylidae . . . . . . . . . . . . . . .. 68, 85

Micropogonias undulatus. . . . .. 52, 60, 80, 82, 84, 90

Mola mola 13,14,77,78,91
_______________________ Hendrix: Monogenea of the Eastern United States 103

Molidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 91

Monocotyle 24, 75, 76, 88, 90

dasybatis 75

dasybatis minimus . . . . . . . . . . . . . . . . . . . . . . . . . 75

diademalis 24, 76, 88

floridana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76

minima 75

pastinacae 75

pricei 76, 88, 90

selachii 75

Monocotylidae . . . . . . . . . . . . . . . . . . . . . . . . .. 18, 75

Monocotylidea . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 75

Monocotyloides 75

dasybatis 75

mznzmus 75

Monogenea 74

Morone 37,48,64,70,79,81,83,85,86,89

americana 37,48,79,81,83,89

saxatilis . . . . . . . . . . . . . . . .. 64, 70, 83, 85, 86, 89

Mugil 39, 73, 79, 85, 89

cephalus 39, 73, 79, 85, 89

lizia 85

Mugilidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79, 89

Mustelus canis 41,81,87

Myliobatidae . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21, 88

Nasicola klawei 15, 78, 90

Negaprion brevirostris 29, 75, 87

Neodermophthirius harkemai 29, 75, 87

Neoerpocotyle 80, 81

maccallumi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80

mavori 81

microstoma 81

Neoheterobothrium 47,82,90

affine 47,82,90

cynoscioni 82, 90

Neohexostoma euthynni 53, 84, 90

Neothoracocotyle acanthocybii 67, 86, 90

Neothoracocotylidae . . . . . . . . . . . . . . . . . . . . . . .. 86

Neothunnus macropterus . . . . . . . . . . . . . . . . . . . . .. 78

Nitzschia 16, 74, 78, 88

elegans 78

elongata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

papillosa 74

sturionis . . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 78, 88

superba. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78, 88

Nudaciraxine gracilis 64, 81, 89

Octobothrium denticulatum 82

Octoplectanum affine 82

Octostoma scombri . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Onchocleidus nactus 37, 79, 89

Onchocotyle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

abbreviata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

mavorz 81

musteli 81

Orbocotyle prionoti 50, 82, 89

Orthopristis 49, 60, 82, 84, 89

chrysoptera 49, 60, 82, 84, 89

ruber 82

Osteichthyes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 88

Papillicotyle floridana 21, 76, 88

Paragaleus gruveli . . . . . . . . . . . . . . . . . . . . . . . . . .. 80

Paralichthys 47,82,90

dentatus 47,82,90

lethostigma 82

Paramonocotyle selachii 75

Pauciconfibula pogoniae 4, 70, 86, 90

Pedocotyle 48, 83, 89

minima 83, 89

morone 48, 83, 89

Peprilus 86, 90

alepidotus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 90

triacanthus 86, 90

Percichthyidae . . . . . . . . . . . . .. 89

Perciformes 72, 89

Philura orata 75

Phylline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77, 78

bumpusii 77

sturionis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 78

Phyllonella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

bumpusii 77

hippoglossi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 77

Pleurocotyle scombri . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Pleuronectes putnami 34, 79, 90

Pleuronectidae . . . . . . . . . . . . . . . . . . . . . . . . .. 34, 90

Pleuronectiformes . . . . . . . . . . . . . . . . . . . . . .. 34, 90

Podocotyle morone. . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Pogonias cromis 70, 86, 90

Pollachius virens 46, 82, 88

Polyopisthocotylida . . . . . . . . . . . . . . . . . . . . . .. 8, 80

Pomatomidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89

Pomatomus saltatrix 64, 72, 83, 86, 89

Prionotus 50, 82, 89

carolinus 50, 82, 89

evolans 82, 89

scitulus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

Protancyrocephaloides liopsettae 34, 79, 90

Protomicrocotyle mirabilis 45, 87, 89

Protomicrocotylidae 87,89

Pseudacanthocotyla verrilli 8, 9, 74, 86

Pseudaxine 62, 83, 90

mexicana 62, 83, 90

texana 83

Pseudoaspinatrium pogoniae 86

Pseudohaliotrema 38, 79, 89

longiphallus 38, 79, 89

mugilinus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 79

Pseudomerizocotyle dasybatis 76

Pseudotagia cupida 60, 84, 89

Pterocleidus nactus. . . . . . . . . . . . . . . . . . . . . . . . . .. 79

Pungitius pungitius 80
104 NOAA Technical Report NMFS 121 _

Pyragraphoridae 61,87,89

Pyrag;raphorus pyrag;raphorus 61,87,89

Raja 19,20,42,75,76,81,88

eglanteria 75, 88

erinacea 20, 75, 76, 88

laevis 42,81,88

ocellata 75, 88

radiata 76, 88

Rajidae 8, 19, 75, 88

Rajiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 75, 88

Rajonchocotyle laevis 42, 81, 88

Remora 79,89

brachyptera 79, 89

remora 79, 89

Rhamnocercus 32, 80, 90

bairdiella 32, 80, 90

stichospinus 80, 90

Rhinoptera bonasus . . . . . . . . . . . . . . . . . . . . 11, 77, 88

Rhizoprionodon terraenovae 75, 87

Sarda sarda 54, 84, 90

Sciaenidae 31,32,80,82,89

Sciaenops ocellatus 74

Scomber 56, 59, 84, 90

japonicus 56, 84, 90

scombrus 59, 84, 90

Scomberocotyle scomberomori 65,87,90

Scomberomorus 62, 64, 83, 87, 90

cavalla 83, 87, 90

maculatus 62,83,87,90

Scombridae 12, 15, 52, 65, 66, 79, 83, 84, 90

Scophthalmus aquosus 35, 77, 90

Scorpaeniformes 89

Serranidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 90

Sparidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 90

Sphoeroides annulatus 78

Sphyrna 17,75,78,81,87

diplana 81

lewini 81

mokarran . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 81

zygaena 17,75,78,81,87

Sphyrnidae 17,43,75,81,87

Squalidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 35, 87

Squaliformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 87

Squalonchocotyle 44,80,87

abbreviata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81, 87

acanthi 81

canis 80

sphyrnae 81

squali 44,81,87

vulgaris 80

Squalus acanthias 23,35,42,44,75,76,81,87

Stenotomus 86, 90

caprinus 86, 90

chrysops 86, 90

Stromateidae. . . . . . . . . . . . . . . . . . . . . . . . . . .. 86, 90

Strongylura 36,64,77,81,89

marina 36,64,77,81,89

notata 77

Swingleus 30, 80, 89

sp 30, 80, 89

polychithroides ..' 30,89

Tagia 84

bairdiella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

cupida 84

micropogoni . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . 84

Tautoga onitis 86, 89

Tetrancistrum longiphallus 79

Tetraodontiformes. . . . . . . . . . . . . . . . . . . . . . . . .. 91

Tetrapturus albidus 18, 77, 78, 89

Tetronchus Van Benedenii 79

Thaumatocotyle 20, 76, 88

concinna 76

dasybatis 20,76,88

Thoracocotyle 66,87,90

crocea 66,87,90

paradoxica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

Thunnus 78, 90

albacares 78,90

atlanticus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

Torpedinidae 33, 76, 88

Torpedo nobiliana 33, 76, 88

Trarhinotus 55, 61, 83, 87, 89

carolinus 55, 61, 83, 87, 89

falcatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 83

rhodopus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83

Tricotyla 14,77,78,91

cutanea mediterranea . . . . . . . . . . . . . . . . . . . . . . . 78

cutanea microcotyla . . . . . . . . . . . . . . . . . . . .. 78

molae 14,77,78,91

Tricotyle scoliodoni 75

Triglidae .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 89

Trionchus dasybatis 75

Tristoma 17,77,78,87,90

coccineum 17,77,78,87,90

cornutum 77

integrum 78, 90

laeve 78

Tristomella 12,78,79,90

laeve 79

laevis 12, 90

lintoni 78, 90

onchidiocotyle 79, 90

Tristomum 78, 79

rocczneum 78

histiophori . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

laeve armata 78

onchidiocotyle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

rotundum 78

rudolphianum 78

Udonella 9, 74, 88, 89, 90
_______________________ Hendrix: Monogenea of the Eastern United States 105

caligorum 9, 75, 88, 89, 90

socialis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

Udonellida 74

Udonellidae 9, 74

Udonellidea 9, 74

Urocleidus nactus. . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

Urophycis 82, 88

chesteri 82

chuss 82, 88

regia 82, 88

Xiphias gladius 17, 78, 90

Xiphiidae 17, 78, 90
106 NOAA Technical Report NMFS 121 _

Acknowledgments _

Preparation of this manual was supported in part by a

grant from the National Science Foundation to the

Editorial Board of the "Marine Flora and Fauna of the

Eastern United States."

The author would like to acknowledge Gettysburg

College for generously awarding three Faculty Development Grants in support ofthis study. I would like to thank

Dr.J. Ralph Lichtenfels for loan of type and other monogenean specimens from the U.S. National Museum Helminth Collection. In addition I would especially like to

thank Mr. Andrew McArdle for drawing many of the

illustrations for the manual from whole mount specimens. Reviews of an early manuscript by Mary BeverleyBurton and Dennis Thoney were greatly appreciated.

Preparation of manuals in the "Marine Flora and

Fauna of the Eastern United States" subseries is coordinated by the following Board:

Coordinating Editor:

Melbourne R. Carriker, College ofMarine Studies, University of Delaware, Lewes, DE 19958

Editorial Advisors:

A. Ralph Cavaliere, Department of Biology, Gettysburg

College, Gettysburg, PA 17325.

Arthur G. Humes, Boston University Marine Program,

Marine Biological Laboratory, Woods Hole, MA

02543.

David L. Pawson, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian

Institution, Washington, D.C. 20560.

Kenneth P. Sebens, Department of Zoology, University

of Maryland, College Park, MD 20742.

Ruth D. Turner, Museum of Comparative Zoology,

Harvard University, Cambridge, MA 02138.

Robert T. Wilce, Department of Botany, University of

Massachusetts, Amherst, MA 01002.

In addition to establishing the format for the "Marine Flora and Fauna of the Eastern United States," the

Board invites systematists to collaborate in the preparation of manuals, reviews manuscripts, and advises the

Scientific Editor ofthe National Marine Fisheries Service.

Coordinating Editor's Comments

Publication ofthe "Marine Flora and Fauna ofthe Eastern

United States" is most timely in view ofthe growing universal emphasis on work in the marine environment and the

crucial need for precise and complete identification of

organisms related to this work. It is essential, if at all

possible, to accurately identifY organisms to species. Accurate scientific names of plants and animals unlock the

great quantities of biological information stored in libraries, obviate duplication of research already done, and

often make it possible to predict the attributes of organisms that have been inadequately studied.

Sherman S. Hendrix's interest in the marine environment began as he grew up near Long Island Sound

in Connecticut. After completing his undergraduate

education at Gettysburg College, he pursued interests

in parasitology in the course of obtaining a M.S. degree

at Florida State University. While at FSU, he was a

member of the scientific crew aboard the USNS Eltanin

on one of its cruises to the Antarctic Ocean. After

obtaining a Ph.D. at the University of Maryland, College Park, he returned to the study of the systematics

and ecology of marine fish parasites, particularly

aspidogastrid trematodes and Monogenea, while teaching biology at Gettysburg College.
________________________ Hendrix: Monogenea of the Eastern United States 107

Published Manuals

Marine Flora and Fauna of the Northeastern United States

Annelida: Oligochaeta David G. Cook and Ralph O. Brinkhurst

Protozoa: Ciliophora Arthur C. Borror

Higher Plants of the Marine Fringe Edwin T. Moul

Pycnogonida Lawrence R McCloskey

Crustacea: Stomatopoda Raymond B. Manning

Crustacea: Decapoda Austin B. Williams

Tardigrada Leland W. Pollock

Cnidaria: Scyphozoa Ronaldj. Larson

Higher Fungi: Ascomycetes, Deuteromycetes,

and Basidiomycetes A.R. Cavaliere

Copepoda: Harpacticoida Bruce C. Coull

Sipuncula Edward B. Cutler

Echinodermata: Holothuroidea David L. Pawson

Copepoda: Lernaeopodidae and Sphyriidae Ju-Shey Ho

Copepoda: Cyclopoids Parasitic on Fishes Ju-Shey Ho

Crustacea: Branchiura Roger F. Cressey

Protozoa: Sarcodina: Amoebae Eugene C. Bovee and Thomas K. Sawyer

Crustacea: Cumacea Les Watling

Arthropoda: Cirripedia Victor A. Zullo

Scleractinia Stephen D. Cairns

Protozoa: Sarcodina: Benthic Foraminifera Ruth Todd and Doris Low

Turbellaria: Acoela and Nemertodermatida Louise F. Bush

Lichens (Ascomycetes) of the Intertidal Region Ronald M. Taylor

Echinodermata: Echinoidea D. Keith Serafy and F. Julian Fell

Echinodermata: Crinoidea Charles G. Messing andJohn H. Dearborn

Erect Bryozoa John S. Ryland and Peterj. Hayward

Marine Flora and Fauna of the Eastern United States

Cephalopoda Michael Vecchione, Clyde F. E. Roper, and Michaelj. Sweeney

Copepoda, Cyclopoida: Archinotodelphyidae, Notodelphyidae,

and Ascidicolidae Patricia L. Dudley and Paul L. lllg

Dicyemida Robert B. Short

Platyhelminthes: Monogenea Sherman S. Hendrix

NOAA Tech. Rep.

NMFS Circular no.

374

378

384

386

387

389

394

397

398

399

403

405

406

409

413

419

423

425

438

439

440

446

NMFSno.

33

91

99

73

96

100

121

NTIS no.

COM 73 50670

73 50888

74 50019

74 50014

74 50487

74 51194

PB 257 987

261 839

268 036

268 714

273 062

274 999

280 040

281 969

222 923

285 538

296 460

297 676

124 520

225 053

219 387

124 735

PC A03/MF A01

PB 86 156 395

PB 91 173 013

PB 89 189 583

PB 91 154 179

PB 92 118 884
NOAA TECHNICAL REPORTS NMFS

The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance

and geographic distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of

these resources, and to establish levels for their optimum use. NMFS is also charged with the development and implementation of policies for managing national fishing grounds, with the development and enforcement of domestic fisheries

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international fishery agreements and policies. NMFS also assists the fishing industry through marketing services and

economic analysis programs and through mortgage insurance and vessel construction subsidies. It collects, analyzes, and

publishes statistics on various phases of the industry.

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edited by Ralph S. Svrjcek. October 1992, 143 p.

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Benway, Jack W. Jossi, Kevin P. Thomas, and Julien R.

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