NOAA Technical Report NMFS 135

Marine Flora and Fauna

of the Eastern United States

Acanthocephala

OmarM.Amin

1fD...

.. .

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

u.s. Department of Commerce

May 1998
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NOAA Technical Report NMFS 135

Marine Flora and Fauna

of the Eastern United States

Acanthocephala

OmarM.Amin

May 1998

u.s. Department of Commerce

Seattle, Washington
Foreward _

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 oforiginal, illlustrated, 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 ofthe 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 ofthe 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 shelffrom

Maine to Florida, but can vary somewhat with each major taxon and the interests of

callaborators. 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, treating a

separate major taxon, includes an introduction, glossary, uniform originally illustrated key, annotated checklist (with information, when available, on distribution,

habitat, life history, and related biology), references to the major literature of the

group, and a systematic index.

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The National Marine Fisheries Service (NMFS) does not approve.

recommend or endorse any proprietary product or proprietary

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made to NMFS, or to this publication furnished by NMFS, in

any advertising or sales promotion which would indicate or

imply that NMFS approves, recommends or endorses any

proprietary product or proprietary material mentioned herein,

or which has as its purpose an intent to cause directly or

indirectly the advertised product to be used or purchased

because of this NMFS publication.

II
Marine Flora and Fauna of the Eastern United States

Acanthocephala

OMARM.AMIN

Institute ofParasitic Diseases

P.O. Box 28372, Tempe, Arizona 85285-8372 and

Department of Zoology

Arizona State University

Tempe, Arizona 85287-1501

ABSTRACT

The phylum Acanthocephala (intestinal worm parasites of vertebrates) of the Atlantic

coast of the United States comprises 43 species and 20 genera belonging to three orders:

Echinorhynchida, Neoechinorhynchida, and Polymorphida. Adults are exclusively intestinal parasites of vertebrates. This study includes those species found in vertebrates of marine

and estuarine environments along the North American Atlantic coast between Maine and

Texas. Species that can be found within that geographical range and those that typically

infect freshwater fishes but that are occasionally present in marine or estuarine hosts are

also included. The taxonamy, anatomy, natural history, and ecology of the phylum Acanthocephala are discussed, and an illustrated key to the genera is presented. Techniques, an

annotated systematic treatment of all 43 species, and a systematic index are included. No

systematic decisions will be made at this time, but areas where such decisions are pending

will be indicated and discussed for future reports.

Introduction Taxonomy

All worms ofthe relatively small phylum Acanthocephala

(= spiny-headed) are, as adults, parasitic in the intestines of all vertebrate groups.

They are pseudocoelomates with bilateral symmetry,

have simplified anatomical structures, and all have similar reproductive processes and life cycles. Their parasitic adaptations to marine, freshwater, and terrestrial

vertebrates render them much more versatile than their

anatomical homogens may suggest. These adaptations

include reduction of the muscular, nervous, circulatory, and excretory systems, and complete loss of the

digestive system. Close phylogenetic relationships with

other invertebrate phyla are not evident. Acanthocephalans bear superficial similarities to the nematodes, and

some anatomical similarities to the cestodes can be

demonstrated. Most genera and species occur in fishes,

particularly freshwater fishes. "The absence of acanthocephalan parasites in elasmobranchs, their relatively

small numbers in marine teleosts, and their extensive

speciation in freshwater teleosts suggest a freshwater

parasitic origin after the separation of elasmobranchs

from their ancestral freshwater stock" (Amin, 1982b).

The presently accepted system for classification of the

Acanthocephala is that of Amin (1985a, 1987a) which

was previously outlined by Amin (1982a). That system,

for the most part, is based on the concepts proposed by

Meyer (1931, 1932, 1933), modified by Van Cleave

(1936), and expanded by Golvan (1959, 1960-1961,

1962, 1969) and Bullock (1969). The schemes of

Petrochenko (1956, 1958) and Yamaguti (1963) are

not recognized; see Amin (1985a, 1987a) for a detailed

discussion.

The phylum Acanthocephala is divided into four classes:

1 Class Polyacanthocephala: includes only one

monogeneric family in one order that parasitizes

fish and crocodiles in South America and Africa.

2 Class Archiacanthocephala: comprises four orders

that primarily infect predacious birds and mammals: order Apororhynchida (with one family,

Apororhynchidae, and one genus parasitizing birds

in Russia, North and South America, and Hawaii);

order Gigantorhynchida (with one family, Gigantorhynchidae, and two genera that parasitize mam-
2 NOAA Technical Report NMFS 135

mals and birds around the world); order Moniliformida (with one family, Moniliformidae, and three

genera that are widely distributed throughout the

world in mammals and occasionally birds); order

Oligacanthorhynchida (with one family, Oligacanthorhynchidae, and nine genera of worldwide distribution in birds, edentates, suids, insectivores, carnivores, simian primates, and marsupials).

3 Class Palaeacanthocephala: is the largest and most

diversified class of acanthocephalans that parasitize

fishes, amphibians, reptiles, birds, and mammals

throughout the world. The class includes two orders: Echinorhynchida and Polymorphida. Members of the order Echinorhynchida are parasites of

fishes, and occasionally amphibians and reptiles,

and are placed in 10 families (Arhythmacanthidae

with six genera in three subfamilies; Cavisomidae

with 10 genera; Diplosentidae with four genera in

two subfamilies; Echinorhynchidae with six genera

in two subfamilies; Fessisentidae with one genus;

Heteracanthocephalidae with four genera in two

subfamilies; Hypoechinorhynchidae with two genera; Illiosentidae with 10 genera; Pomphorhynchidae

with four genera; Rhadinorhynchidae with 22 genera in five subfamilies). Members of the order

Polymorphida are parasites of amphibians, reptiles,

birds, and mammals. The order includes three families (Centrorhynchidae with two genera; Plagiorhynchidae with nine genera in three subfamilies; and

Polymorphidae with nine genera).

4 Class Eoacanthocephala: includes parasites offishes

and less frequently parasites of amphibians and reptiles. It consists of two orders: Order Gyracanthocephala (with eight genera in one family,

Quadrigyridae, and two subfamilies) and order

Neoechinorhynchida with three families (Dendronucleatidae with one genus; Neoechinorhynchidae

with 18 genera in three subfamilies; and Tenuisentidae with two genera).

The acanthocephalan species treated in this manual

are known from fishes, mammals, and birds that are

naturally found in marine as well as estuarine habitats.

Freshwater acanthocephalans that have been reported

occasionally from fish hosts in marine or estuarine locations within the geographical range covered by this

manual are also included. Marine acanthocephalans

reported outside ofthis range, but that may prove to be

present within it, are also discussed. A total of 43 species in 20 genera of acanthocephalans are treated in

this manual in the following sections. The genera belong to three orders as follows:

Order Echinorhynchida: Acanthocephalus Koelreuther,

1771; Dollfusentis Golvan, 1969; Echinorhynchus Zoega in

Muller, 1776; Filisoma Van Cleave, 1928; Gorgorhynchoides

Cable and Linderoth, 1963; Gorgorhynchus Chandler,

1934; Leptorhynchoides Kostylev, 1924; Pomphorhynchus

Monticelli, 1905; Serrasentis Van Cleave, 1923;

Tegorhynchus Van Cleave, 1921.

Order Neoechinorhynchida: AtactorhynchusChandler,

1935; Floridosentis Ward, 1953; Neoechinorhynchus Stiles

and Hassall, 1905; Paratenuisentis Bullock and Samuel,

1975.

Order Polymorphida: Andracantha Schmidt, 1975;

Arhythmorhynchus Luhe, 1911; Bolbosoma Porta, 1908;

Corynosoma Luhe, 1904; Polymorphus Southwellina

Witenberg, 1932.

Anatomy of Adults _

Like cestodes, adult acanthocephalans are obligate intestinal parasites that lack a digestive system. They are

dioecious and females are usually larger than males of

the same species. The body consists of the proboscis,

neck, and trunk. The proboscis may be short and stubby

with few hooks or cylindrical with many hooks. Except

for one order, Apororhynchida, the proboscis is armed

with recurved sclerotized hooks and is retractable into

a receptacle that houses the "brain" (cerebral ganglion).

Proboscis hooks are often arranged in longitudinal

rows and may vary in shape, size, and root structure

anterioposteriorly or dorsoventrally. The growth pattern of proboscis hooks and the relationship of this

growth pattern to acanthocephalan establishment and

attachment to host intestinal mucosa have been studied

in a few species (e.g. see Amin, 1986b, 1987c). Proboscis hooks are usually rooted, and these roots are often

simple and directed posteriorly; many variations exist.

An apical organ at the anterior tip ofthe proboscis of a

few species, which may have a sensory or secretory

function, has been described.

The neck is the unarmed area directly following the

posteriormost circle of hooks on the proboscis. It may

be practically nonexistent or very long, well-developed,

and capable of of being enlarged into a bulb at some

point in its length, as in the palaeacanthocephalan

family Pomphorhynchidae. The proboscis receptacle

may be single-walled (e.g. in Eoacanthocephala and

some archiacanthophalans) or double-walled. The pattern of proboscis receptacle musculature is variable

and may assume spiral or diagonal arrangement; see,

for example, the archiacanthocephalan order Moniliformida. Some palaeacanthocephalans (e.g. family

Fessisentidae) have a nuclear pouch at the posterior

extremity of the proboscis receptacle,

The central structure of the nervous system is the

"brain" (cerebral ganglion) which is invariably present

within the proboscis receptacle at various levels in different families and genera. The ganglia are associated
________________________ Amin: Acanthocephala of the Eastern United States 3

with the peripheral nervous system which is more complex in male than female worms. The genital and bursar ganglia are found at the posterior end of males and

are associated with reproductive activity. The sensory

receptor system includes the apical structure which is

located at the anterior tip of the proboscis of some

acanthocephalans and may vary from a simple papilla

to an extrudable depression surrounded by specialized

cells and ducts. Sensory support cells and sensory pits

also have been described from a few acanthocephalan

species.

The trunk is usually cylindrical but may (rarely) be

ribbonlike or have superficial annulations. It is often

unarmed (spineless) but may be armed with spines of

many shapes and sizes that are variously distributed.

Most adults vary from a few mm to 1 or 2 cm in length.

However, some oligacanthorhynchid archiacanthocephalans, e.g. Macracanthorhynchus hirudinaceus from

pigs and Oligacanthorhynchus longissimus from aardvarks

may reach 45 and 93 cm in length, respectively. Absorption of nutrients occurs through the body wall which

contains a lacunar (circulatory) system with lateral longitudinal canals (in Palaeacanthocephala) or dorsoventral longitudinal canals (in other classes). The body

wall contains hypodermal nuclei that may fragment in

adults. Giant nuclei, at least in the genus Neoechinorhynchus, appear to be associated with the reproductive

activity of worms (Amin and Vignieri, 1986a, 1986b;

Amin and Gunset, 1992). The tegument of adult acanthocephalans includes the following layers from the

surface: glycocalyx; outer limiting membrane; striped,

vesicular, felt, and outer radial layers; inner membrane;

and connective layer. A detailed discussion of the functional morphology of the various anatomical structures

and organ systems of the Acanthocephala is presented

by Miller and Dunagan (1985).

A pair of lemnisci originate at the base of the neck

and are believed to have absorptive function comparable to that of the body wall. They may be equal or

unequal in length, short or very long, nucleated or

nonnucleated, and variously shaped.

Excretion and osmoregulation are accomplished by

diffusion in all acanthocephalans, except in members

of Oligacanthorhynchidae, which have two specialized

protonephridial organs. Two types of protonephridia

are known: dendritic or capsular saclike.

The reproductive system in both sexes is suspended

in the body cavity by one or two long hollow ligament

sacs that extend between the base of the proboscis

receptacle to near the distal genital pore. The ligament

sacs mayor may not persist in adults. The female reproductive system consists of gonads from which ovarian

balls develop and, after fertilization, produce oocytes

and eggs. In a few precocious species, the ovarian balls

may begin to develop in the last larval stage (the

cystacanth) while still in the body cavity of the intermediate host (see Amin, 1982b). The efferent duct of the

female system includes the uterine bell, uterus, and

vagina. The anterior bell chamber opens posteriorly

into the selector apparatus which allows the passage of

only fertilized, ripe eggs through the system. Rejected

eggs may be returned to the pseudocoelom through a

ventral bell opening. Entrance to the variably long uterus

is controlled by the selector apparatus and by a posterior vaginal sphincter. The vagina opens into the genital pore which may be terminal or subterminal and

which may occasionally be bordered or surrounded by

accessory structures, such as lips, papillae, or spines.

The male reproductive system usually includes two

testes; species with a single testis are also known, e.g.

those in the echinorhynchid family Fessisentidae. The

testes are often spheroid-ovoid and arranged in tandem but may also be long and tubular. Monorchid

males have been described in a number ofspecies (see

Amin, 1975a, 1989; Bullock, 1962). The sperm ducts

may be enlarged in some species to form a seminal

vesicle. There are usually four, six, or eight cement

glands that may be ovoid, pyriform, elongate, tubular,

or intestiniform. They are grouped in three categories:

1) a single syncytial structure with giant nuclei

(Eoacanthocephala); 2) up to eight pyriform glands

each with a giant nucleus (Archiacanthocephala); or 3)

spheroid or tubular glands with fragmented nuclei

(Palaeacanthocephala). Paired cement ducts originate

from the cement glands and occasionally fuse to form

the cement reservoir which opens into the common

genital canal. Cement gland secretions ultimately form

the copulatory cap that plugs the female gonopore

after copulation. Saefftigen's pouch, the eversible copulatory bursa, and the penis make up the copulatory

structures of male acanthocephalans.

The taxonomic treatment and key included in the

following sections are based on adult anatomical features outlined above. For a discussion of the anatomy

and development oflarval stages (acanthor, acanthella,

cystacanth), see Amin (1982b) and Schmidt (1985).

The broad range of morphological variability and

anomalies in attachment and reproductive structures,

among others, have been reported infrequently (see

Bullock, 1962; Amin, 1975a, 1989).

Natural History

Only adult forms of acanthocephalans inhabit the intestinal tract of all vertebrate groups. They usually live

less than one year in their definitive hosts where they

often exhibit seasonality in numbers and maturity. In

riverine species, new generations are often recruited by

hosts in the autumn after ingestion of crustacean inter-
4 NOAA Technical Report NMFS 135

mediate hosts parasitized by cystacanths. Young recruits

undergo growth and development and reach peak size

and maturity in the spring. Migration to more posterior

intestinal locations has been associated with seasonal

development and maturation in many riverine species.

The short-lived males are less common in spring collections. Reproductive activity increases in winter and

spring as evidenced by greater numbers offemales with

eggs and copulatory caps (cement plugs) during these

seasons. In freshwater acanthocephalans whose hosts

inhabit large and deep oligotrophic lakes, seasonality

in recruitment, development, maturity, and numbers

are not evident (see Amin and Burrows [1977] and

other references therein). The absence of cyclic fluctuations is related to the lack of pronounced seasonal

temperature oscillations that are known to affect recruitment, maturation, and seasonal availability of intermediate hosts. These seasonal aspects of marine acanthocephalans have been rarely investigated and reported.

Eggs are passed with the definitive host's feces and

remain free in the environment until ingested by an

arthropod intermediate host. The eggs of aquatic

acanthocephalns are usually oval to oblong and show a

polar prolongation of the middle membrane. Those of

terrestial worms are typically oval, thick-shelled, and

show no prolongation ofthe middle membrane. Aquatic

intermediate hosts usually consist of Amphipoda,

Isopoda, Copepoda, and Ostracoda, whereas terrestial

hosts include insects such as Orthoptera and Coleoptera.

Ingested eggs (shelled acanthors) develop into young

and (later) older acanthellae in the body cavity of the

arthropod intermediate host, where they later develop

to the larger infective cystacanth stage. In some acanthocephalan species, larval forms induce alteration in

the behavior and/or coloration of the in termediate

host (in contrast with noninfected arthropods), making the intermediate host much more readily captured

and eaten by the definitive host. Adult breeding and

egg production appear to be timed to make eggs available to the intermediate host when its populations are

seasonally available and well established. This seasonality is often evident in late spring and in summer in

riverine species (see Amin et aI., 1980).

Aspects of life history outlined above are universal

and are shared by other known acanthocephalan species. In a few species, however, an additional paratenic

(transport) host occupies a position between the arthropod intermediate host and the vertebrate definitive host (in which acanthocephalans reach sexual maturity and breed). Paratenic hosts become ecologically

indispensible for the completion of the life cycle of

these acanthocephalans whose hosts have no direct

environmental contact with intermediate hosts. Less

frequently, a definitive host may become a paratenic

host if the infected intermediate host contained larvae

that had not yet reached the infective cystacanth stage.

In paratenic hosts, immature worms are usually encysted in or on body cavity organs. Transmission of

adults from one definitive host to another through

predation also is known to occur.

Techniques _

The following methods of processing acanthocephalans for whole mounting were originally adapted from

Bullockl :

1 Remove adults carefully from host intestinal mucosa without injuring or losing any of the parts,

particularly the proboscis, which may need to be

removed separately. Record worm distribution, pathology, and related observations. Cut intestinal

sections with intact worms and clear pathology and

preserve in 4% formaldehyde (10% formalin) for

future histopathological studies.

2 Refrigerate freshly recovered specimens overnight

to extend the proboscis, then fix in refrigerated

AFA (alcohol-formalin-acetic acid) or in 75% ethanol if AFA is not available.

3 Prick specimens individually with a fine needle such

as an insect mounting pin. Prick large and thick

specimens more than once. Do not prick through

reproductive or attachment structures.

4 Stain in Mayer's acid carmine overnight.

5 Destain next morning with 4% HCI in 70% ethanol

until pink.

6 Rinse three times in 75% ethanol or until no more

stain leaches out ofspecimen.

7 Place 12 to 24 h in 85% ethanol.

8 Place 12 to 24 h in 95% ethanol.

9 Repeat step 8.

10 Place 12 to 24 h in 100% ethanol; longer in large

worms.

II Place 12 to 24 h in 25% terpineol/75% absolute

ethanol (by volume).

12 Place 12 to 24 h in 50% terpineol/50% absolute

ethanol (by volume).

13 Place 12 to 24 h in 75% terpineol/25% absolute

ethanol (by volume).

14 Place 12 to 24 h in 100% terpineol.

15 Place 12 to 24 h in pure terpineol mixed with

Canada balsam in equal proportions by volume.

Alternative clearing agents: xylene or oil or winter

green.

16 Whole mount in Canada balsam.

I Bullock, W. L. 1973. 13 Thompson Lane, Durham, H 03824.

Personal. commun.
_________________________ Amin: Acanthocephala of the Eastern United States 5

Important notes: Specimens should not be removed

from vials or taken out of solution. Change fluids only

by withdrawing or adding with uncontaminated Pasteur pipettes. After use, discard all alcohol concentrations. All fluids containing terpineol are reusable. Do

not leave solution bottles or specimen vials open longer

than absolutely necessary. Air moisture may cause total

and irreversible opaqueness in specimens when

mounted in balsam. In such cases, returning specimens

to the 100% ethanol stage and repeating subsequent

steps may save the day.
6 NOAA Technical Report NMFS 135

Key to Genera _

The following key includes mostly marine genera known

from marine fish and mammals. The key also includes the

following categories of acanthocephalans: 1) genera reported from birds captured in Atlantic coastal areas or

whose intermediate hosts comprise marine invertebrates;

2) genera utilizing marine invertebrates as paratenic hosts;

3) marine genera (determined from host distribution) from

waters of neighboring countries adjacent to the study area

covered by this manual; and 4) freshwater genera with

occasional records in marine or estuarine hosts. The key is

based on the concepts reported by Amin (l987a). Most

figures were made with the aid of original descriptions.

la Main longitudinal lacunar canals lateral. Lemniscal, cement gland, and hypodermal nuclei fragmented. Ligament sacs in females single, not persistent. Proboscis cylindrical to spheroid usually with

many hooks in alternating longitudinal rows (Figs. 1, 2). Proboscis receptacle double-walled

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Class Palaeacanthocephala 2

Ib Main longitudinal lacunar canals median. Lemniscal, cement gland, and hypodermal nuclei gigantic.

Ligament sacs in females double, persistent. Proboscis usually small with few radially arranged hooks

(Fig. 3) Proboscis receptacle single-walled Class Eoacanthocephala, Order Neoechinorhynchida 8

jYUA~". . ' . .. .

)~ V

3

:ffi-\,--- Lemniscus

--+f--- Genital ligament

H-f---Uterine bell

IfI--t,l----- Uterus

liH~--Proboscis receptacle

:;J;I----- Vagina

~/~J!I-3"'-1'\-- Brain

#---Ir+-- Retractor muscles

i

2

--++--- Testis

'-"t-ff--- Cement gland

~-- Bursa

Ht--- Saefftigen's pouch

----Proboscis

---¥;~r--- Proboscis receptacle

IfX.."I-Ttl--- Lemniscus

~.~\\t\---Brain

//r-~I+--Genital ligament

IHIJ'"+---tt/-- Retractor muscles

Figures 1-3

(I) Adult male Acanthocephalus dirus from New Hampshire. (2) Adult female Acanthocephalus dirus from New Hampshire. (3) Proboscis of Neoechinorhynchus cylindratus.
_________________________ Amin: Acanthocephala of the Eastern United States 7

2 Parasites of fishes and amphibians

Parasites of reptiles (rarely), birds, and mammals

Order Echinorhynchida

Order Polymorphida, Family Polymorphidae

3

15

3a Neck very long and cylindrical or spirally twisted with or without bulbous swelling. Proboscis receptacle

long and inserted at base of proboscis (Fig. 4). Parasites of freshwater and marine fishes

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Family Pomphorhynchidae, Genus Pomphorhynchus

3b eck and proboscis receptacle not long (Figs. 1-3) 4

4a Trunk usually unarmed (Figs. 1,2); armed individuals with anterior minute cuticular spines 5

4b Trunk usually armed (Figs. 5-7); cuticular spines may rarely be replaced by sclerotized plaques or

absent. Unarmed worms with dendritic hypodermal nuclei, subcylindrical proboscis, long proboscis

receptacle with anterior brain, and 6-8 compact cement glands. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6

4

7

Figures 4-7

(4) Anterior trunk, neck, and proboscis of Pomphorhynchus bulbocolli. (5) Anterior trunk of Tegorhynchus. (6) Young

male Serrasentis sagittifer. (7) Anterior trunk and proboscis of Gorgorhynchus medius.
8 NOAA Technical Report NMFS 135

5a Proboscis cylindrical with many hooks or spheroid with few hooks. Cement glands 6 or 8, usually

pyriform to spherical, and compact (Fig. 1). Parasites of freshwater or marine fishes, and occasionally

amphibians Family Echinorhynchidae, Subfamily Echinorhynchinae 9

5b Proboscis claviform to long and slender. Cement glands 4, elongate, and tubular, or filiform (Fig. 8).

Parasites of freshwater and marine fishes Family Cavisomidae, Genus Filisoma

6a Trunk spined in one undivided region anteriorly (Fig. 5) and often at posterior extremity. Cement

glands 8, elongate pyriform. Parasites of marine and freshwater fishes Family Illiosentidae 10

6b Trunk usually spined differently (Figs. 6-7), rarely unspined. Cement glands 4 or 6, elongate and

tubular or short and pyriform; 6 or 8 in some unspined species. Parasites of marine and freshwater

fishes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family Rhadinorhynchidae 7

7a Trunk with many ventral-transverse combs ofspines behind anterior collar of spines (Fig. 6)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Subfamily Serrasentinae, Genus Serrasentis

7b Trunk spines in 1 or 2 regions anteriorly (Fig. 7), or infrequently absent or replaced by sclerotized

plaques Subfamily Gorgorhynchinae 11

8a Proboscis short and subglobular or subcylindrical, armed with a small number of hooks arranged in

spiral, circular, or diagonal rows (Fig. 3). Parasites of fishes and amphibians

.............................................................. Family Neoechinorhynchidae 13

8b Proboscis relatively long and cylindrical, armed with many hooks arranged quincuncially in longitudinal rows (Fig. 9). Parasites of fishes Family Tenuisentidae Genus Paratenuisentis

9a "Brain" at posterior end of proboscis receptacle (Figs. 1, 2) . . . . . . . . . . . . . . . . . . . .. Genus Acanthocephalus

9b "Brain" near middle of proboscis receptacle Genus Echinorhynchus

9

Figures 8 and 9

(8) Tubular cement glands and posterior end of

Filisoma. (9) Proboscis of Paratenuisentis ambiguus.
________________________ Amin: Acanthocephala of the Eastern United States 9

lOa Proboscis ensheathed in a conspicious hyaline membrane (Fig. 10) Genus Tegorhynchus

lOb Proboscis without a hyaline membrane (Fig. 11) Genus Dollfusentis

11a Trunk with dorsal bulboid swelling anteriorly (Fig. 12) Genus Gorgorhynchoides

Trunk without bulboid swelling. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12

l2a Trunk armed anteriorly with irregularly arranged spines in pits (Fig. 7) Genus Gorgorhynchus

l2b Trunk unarmed Genus Leptorhynchoides

l3a Proboscis with 3 circles of hooks each with 6 hook (Fig.3) . . . . . . . . . . . . . . . . . . .. Genus Neoechinorhynchus

l3b Proboscis with more hooks (Figs. 13, 14) 14

10

13

I )1 ;! 'I)

Figures 10-14

(10) Proboscis hooks and hyaline sheath of Tegorhynchus. (11) Anterior trunk and proboscis of Dollfusentis

chandleri. (12) Anterior trunk and proboscis of Corgorhynchoides elongatus. (13) Proboscis of Floridosentis elongatus.

(14) Proboscis of Atactorhynchus verecundus.
10 NOAA Technical Report NMFS 135

I4a Proboscis with 8 diagonal rows of about 7 hooks each (Fig. 13) . . . . . . . . . . . . . . . . . . . .. Genus Floridosentis

14h Proboscis with 8 diagonal rows of about 8 hooks each anteriorly and about 16 smaller hooks posteriorly

(Fig. 14) Genus Atactorhynchus

15a Posterior trunk with genital spines in at least one sex (Fig. 15) . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

I5h No genital spines in either sex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

16a Swollen anterior trunk with 2 fields of spines (Fig. 16) Genus Andracantha

16h Swollen anterior trunk with 1 field of spines (Fig.17) Genus Corynosoma

17a Anterior trunk greatly swollen and with 1 field of spines, separated from rest of trunk by narrow

constriction (Fig. 18) Genus Bolbosoma

17h Anterior and posterior portions of trunk not separated by constriction, with 1 or 2 fields of trunk

spines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

17

Figures 15-19

(15) Posterior end of Andracantha mergi showing genital spine. (16) Female Andracantha gravida. (17) A female

Corynosoma sp. (18) Anterior trunk and proboscis of Bolbosoma turbinella.
______________________ Amin: Acanthocephala of the Eastern United States II

18a Two fields of trunk spines in at least one sex (Fig. 19); trunk somewhat short. Four cement glands

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Genus Southwellina

18b One field of trunk spines (Fig. 18); trunk variable in length, cement glands 2 or 4 19

19a Trunk extremely long; 2 or 4 cement glands. Middle of proboscis usually enlarged with greatly

enlarged ventral hooks (Fig. 20) Genus Arhythmorhynchus

19b Trunk often short; 4 cement glands. Proboscis cylindrical, ovoid, spindle-, or pear-shaped with no

dorsa-ventral differentiation or abrupt extreme enlargement of hooks (Figs. 21, 22)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Genus Polymorphus

(~~ i¥'rf1( 21 19 ~AII(,\ hl 1/\ i\

~'V~

~~~U18.

Jr~~ j%6,} P 'I '" J P

" V \~

r~~ ~~* ~f~jl

~g~~ ~u~~ 20 22

Figures 20-22

(19) Female Southwellina dimorpha. (20) Proboscis of Arhythmorhynchus uncinatus. (21) Proboscis of Polymorphus

(Polymorphus) brevis. (22) Proboscis of Polymorphus (Profilicollis) altmani.
12 NOAA Technical Report NMFS 135

Annotated Systematic List

The following list of primarily marine acanthocephalans from the eastern seaboard of the United States is

arranged systematically in two palaeacanthocephalan

orders (Echinorhynchida [with five families] and Polymorphida [with one family]), and one eoacan thocephalan order (Neoechinorhynchida [with two families]) according to the classification system of Amin

(1982a, 1985a). The list includes 43 species in 20 genera, most of which are distributed in marine or estuarine waters along the Atlantic coast ofthe United States.

Orders, families, genera, and species are listed alphabetically under classes Palaeacanthocephala and

Eoacanthocephala, irrespective of their marine affinities,

hosts, or distribution. Available type and voucher material

of the listed species are obtainable from the Biosystematics and National Parasite Collection unit of the Agricultural Research Service, USDA, Beltsville, MD. (J. Ralph

Lichtenfels, curator) and from the Harold W. Manter

Laboratory, University of Nebraska State Museum (Mary

H. Pritchard, curator). Host and distributional data from

museum records have been useful in text treatment.

The following text emphasizes host records and geographical distribution. Systematic notes will also be included when relevant but no systematic decisions will

be made at this time. Areas where such decisions need

to be made will, however, be pointed out for future

studies (for synonyms, see Amin, 1985a).

All echinorhynchid and neoechinorhynchid acanthocephalans discussed below are known from marine and

estuarine fishes and occasionally from freshwater fishes

collected in Atlantic coastal streams ofNorth America. All

polymorphid acanthocephalans were taken from birds

captured over, in, or near the same geographical regions.

Class PALAEACANTHOCEPHALA Meyer, 1931.

Order ECHINORHYNCHIDA Southwell and McFie,

1935.

Family CAVISOMIDAE Meyer, 1932.

Filisoma fidum Van Cleave and Manter, 1947, was described from Bermuda chub, Kyphosus sectatnx, in the

Dry Tortugas, Florida. The host genus, Kyphosus, is represented on the Atlantic and Pacific coasts of North

America by distinct species that might have gradually

diversified from common ancestors during isolation

(Van Cleave and Manter, 1947). Eight species of the

genus FilisomaVan Cleave, 1928, are currently recognized

and a key distinguishing F. fidum from other species in the

genus is included in Amin and Nahhas (1994).

Family ECHINORHYNCHIDAE Cobbold, 1876.

The genera AcanthocephalusKoelreuther, 1771, and Echinorhynchus Zoega in Muller, 1776, are two of the largest

echinorhynchid genera with over 45 and 41 species,

respectively, reported from freshwater and marine fishes

and occasionally from amphibians (Amin, 1982a,

1985a).

Acanthocephalus dims (Van Cleave, 1931) Van Cleave

and Townsend, 1936, is a freshwater species that has

been redescribed and documented from 65 species and

16 families of fishes (Amin, 1984). It has three distinct

populations: 1) in the Mississippi River drainage system; 2) Wisconsin and Lake Michigan; and 3) the New

England states (Amin, 1985b, 1986a; Amin and

Huffman, 1984). Its New England population has been

reported from 14 species and 9 families offishes including Atlantic tomcod, Microgadus tomcod, and American

eel, Anguilla rostrata, in the Oyster River, a New Hampshire estuary, downstream from the dam separating it

from a freshwater Mill Pond (Bullock, 1962; Bullock2).

Host and geographical distribution of adults depends

on the extent of dispersal of its intermediate host(s)

(Amin, 1985b). The intermediate hosts of Acanthocephalus dirus are the isopod Asellus intermedius (= Caecidotea militans in Wisconsin) in riverine situations and

the amphipod Pontoporeia aJfinis in Lake Michigan

(Amin, 1978). The ecology of A. dirus has been studied

in various parts of its range by a number of authors

including Seidenberg (1973), Amin (1975b), and Camp

and Huizinga (1980), and its larval development by

Amin (1982b).

Echinorhynchus gadi Zoega in Miiller, 1776. Since its

original description, E. gadi has been reported from a

considerable number of marine and estuarine fish species from Russia; Barents, White, Baltic, and North seas;

Gulf ofFinland; Scandanavian waters; Sea ofJapan; and

the Atlantic coast of North America. The large number

of host species (see Petrochenko, 1956; Yamaguti, 1963;

Valtonen and Crompton, 1990, for partial lists) from

diverse families and their wide range of geographical

distribution clearly reflects the great degree of morphological variability in this species that is perhaps one

of the widest among acanthocephalan species. This

variation has contributed to a long list of synonymies

describing different populations of E. gadi as new species. Van Cleave recognized this fact and cautioned that

"Failure to recognize the extreme plasticity of this form

has caused many workers to describe as new species

forms which on the basis of essential characteristics are

wholly indistinguishable from one another." At least 12

synonyms are listed in synonymies (Amin, 1985a). In

addition, the subspecies callanae Viborg, 1795, and

virentis Rathke, 1799, are regarded as E. gadi; E. gadi

2 Bullock, W. L. 1994. 13 Thompson Lane, Durham, NH 03824.

Personal commun.
________________________ Amin: Acanthocephala of the Eastern United States 13

sensu Van Cleave (1924a) (= E. vancleavei Golvan, 1969);

E. gadi sensu Yamaguti, 1935 (= E. yamagutii Golvan,

1969); andE. gadisensu Southwell and MacFie, 1925 (=

another species).

Whether E. gadi is a species complex is an often asked

question. Huffman and Bullock (1975) distinguished

with meristogram analysis two "populations" of E. gadi

obtained from two distinct sets of hosts from the American Atlantic coast. Amin and Redlin (1980) and Shostak

et al. (1986) demonstrated that worm age, host species,

and geographical location contribute to the morphological variability in taxonomically important characters. The latter authors further showed that meristogram

analysis alone is not sufficient to delineate taxa and

hypothesized that "geographical variability may be a

consequence of restricted gene flow." The taxonomic

identity of E. gadi is too complex to be realistically

resolved within the scope of the present study.

In addition to the definitive hosts (including those

from North American Atlantic coast fishes) listed in

Petrochenko (1956) and Yamaguti (1963), see lists included in Huffman and Bullock (1975), Meyers (1978),

Shostak et al. (1986), Gartner and Zwerner (1989),

Arai (1989), Landry et al. (1992), Linton,3 and museum records. The fish definitive hosts from the Atlantic coast of North America (New Brunswick-Canada,

Maine, New Hampshire, New Jersey, Virginia), drawn

in part from above sources, include: Acipenseroxyrhynchus,

Alosa aestivalis, A. pseudoharengus, A. sapidissima,

Ammodytes hexapterus, Anarhichas lupus, Clupea harengus,

Coregonus clupeaformis, Coryphaenoides rupestris, Cymatogaster aggregate, Gadus macrocephalus, G. morhua, G.

ogac, Glyptocephalus cynoglossus, Hemilepidotus hemilepidotus,

Hemitripterus americanus, Hippoglossoides platessoides,

Hippoglossus hippoglossus, Icelinusfilamentosus, Leptocottus

armatus, Liparis atlanticus, Lycodes cortezianus, Mallotus

villosus, Macrourus berglax, Melanogrammus aeglefinus,

Merluccius productus, Microgadus proximus, M. tomcod,

Morone saxatilis, Myoxocephalus octodecemspinosus, M.

polycanthocephalus, M. quadricomis, Nemichthys scolopaceus,

Nnumia bairdi, Oncorhynchus gorbuscha, O. keta, O. kisutch,

0. nerka, O. tshawytscha, Paralichthys dentatus, Platichthys

stellatus, Pleuronectes americanus, P. ferrugineus, Pleuronectes

putnami, Podothecus acipenserinus, Pollachius virens, Raja

radiata, Reinhardtius hippoglossoides, Salmo salar, Salvelinus

alpinus, S. fontinalis, S. malma, S. namaycush, Scophthalmus

aquosus, Sebastes aleutianus, S. alutus, S. babcocki, S. borealis, S. caurinus, S. diploproa, S. Jlavidus, S. helvomaculatus,

S. proriger, S. ruberrimus, S. zacentrus, Tautoglabrus

adspersus, Theragra chalcogramma, Urophycis chuss, U. tenuis.

3 Linton, E. 1933. otes on Acanthocephala mainly from fishes

and birds of the Woods Hole region; submitted to Proc. U.S. Natl.

Mus. Wash., D.C., 81 p., 11 pI. Unpubl. manuscr.

The intermediate hosts of E. gadi are amphipods of

the genera Aeginia, Amphithoe, Calliopius, Caprella,

Cyphocaris, Gamarellus, Gammarus, and Pontoporeia, and

the mysid, Mysis. Intermediate hosts known from the

North American Atlantic Coast include Aeginina longicomis, Gamarellus angulosus, Calliopius laeviusculus (Marcogliese (1994); Bullock2).

Echinorhynchus laurentianus Ronald, 1957, was reported

in the Gulf ofSt. Lawrence from marine Heterostomata.

These hosts include Glyptocephalus cynoglossus, Hippoglossoides platessoides, Hippoglossus hippoglossus, Pleuronectes

americanus, P. ferrugineus, P. putnami, Reinhardtius

hippoglossoides, Scophthalmus aquosus, and various species

of flounders (see Arai, 1989, for a review of these

records). Many of these fish hosts also are found in the

northern U.S. Atlantic waters. This common distribution probably encompasses shared parasite faunas including E. laurentianus.

Family ILLIOSENTIDAE Golvan, 1960.

Genus dollfusentis Golvan, 1969, was erected to accommodate a species originally described by Linton (1905)

and to relegate three illiosentid species described by

Cable and Linderoth (1963) to it. All four species are

included in the present review.

Dollfusentis chandleri Golvan, 1969 (= Telosentis tenuicomis

(Linton, 1905) Van Cleave, 1918) has the widest range

of host and geographical distribution of all species of

Dollfusentis. Chandler (1934) provided a detailed

emended description of the species from Galveston

Bay, Texas. It was later redescribed by Salgado-Maldonado (1976) who also described the fifth species of

the genus, Dollfusentis bravoaeSalgado-Maldonado, 1976,

from Cozumel, Quin tana, Mexico. This species has been

reported from many marine and estuarine localities

north and south of the Gulf of Mexico. Southernmost

locations extend into Brazil from Haemulon sciurus (see

Kohn and Macedo, 1984). Its North American range

includes the Atlantic Coast of Massachusetts, Maryland,

Virginia, orth Carolina, Florida, Mississippi, Louisiana, Alabama, and Texas.

Definitive hosts from the above locations include

Acipenser oxyrhynchus, Archosargus probatocephalus,

Bairdiella chrysura, Cynoscion arenarius, C. nebulosus, C.

regalis, Eucinostomus argenteus, Larimusfasciatus, Leiostomas

xanthurus, Lobotes surinamensis, Micropogonias undulatus,

Monacanthus hispidus, Morone americana, M. saxatilis,

Orthopristis chrysoptera, Paralichthys dentatus, Polydactylus

octonemus, Tylosurus acus and T. carribaeus. Sources for

the above geographical and host records include Chandler (1934, 1935), Van Cleave (1947), Bullock (1957,

1960), Huizinga and Haley (1962), Yamaguti (1963),
14 NOAA Technical Report NMFS 135

Bullock and Mateo (1970), Williams and Gains (1974),

Buckner et aI. (1978), Overstreet (1978), Jansen and

Burreson (1990), Thoney (1991), Linton,3 and museum records.

O'Rourk (1949), Sogandares-Bernal (1955), Bullock

(1957), and Huizinga and Haley (1962) studied

prevalences of D. chandleri in North Carolina, Louisiana, Texas, and Maryland coastal waters. Amphipod

intermediate hosts include Lepidactylussp., Grandidierella

bonnieroides, and Corophium lacustra. These hosts were

reported from high and low salinity waters of the coast

of Alabama, Mississippi, and Louisiana by Buckner et

aI. (1978) and Overstreet (1978). Amphipod hosts are

usually eaten directly by fish definitive hosts, and

paratenic transport hosts are not involved in the life

cycle of this acanthocephalan.

Dollfusentis centmrhynchus (Cable and Linerotb, 1963)

Golvan, 1969, was originally described from Mulloidichthys martinicus in Jamaica by Cable and Linderoth

(1963) and later reported from Florida by Bullock and

Mateo (1970). No other records of this acan thocephalan were published since.

Dollfusentis heteraoonthus (Cable and Linderotb, 1963)

Golvan, 1969, was originally described from Platophrys

ocellatus, Gerres cinereus, Bathygobius soporator, and

Labrisomus nuchipinnis in Cura~ao by Cable and

Linderoth (1963) and later reported from Florida by

Bullock and Mateo (1970).

Dollfusentis longispinus (Cable and Linderoth, 1969)

Golvan, 1969, was originally described from Anisotremus

virginicus in Jamaica and probably extends into Florida

waters. The validity of this species was questioned by

Bullock and Mateo (1970). Its suggested synonymy with

D. chandleri may be validated upon careful study of a

long series ofspecimens and data analysis.

Tegorhynchusfurootus (Van Cleave and Lincicome, 1939)

Bullock and Mateo, 1970, was described as Illiosentis

furcatus from Menticirrhus americanus off Grand Isle,

Louisiana. It has since been reported from the same

host species in coastal Mississippi and Alabama

(Overstreet, 1978) and from Fundulus similis, Dasyatis

sabina, D. say, and D. americana off Horn and Ship

islands, Mississippi (Buckner et aI., 1978). Museum (the

Manter Laboratory) data include records of T. furcatus

from D. say, and Menticirrhus americanus in the Caribbean, from Trachinotus carolinus in Mississippi, from M.

saxatilisin Massachusetts, and from Polydactylus octonemus

and M. americanus in Texas.

Intermediate hosts of T. furcatus include Lepidactylus

sp. and Haustorius sp. Both amphipod genera were

reported off Horn Island, Mississippi, where adults were

collected from fish definitive hosts (Buckner et a\.,

1978) as well as from Lepidactylus sp. in the northern

Gulf of Mexico (Overstreet, 1978).

Family Pomphorhynchidae Yamaguti, 1939.

Genus pomphorhynchus Monticelli, 1905, is represen ted

by four species in this review. Two species, P. rocci

Cordonier and Ward, 1967, and P. tereticollis (Rudolphi,

1809) Meyer, 1932, primarily infect marine fishes but

also may be found in freshwater coastal fishes. The

other two species, P. bulbocolli Linkins in Van Cleave,

1919, and P. lucyi Williams and Rogers, 1984, are freshwater species that occasionally infect marine or brackish water fishes. The first two species have been very

difficult to distinguish by using morphological characteristics or host ecology; see Samuel et aI. (1976),

Huffman and Nickol (1978), and Gleason and Huffman

(1981). Differentiation based on comparative length of

proboscis hooks in the 60% to 80% position region and

a certain hook ratio appear to have resolved the problem (Gleason and Huffman, 1981).

Pomphorhynchus bulbocolli Linkins in Van Cleave, 1919,

is widely distributed in North American freshwater fishes

and has been reported from at least 81 host species

(Samuel et aI., 1976). See Amin (1987b) for a discussion of its ecology and host relationships. Reference is

made here to only marine fish hosts of P. bulbocolli. Van

Cleave (1924a) reported P. bulbocollifrom Roccus lineatus

(= Morone saxatilis), the type host of P. rocci. Pomphorhynchus bulbocolli also infected the marine Microgadus

tomcod and Fundulus sp. from the brackish waters directly below the last dam on the Oyster River, New

Hampshire (Bullock, in Huffman and Nickol, 1978);

infected intermediate hosts may have washed over the

dam. Pomphorhynchus bulbocolli also was reported from

Anguilla rostrata in New Hampshire by Muzzall (1981).

Overstreet (1978) also reported P. bulbocolli from Amia

calva and other unnamed fishes from low-salinity habitats in the northern Gulf of Mexico.

The intermediate hosts ofP. bulbocolli include Hyalella

azteca and Gammarus pseudolimnaeus (Amphipoda), and

Caecidotea communis (Isopoda) (Gleason, 1987, 1989;

Muzzall, 1981, 1982). Muzzall (1981) also reported P.

bulbocolli infections in "estuarine amphipods of the genus Gammarus" from New Hampshire. Attempts to infect H. azteca with P. rocci failed (Johnson, 1973, personal commun. in Samuel et aI, 1976) thus lending

credibility to the distinctive identity of each of P. rocci

and P. bulbocolli.

Pomphorhynchus lucyi Williams and Rogers, 1984, appears to have little host specificity. It was described

from 14 mostly freshwater species offish belonging to 7

families and 6 orders from northern Florida and south-
________________________ Amin: Acanthocephala of the Eastern United States 15

ern Alabama. Fish hosts with marine or brackish affinities included Atlantic needlefish, Strongylura marina,

and American eel, Anquilla rostrata. No other reports of

p. lucyi are published to date.

Pomphorhynchus rocci Cardonnier and Ward, 1967, was

described from Morone saxatilis off the North Carolina

coast. Other marine hosts include Paralichthys dentatus

and Merluccius bilinearis from Raritan Bay, New Jersey

(Meyers, 1978) and Pleuronectes putnami from Great Bay

Estuary, New Hampshire (Burn, 1980). Brackish water

records include infections in Micropterus salmoides in

North Carolina and freshwater records involved letalurus

furcatus, Pylodictus olivaris, and Aplodinotus grunniens in

Louisiana and Mississippi (Huffman and Nickol, 1978).

A curious record from A. grunniens in a noncoastal

freshwater stream, the Ohio River in West Virginia, was

reported by Joy et al. (1986). The brackish water marine amphipod Gammarus tigrinus is the known crustacean intermediate host of P. rocci (see Samuel et aI,

1976; Muzzall, 1981).

Pomphorhynchus tereticollis (Rudolphi, 1809) Meyer, 1932,

is a Nearctic species that also is reported in Ireland

from Gadus sp., Acipensersturio, and Salmo fario. In North

America, it was reported "in great numbers" from Morone

saxatilis as well as from Cynoscion regalis and Synodus

foetens off Woods Hole, Massachsetts and North Carolina by Linton (1889) as Echinorhynchus proteus Westrumb,

1821, and Meyer (1932), respectively.

Family RHADINORHYNCHIDAE Travassos, 1923.

Subfamily GORGORHYNCHINAE Van Cleave and

Lincicome, 1940.

Genus Gorgorhynchoides Cable and Linderoth, 1963, is

distinguished from other rhadinorhynchid genera by

an anterior dorsal swelling of the trunk; the swelling

mayor may not be spined. The description of much of

the Gorgorhynchoides material by Cable and Mafarachisi

(1970) was based on collections by Linton deposited at

the USNM Helm. ColI., catalogue nos. 8044, 8045. Cable

and Mafarachisi (1970) reported that these specimens

were not mentioned in Linton's publications and indicated that they were aware that Linton "intended to

erect a new genus and had selected a type male and a

type female." They were not aware that he had already

described the genus as "Aulorhynchus n. gen." (with

trumpet snout) in a significant contribution that was

not published because of his death (Linton3). Linton

named his type species "Aulorhynchus longulus n. sp.,"

the description and figures (64-71) of which are very

similar to those of Gorgorhynchoides lintoni Cable and

Mafarachisi, 1970. It was collected from Seriola lalandi in

1910 near the Woods Hole region, Massachusetts.

The species ofthe genus Gorgorhynchoides, as emended

by Cable and Mafarachisi (1970), are treated independently as valid in this study. The validity of these species

is, however, not unquestionable and separate study of

their systematic identity is warranted.

Gorgorhynchoides bullocki Cable and Mafarachisi, 1970,

was originally collected from Caranx hippos, Lutjanus

griseus, and Bairdiella chrysura off the Florida Gulf coast

and farther south into Chetumal Bay, Mexico.

Gorgorhynchoides elongatus Cable and Linderoth, 1963

(description emended by Cable and Mafarachisi, 1970),

was initially collected from Caranx crysos in Curat,;:ao but

probably extends farther north into the geographical

range covered by this study.

Gorgorhynchoides lintoni Cable and Mafarachisi, 1970,

was reported from Caranx crysos, Seriola lalandi, and

Trachinotus falcatus from the northern Atlantic coast of

the United States, see discussion of the genus (above).

Genus gorgorhynchus Chandler, 1934, was established to

contain Echinorhynchus medius Linton, 1908 (whose cement glands were reinterpreted by Chandler, 1934)

and to describe G. gibber which was considered synonymous with G. medius (see Amin, 1985a). Three species

of Gorgorhynchus are reported here.

Gorgorhynchus cablei Golvan, 1969, was described from

Lutjanus jocu from Jamaica and is probably present in

the southern range of the area covered by this study.

Gorgorhynchus clavatus Van Cleave, 1940, was collected

from Paralabrax humeralis in the Galapagos andJamaica

and probably extends into the geographical range covered by this study; Lutjanus appears to serve as a

paratenic host.

Gorgorhynchus medius (Linton, 1908) Chandler, 1934 (=

Gorgorhynchus gibber, Chandler, 1934), was reported from

Ariusfelis, MycterCYfJerca venenosa, and OrthCYfJristis chrysCYfJtera

from the Atlantic coast of Florida and Texas and farther south into the Gulf of Mexico and Bermuda Islands. The species was redescribed from Arius melanCYfJus

collected in Veracruz, Mexico, by Salgado-Maldonado

(1978). Encapsulated immature worms were found

"among the viscera ... ofspiny rayed fishes" (Chandler,

1935) .

Leptorhynchoides thecatus (Linton, 1891) Kostylev,

1924, is a common parasite found in at least 6 families

of freshwater fishes, particularly centrarchids, and extends over a wide geographical range in North America.

It is often found encysted in the body cavity of fish
16 NOAA Technical Report NMFS 135

paratenic hosts (see Petrochenko, 1956; Yamaguti, 1963;

Hoffman, 1967; and museum records for partial host

lists). Its ecology and host associations were reported by

Amin (1988) in Wisconsin and by others elsewhere. Its

intermediate hosts include amphipods of the genus

Hyalella. It has been reported from "fresh-brackish water interface" in a "variety of marine intruders" in the

northern Gulf of Mexico in Mississippi and Alabama by

Overstreet (1978).

Subfamily SERRASENTINAE Petrochenko, 1956.

Serrasentis sagittifer (Linton, 1889) Van Cleave, 1923, is

the only species of the genus Serrasentis Van Cleave,

1923, that is reported from the study area. It was most

completely described by Travassos (1966) and redescribed by Salgado-Maldonado (1978). Synonymies

were reported by many authors such as Travassos (1966),

Jupta and Jain (1985), Amin (1985a), and Amin et al.

(1984). It has been reported from various marine waters in the world including the tropical Atlantic with

records extending from Texas and Florida in Ariusfelis,

Bairdiella chrysura, Paralichthys dentatus, Rachycentron

canadum, Cynoscion regalis to New Jersey in P. dentatus

and Pomatomus saltatrix (see Meyers, 1978; Overstreet,

1978; and museum records). Reservoir (paratenic) hosts

in Atlantic North America include B. chrysura,

Centropristis striata, Clupea harengus, Coryphaena hippurus,

Lagodon rhomboides, Leiostomus xanthurus, Micropogonias

undulatus, Orthopristis chrysoptera, Paralichthys albigutta,

Pomatomus saltatrix, Prionotus tribulus, Stenotomus chrysops,

Synodus foetens, and Cobiomorus fuscus; and see Linton

(1889,1901,1905; Overstreet, 1978).

Order NEOECHINORHYNCHIDA Southwell and

MacFie, 1925.

Family NEOECHINORHYNCHIDAE Ward, 1917.

SubfamilyATACTORRYNCHINAE Petrochenko, 1956.

Atactorhynchus verecundus Chandler, 1935, was described

from Paralichthys lethostigma in Galveston Bay, Texas,

and subsequently reported from Cyprinodon variegatus

in other Texas locations (Bullock, 1957), the west coast

of Florida (Bullock, 1960), Mobile Bay, Alabama (Williams and Gaines, 1974), Mississippi (Overstreet, 1978),

and Long Island, New York (Bullock, 1962). This acanthocephalan species appears to be very host specific to

C. variegatus.

Floridosentis elangatus Ward, 1953, was described from

Mugil cephalus in Biscayne Bay, Miami, Florida, and

later collected from the same host species in southern

Texas (Bullock, 1957) and the the southwestern coast

of Florida (Bullock, 1960). It was redescribed from the

same host species in Mexico by Salgado-Maldonado

and Barquin-Alvarez (1978). Other hosts include M.

Curema from Puerto Rico (Cable and Quick, 1954) and

from Mobile Bay, Alabama (Williams and Gaines, 1974).

The same two species of Mugil also are reported hosts

of F. elongatus in the Pacific Ocean, at least off the

Mexican coast at La Paz, Baja California, and Mazatlan,

Sinaloa (Bravo-Hollis, 1969).

Subfamily NEOECHINORHYNCHINAE Travassos,

1926.

Neoechinorhynchus daryphorus Van Cleave and Bangham,

1949, is a rarely reported species that was originally

described from Jordanella floridae in the Cape HazeEnglewood area of Florida. The recovery of similar

forms from marine waters elsewhere in the world (Amin

et aI., 1984) suggests that the geographical distribution

ofNeoechinorhynchusforms with unequal proboscis hooks

in the anterior circle is considerably wider than previously thought. Extensive collections at least in the Florida

estuaries should prove their presence there. Larval stages

were recovered from the mesenteries ofFundulus majalis

(see Van Cleave and Bangham, 1949), Lucania parva

and Notropis sp. in Florida brackish waters at Cape

Haze-Englewood (Bullock, 1960).

Neoechinorhynchus rostratum Amin and Bullock, 1998.

There are about 18 species of Neoechinorhynchus Stiles

and Hassall, 1905, that occur in North American freshwater hosts, mostly centrarchids and turtles. The

Neoechinorhynchus material collected from American eel,

Anguilla rostrata, along the New England and Canadian

coasts is very similar to the freshwater N. cylindratus

(Van Cleave, 1913) Van Cleave, 1919, but differs in the

following features. It has an estuarine geographical

distribution in eel, does not occur as adults in the

nearby freshwater centrarchids, and has a slightly different morphology of the egg and female posterior

end. This new species maintains breeding populations

in eel. Adults collected from Microgadus tomcod do not

reach sexual maturity in this marine-estuarine host.

Immature individuals were frequently found in liver

and mesenteries of Fundulus heteroclitus.

Family TENUISENTIDAE Van Cleave, 1936.

Paratenuisentis ambiguus (Van Cleave, 1921) Bullock and

Samuel, 1975, the only species ofthe genus, was erected

for Tanaorhamphus ambiguus Van Cleave, 1921, from

Alosa pseudoharengus, Anguilla rostrata, Fundulus

diaphanus, and Morone americanus. The natural definitive host is A. rostrata from Maryland, Massachusetts,

and New Hampshire coastal areas. Unnatural hosts include Microgadus tomcod and Fundulus heteroclitus from
________________________ Amin: Acanthocephala of the Eastern United States 17

New Hampshire and Alosa pseudoharengus, Marone

americanus, and F diaphanus. Intermediate hosts include Cammarus mucronatus from Maryland and C.

tigrinus from New Hampshire (Bullock and Samuel,

1975).

Order POLYMOPHIDA Petrochenko, 1956.

FamilyPOLYMORPHIDAE Meyer, 1931.

Six polymorphid genera from birds and mammals represen t the only acan thocephalans reported from

nonpiscine hosts in this review.

Genus Andracantha Schmidt, 1975, was erected for three

polymorphid acanthocephalans having genital spines

and two fields of trunk spines. Two of these species had

been assigned previously to Corynosoma Liihe, 1904, and

occur within the coastal areas covered by this study.

The third species, Andracantha phalacrocoracis (Yamaguti,

1939) Schmidt, 1975, is known from the pelagic cormorant, Phalacrocorax pelagicus, from Alaska and Japan

(Schmidt, 1975) and from the common cormorant, P.

carbo, in Czechoslovakia (Scholz et aI., 1992). Three

other species ofAndracantha are found elsewhere in the

world.

Andracanthagravida (Alegret, 1941) Schmidt, 1975, was

reported from the double-crested cormorant, P. auritus

floridanus, and brown pelican, Pelicanus occidentalis

carolinensis, in Cuba, Florida, and Louisiana (Schmidt,

1975), and the Gulf of Mexico.

Andracantha mergi (Lundstrom, 1941) Schmidt, 1975, is

less host specific than A. gravida. It was reported from

the red-breasted merganser, Mergus serrator, and

red-throated loon, Cavia stellata, in Alaska; from the

lesser scaup duck, Aythya affinis, in New York; from the

black-crowned night heron, Nycticorax nycticorax, in New

Hampshire Uuvenile worms) (Schmidt, 1975); and from

the American bald eagle, Haliaeetus leucocephalus, in

ewJersey and Nova Scotia (Nickol and Kocan, 1982).

Additional hosts include P. auritus in Maine and Pholis

gunnellus (rock eel, paratenic host) in New Brunswick,

Canada (museum collections). The original description was based on specimens collected from M. serrator

in Sweden.

Arhythmorhynchus uncinatus (Kaiser, 1893) LUbe, 1912,

was one of three species of Acanthocephala described

from unnamed hosts in Florida. It was the "object of

conjecture" among taxonomists until it was later more

completely described by Van Cleave (1924b) from some

of Kaiser's original material. Adults have not been reported since and the definitive host(s) remain unknown.

Immature worms were recovered from the mesenteries

of a marine fish, sheepshead, Archosargus probatocephalus,

in Florida by Bullock (1960).

Genus Bolbosoma Porta, 1908, includes parasites of

whales, and as immature worms, may occasionally infect humans (Beaver et aI., 1983; Tada, et aI., 1983).

Two species are reported here.

Bolbosoma turbinella (Diesing, 1851) Porta, 1908, was

primarily reported from the blue whale, Balaenoptera

musculus, in North American Atlantic waters. It is not,

however, restricted to Atlantic blue whales and is also

found in the Pacific waters of North America. Other

definitive hosts include sei whale, fin whale, humpback

whale, North Atlantic right whale, and North Atlantic

bottlenose whale (Measures, 1992, 1993). The sei whale

and right whale feed mostly on copepods. The fin whale

and the humpback whale are euryphagous and feed on

swarming fish (Nemato, 1959; Mitchell, 1975). Ten families offish from various geographical locations include

known paratenic hosts of B. turbinella (see review by

Measures, 1992).

Bolbosoma vasculosum (Rudolphi, 1819) Porta, 1908, has

been reported from dolphins, Delphinus delphis and

Mesoplodon bidens in North American Atlantic waters,

Mediterranean Sea, and Sea ofJapan as well as from the

roundnose grenadier, Coryphaenoides rupestris in the North

Atlantic Ocean (Bakey and Zubchenko, 1984). Fish

paratenic hosts from various geographical locations are

listed in Petrochenko (1958) and Yamaguti (1963)

Carynosoma constrictum Van Cleave, 1918, was described

from American scoter, Oidemia americana, in Yellowstone

Lake, Wyoming, by Linton (1892) as Echinorhynchus

striatus. It has since been reported from various species

of ducks in inland and coastal North American locations. Hosts include Anas acuta, A. americana, A.

carolinensis, A. clypeata, A. discars, A. platyrhynchos, A.

rubripes, Anas spp., Aythya affinis, A. marila, Bucephala

albeola, Cygnus alar, C. buccinator, Fulica americana,

Erismatura jamaicensis rubida, Melanitta deglandi, M. niger,

Oidemia americana, Phalaropus tricolor, Podiceps grisegena,

and Recurvirostrata americana (see Van Cleave, 1918;

Van Cleave and Starrett, 1940; Petrochenko, 1956;

Yamaguti, 1963; and museum records). The North

American Atlantic distribution is often more northerly,

e.g. in New Hampshire, New York, and into Canada in

Anas, Aythya, and Cygnus.

Corynosoma wegeneri Heinze, 1934, adults commonly

infect gray and ringed seals, Erignathus barbatus, Halichoerus grypus, Phoca fasciata, P. hispida, P. richardi, and

P. vitulina from the northwest Atlantic to Greenland.
18 NOAA Technical Report NMFS 135

Paratenic fish hosts include Gadus morhua, Hemitripterus

americanus, Hippoglossoides platessoides, Macrozoarces

americanus, Melanogrammus aeglefinus, Myoxocephalus

octodecemspinosus, M. scorpius, Osmerus mordax, Pleuronectes

americanus, P. ferrugineus, and Scophthalmus aquosusfrom

New Hampshire, Maine, the Canadian New Brunswick

and Scotian Shelf, and Greenland (Meyer, 1954;

Petrochenko, 1956; Marcogliese and McClelland, 1992;

and museum records). Recent increases in C. wegeneri

populations in ground fish appear to be related to

observed marked growth in seal populations (Marcogliese and McClelland, 1992). Similar observations were

made on other species of Corynosoma of the Antarctic

fur seal in South Georgia and South Drkneys by

Zdzitowiecki (1987).

Genus Polymarphus Liihe, 1911.

The position, concept, and composition of the genus

Polymorphus were critically evaluated and a new diagnosis of the genus was made based on the designation of

Hexaglandula Petrochenko, 1950, and Subcorynosoma

Hoklova, 1967, asjuniorsynonyms (Amin, 1992). Other

taxonomic changes were proposed and two subgenera

are now recognized: Polymorphus Liihe, 1911, with 36

species and Profilicollis Meyer, 1931, with 10 species. A

key to the species of the genus is included in Amin's

(1992) critical revision of this genus.

Eight polymorphic species are recognized from birds

whose range includes Atlantic North American coastal

areas; four in each of the subgenera Polymorphus and

Profilicollis.

Polymarphus (Polymarphus) acutisVan Cleave and Starrett,

1940, is a Nearctic species that usually infects ducks,

grebe, gull, coot, and godwit in coastal and inland

North American and Siberian locations. It was originally described from mallard, Anas platyrhynchos, and

pintail, A. acuta, taken from the Illinois River, Illinois.

Other host records include Anas clypeata, A. strepera,

Aythya ferina, A. fu ligula, Bucephala clangula, Fulica

americana, Larus argentatus, Limosa limosa, Oxyura

leucocephala, and Podiceps grisegena (see Petrochenko,

1958; Amin, 1992). For a comprehensive listing of Russian hosts see Hoklova (1978).

Polymarphus (Polymarphus) brevis (Van Cleave, 1916)

Travassos, 1926, is a parasite of osprey, bitterns, and

herons in North America. It was originally described as

Arhythmorhynchus brevis from the bittern Botaurus

lentiginosus in Baltimore, Maryland. Travassos (1926)

assigned the'species to the genus Polymorphusand Meyer

(1932) accepted this assignment. Van Cleave (1945)

reassigned it back to Arhythmorhynchus and SalgadoMaldonado (1980) redescribed it from Nycticorax

nycticorax in Michoacan, Mexico. The species is herein

recognized in Polymorphus (see Amin, 1992, for a detailed discussion).

Definitive hosts from Atlantic coastal states include

N. nycticorax from New Hampshire and Ardea herodias

from Louisiana. Other hosts from elsewhere in the

United States include Botaurus virescens, Gavia immer,

Pandion haliaetus carolinensis, and Pelecanus erythrorhynchos.

Paratenic fish hosts include Gambusia affinis, lctalurus

nebulosus, Leiostomus xanthurus, Lepomis cyanellus, and

Menidia beryllina in Florida and New Hampshire. For

host records see Van Cleave (1916, 1945), Petrochenko

(1958), Yamaguti (1963), Amin (1992), and museum

records.

Polymarphus (Polymarphus) obtusus Van Cleave, 1918,

was described from water turkey, Anhinga anhinga, in

Florida. It has since been reported from various aquatic

birds in coastal states extending north to New York

where it was recorded from Anas platyrhynchos. Other

hosts include Ardea herodias, Erismatura jamaicensis,

Mergus merganser, and Phoenicopterus ruber (see Van

Cleave, 1918, 1924£1; Petrochenko, 1958; Yamaguti, 1963;

Amin, 1992; and museum records).

Polymarphus (Polymarphus) trochus Van Cleave, 1945, is

a common Nearctic parasite of coot, Fulica americana

(type host) but rarely infects the mallard duck, Anas

platyrhynchos, and common moorhen, Gallinula chloropus,

in North America (Priebe, 1952; Podesta and Holmes,

1970; McDonald, 1988). It is not the same as "Polymorphus

trochus" of Hoklova (1966) which is recovered mostly

from ducks, Anas spp., in Asia. The latter is probably a

new species (Amin, 1992) In North America, P. (P.)

trochuswas reported from Atlantic coastal states extending

from Florida to New York, as well as from other inland

states (see above references for more information).

Polymorphus (Profllicollis) altmani (Perry, 1942) Van

Cleave, 1947, is a parasite of surf scoter and sea diving

ducks Melanitta perspicillata and M. deglandi in California but has also been reported from the former species

on the North West Atlantic coast off New Brunswick

and Newfoundland (Bourgeois and Threlfall, 1982).

Directed collections of coastal New England states will

probably reveal its presence there.

Polymarphus (Profilicollis) botulus (Van Cleave, 1916)

Witenberg, 1932, was originally described as Filicollis

botulus from the circumboreal eider duck Somateria

dresseri in Maine. Schmidt and Kuntz (1967) and Amin

(1985£1) recognized that species in Polymorphus as the

type of the subgenus Profilicollis. Hoklova (1974) redescribed the species and reinstated it in her family Filicollidae. This placement is not recognized here.
_________________________ Amin: Acanthocephala of the Eastern United States 19

Polymorphus (P.) botuluswas reported from a variety of

sea ducks on both coasts of North America as well as

from Eurasia. On the North American Atlantic coast, it

also infects the Holarctic common eider duck, S.

mollissima, where its seasonality and host relationships

were reported by Bishop and Threlfall (1974). See also

Bourgeois and Threlfall (1982), Clark et al. (1958), and

Threlfall (1968) for additional geographical records

from eider ducks and scoters (Melanitta perspicillata,

M. nigra, M. fusca) on the North West Atlantic coast.

Records from the North American Pacific coast and

Eurasia include a wider variety of birds other than eider

ducks and scoters. See Yamaguti (1963) and Ching

(1989), and others quoted therein, for records of the

shore crab intermediate hosts of P. (P.) botulus in these

locations.

Polymorphus (Profilicollis) kentiVan Cleave, 1947 (= F"ilicollis sphaerocephalus Harrington and Pillsbury, 1938,

and Harrington, 1939) was described from Larus

argentatus smithsonianus in Kent Island, New Brunswick,

Canada. Bullock3 found acanthocephalans tentatively

identified as P. (P.) kenti in the sand crab, Emerita sp.,

from Florida. These specimens are in the Nebraska

State Museum, Lincoln, HWML Coll., nos. 33481 and

33482, as "Polymorphus sp. ex Emerita sp." In 1960, Bullock2 attempted to collect P. (P.) kenti from 17 Kent

Island gulls that did not yield any worms. Absence of

worms is not a rare phenomenon. Hosts from which

certain acanthocephalans were initially identified occasionally yield no parasites upon examination in the

same locality years later. The distribution ofP. (P.) kenti

is not limited to the east coast. Reish (1950) described

it from gulls in Oregon and from the sand crab Emerita

analoga (in larval form).

Polymorphus (Profilicollis) mtYorLundstrom, 1942, adults

were described from goldeneye duck, Bucephala clangula,

in Sweden and have since been reported from various

Scandinavian locations. Schmidt and MacLean (1978)

found it in New England coastal locations in other

diving ducks including white-winged scoter, Melanitta

deglandii, lesser scaup, Aythya affinis, bufflehead,

Bucephala albeola, and canavasback, A. valisineria. Juvenile P. (P.) major were also found by Schmidt and

MacLean (1978) from rock crabs, Cancer irroratus, in

Maine shore waters.

Genus Southwellina Witenberg, 1932, was erected by

Witenberg (1932) who designated Arhythmorhynchus

hispidus Van Cleave, 1925, as its type species. The genus

was not seriously recognized by acanthocephalan taxonomists until Schmidt (1973) redefined and resurrected it, and recognized three species in it; two species

are included in this treatment.

Southwellina dimorpha Schmidt, 1973, was described from

white ibis, Eudocimus albus, in Florida. Intermediate

hosts include the crayfish Procambarus clarkii from Pecan Island, Louisiana (Schmidt, 1973).

Southwellina hispida (Van Cleave, 1925) Witenberg, 1932,

was first described from juveniles collected from frogs

inJapan. Adults were subsequently described from specimens found in the heron Nycticorax n. nycticorax in Japan

by Fukui (1929). Synonyms of S. hispida include Arhythmorhynchus duocinctusChandler, 1935, which was described

from juveniles found in the mesenteries of the southern flounder, Paralichthys lethostigma, from Galveston

Bay, Texas. Lincicome (1943) subsequently described

the same species from N. nycticoraxfrom the Bronx Zoo.

Additional definitive hosts include Pelecanus occidentalis

and Phalacrocorax auritus from Florida, and paratenic

fish hosts include Menidia beryllina and Fundulus grandis

from Florida and Texas (museum records). Meyers

(1978) reported additional fish hosts (Paralichthys

dentatus, Pomatomus saltatrix, Cynoscion regalis) from

Raritan Bay, NewJersey. Synonymies of S. hispida from

N. nycticorax and other definitive bird hosts (Ardea,

Egretta, Phalacrocorax) from Japan, from Primorye and

the Crimea (Russia), and from the Galapagos Islands

were discussed by Schmidt (1973). This species' broad

geographical distribution (Van Cleave, 1940) extends

into Europe, e.g. Hungary (Dimitrova, 1991).

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Systematic Index of Acanthocephala _

Acanthocephala. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1

Acanthocephalus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2, 8

Acanthocephalus dims . . . . . . . . . . . . . . . . . . . . . . .. 6, 12

Andracantha. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2,10, 17

Andracantha gravida. . . . . . . . . . . . . . . . . . . . . . .. 10, 17

Andracantha mergi. . . . . . . . . . . . . . . . . . . . . . . . .. 10, 17

Andracantha phalacrocoracis . . . . . . . . . . . . . . . . . . .. 17

Apororhynchida. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Aporohynchidae . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1

Archiacanthocephala 1,2

Arhythmacanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

Arhythmorhynchus 2, 11, 18

Arhythmorhynchus brevis. . . . . . . . . . . . . . . . . . . . . . . .. 18

Arhythmorhynchus duocinctus . . . . . . . . . . . . . . . . . . .. 19

Arhythmorhynchus uncinatus 11, 17

Atactorhynchinae. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Atactorhynchus 2, 10

Atactorhynchus verecundus. . . . . . . . . . . . . . . . . . . .. 9, 16

Aulorhynchus longulus . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Bolbosoma 2, 10, 17

Bolbosoma turbinella. . . . . . . . . . . . . . . . . . . . . . . .. 10, 17

Bolbosoma vasculosum . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Cavisomidae 2, 8, 12

Centrorhynchidae . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

Corynosoma 2, 10, 17

Corynosoma constrictum . . . . . . . . . . . . . . . . . . . . . . . .. 17

Corynosoma wegeneri. . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Dendronucleatidae . . . . . . . . . .. 2

Diplosentidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

Dollfusentis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2, 9, 13

Dollfusentis chandleri . . . . . . . . . . . . . . . . . . . . . . . .. 9, 13

Dollfusentis bravoae . . . . . . . . . . . . . . . . . . . . . .. 13

Dollfusentis centrorhynchus. . . . . . . . . . . . . . . . . . . . . .. 14

Dollfusentis heteracanthus. . . . . . . . . . . . . . . . . . . . . . .. 14

Dollfusentis longispinus . . . . . . . . . . . . . . . . . . . . . . . .. 14

Echinorhynchida 2,7,12

Echinorhynchidae 2,8, 12

Echinorhynchinae. . . . . . . . . . . . . . . . . . . . . . . . . . .. 8

Echinorhynchus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2, 8

Echinorhynchus gadi 12,13

Echinorhynchus gadi callariae . . . . . . . . . . . . . . . . . . .. 12

Echinorhynchus gadi virentis . . . . . . . . . . . . . . . . . . . .. 12

Echinorhynchus laurentianus . . . . . . . . . . . . . . . . . . . .. 13

Echinorhynchus medius. . . . . . . . . . . . . . . . . . . . . . . . .. 15

Echinorhynchus proteus. . . . . . . . . . . . . . . . . . . . . . . . .. 15

Echinorhynchus vancleavei . . . . . . . . . . . . . . . . . . . . . .. 12

Echinorhynchus yamagutii . . . . . . . . . . . . . . . . . . . . . .. 13

Eoacanthocephala 26

Fessisentidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

Filisoma. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2, 8

Filisoma fidum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12

Filicollis botulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Floridosentis 2, 10
24 NOAA Technical Report NMFS 135 _

Floridosentis elongatus . . . . . . . . . . . . . . . . . . . . . . .. 9, 16

Gigantorhynchida . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1

Gigantorhynchidae 1

Gorgorhynchinae 8,15

Gorgorhynchoides 2,19, 15

Gorgorhynchoides bullocki . . . . . . . . . . . . . . . . . . . . . . .. 15

Gorgorhynchoides elongatus . . . . . . . . . . . . . . . . . . .. 9, 15

Gorgorhynchoides lintoni. . . . . . . . . . . . . . . . . . . . . . . .. 15

Gorgorhynchus 2,9, 15

Gorgorhynchuscabwi 15

Gorgorhynchus clavatus . . . . . . . . . . . . . . . . . . . . . . . .. 15

Gorgorhynchus medius . . . . . . . . . . . . . . . . . . . . . . .. 7, 15

Gorgorhynchus gibber . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Gyracanthocephala . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

Heteracanthocephalidae. . . . . . . . . . . . . . . . . . . . . . .. 2

Hexaglandula 18

Hypoechinorhynchidae 2

Illiosentidae 2,8, 13

Illiosentis furcatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Leptorhynchoides 2,9

Leptorhynchoides thecatus . . . . . . . . . . . . . . . . . . . . . . .. 15

Macracanthorhynchus hirudinaceus . . . . . . . . . . . . . . . . 3

Moniliformida 2

Moniliformidae 2

Neoechinorhynchida. . . . . . . . . . . . . . . . . . . . . . .. 2, 16

Neoechinorhynchidae 2,6,8,16

Neoechinorhynchinae. . . . . . . . . . . . . . . . . . . . . . . .. 16

Neoechinorhynchus . . . . . . . . . . . . . . . . . . . . . . . . . . 2, 9

Neoechinorhynchus cylindratus. . . . . . . . . . . . . . . . .. 6, 16

Neoechinorhynchus doryphorus . . . . . . . . . . . . . . . . . . .. 16

Neoechinorhynchus rostratum . . . . . . . . . . . . . . . . . . . .. 16

01igacanthorhynchida 2

01igacanthorhynchidae 2

Oligacanthorhynchus longissimus. . . . . . . . . . . . . . . . . .. 3

Palaeacanthocephala 2, 6, 12

Paratenuisentis 2, 8

Paratenuisentis ambiguus. . . . . . . . . . . . . . . . . . . . .. 8, 16

Plagiorhynchidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

Polyacanthocephala. . . . . . . . . . . . . . . . . . . . . . . . . . .. 1

Polymorphida 2,7, 17

Polymorphidae 7, 17

Polymorphus 2,11,18

Polymorphus (Polymorphus) acutis. . . . . . . . . . . . . . . .. 18

Polymorphus (Polymorphus) brevis. . . . . . . . . . . . .. 11, 18

Polymorphus (Polvmorphus) obtusus . . . . . . . . . . . . . .. 18

Polymorphus (Polymorphus) trochus. . . . . . . . . . . . . . .. 18

Polymorphus (Profilicollis) altmani 11, 18

Polymorphus (Profilicollis) botulus. . . . . . . . . . . . . . . .. 18

Polymorphus (Profilicollis) kenti. . . . . . . . . . . . . . . . . .. 19

Polymorphus (Profilicollis) major. . . . . . . . . . . . . . . . .. 19

Pomphorhynchidae 2,7,14

Pomphorhynchus 2,7

Pomphorhynchus bulbocolli . . . . . . . . . . . . . . . . . . . .. 7, 14

Pomphorhynchus lucyi. . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Pomphorhynchus rocci. . . . . . . . . . . . . . . . . . . . . . .. 14, 15

Pomphorhynchus tereticollis. . . . . . . . . . . . . . . . . . .. 14, 15

Profilicollis 18

Quadrigyridae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2

Rhadinorhynchidae . . . . . . . . . . . . . . . . . . . . . .. 2, 8, 15

Serrasentinae 8, 16

Serrasentis 2,8, 16

Serrasentis sagittifer. . . . . . . . . . . . . . . . . . . . . . . . . .. 7, 16

Southwellina 10,11,19

Southwellina dimorpha . . . . . . . . . . . . . . . . . . . . . .. 10, 19

Southwellina hispida. . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Subcorynosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Tanaorhamphus ambiguus. . . . . . . . . . . . . . . . . . . . . .. 16

Tegorhynchus 2,7,9

Tegorhynchusfurcatus . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Telosentis tenuicomis . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Tenuisentidae 2,8, 16

Systematic Index of Invertebrate

Intermediate Hosts

Aeginina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Aeginina longicomis. . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Amphithoe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Asellus intermedius. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12

Caecidotea communis . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Caecidotea militaris. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12

Calliopius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Calliopius laeviusculus . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Cancer irroratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Caprella. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Corophium lacustra. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Cyphocaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Emerita sp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Emerita analoga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Gamarellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Gamarellus angulosus. . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Gammarus 13,14

Gammarus mucronatus. . . . . . . . . . . . . . . . . . . . . . . . .. 17

Gammarus pseudolimnaeus . . . . . . . . . . . . . . . . . . . . .. 14

Gammarus tigrinus. . . . . . . . . . . . . . . . . . . . . . . . .. 15, 17

Grandidierella bonnieroides. . . . . . . . . . . . . . . . . . . . . .. 14

Haustorius sp. 14

Hyalella. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Hyalella azteca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Lepidactylus sp. 14

Mysis 13

Pontoporeia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Pontoporiea a.fJinis . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12

Procambarus clarkii. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Systematic Index of Vertebrate Definitive and

Paratenic Hosts

Acipenser oxyrhynchus. . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Acipenser sturio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15
________________________ Amin: Acanthocephala of the Eastern United States 25

Alosa aestivalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Alosa pseudoharengus 13, 16, 17

Alosa sapidissima . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Amia calva 14

Ammodytes hexapterus. . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Anarhichas lupus 13

Anas 17

Anas acuta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17, 18

Anas americana. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Anas carolinensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Anas clypeata . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17, 18

Anas discors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Anas platyrhynchos . . . . . . . . . . . . . . . . . . . . . . . . .. 17, 18

Anas rubripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Anas strepera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Anas spp 17

Anguilla rostrata 12,14,15,16

Anhinga anhinga. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Anisotremus virginicus . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Aplodinotus grunniens . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Archosargus probatocephalus. . . . . . . . . . . . . . . . . .. 13, 17

Ardea 19

Ardea herodias. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Ariusfelis 15,16

Arius melanopus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Aythya. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Aythya aJfinis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17, 19

Aythya ferina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Aythyafuligula 18

Aythya marila . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Aythya valisineria . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Bairdiella chrysura 13, 15, 16

Balaenoptera muscularis. . . . . . . . . . . . . . . . . . . . . . . .. 17

Bathygobius soporator . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Botaurus lentiginosus . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Botaurus virescens. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Bucephala albeola . . . . . . . . . . . . . . . . . . . . . . . . . .. 17, 19

Bucephala clangula 18, 19

Caranx crysos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Caranx hippos. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Centropristis striata. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Clupea harengus . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13, 16

Coregonus clupeaformis. . . . . . . . . . . . . . . . . . . . . . . . .. 12

Coryphaena hippurus . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Coryphaenoides rupestris . . . . . . . . . . . . . . . . . . . . .. 13, 17

Cygnus 17

Cygnus olor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Cygnus buccinator. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Cymatogaster aggregata. . . . . . . . . . . . . . . . . . . . . . . . .. 13

Cynoscion arenarius . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Cynoscion nebulosus . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Cynoscion regalis 13,15,16,19

Cyprinodon variegatus . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Dasyatis americana. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Dasyatis sabina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Dasyatis say. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Delphinus delphis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Egretta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Erignathus barabatus . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Erismatura jamaicensis. . . . . . . . . . . . . . . . . . . . . . . . .. 18

Erismatura jamaicensis rubida . . . . . . . . . . . . . . . . . . .. 17

Eucinostomus argenteus . . . . . . . . . . . . . . . . . . . . . . . .. 13

Eudocimus albus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Fin whale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Fundulus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Fundulus diaphanus 16, 17

Fundulus grandis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Fundulus heteroclitus . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Fundulus majalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Fundulus similis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Fundulus sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Fulica americana. . . . . . . . . . . . . . . . . . . . . . . . . . .. 17, 18

Gadus 15

Gadus macrocephalus. . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Gadus morhua . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13, 18

Gadus ogac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Gallinula chloropus. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Gambusia aJfinis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Gavia immer. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Gavia stellata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Gerres cinereus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Glyptocephalus cynoglossus . . . . . . . . . . . . . . . . . . . . . .. 13

Gobiomorusfuscus . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Haliaeetus leucocephalus. . . . . . . . . . . . . . . . . . . . . . . .. 17

Halichoeres grypus. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Hemilepidotus hemilepidotus. . . . . . . . . . . . . . . . . . . . .. 13

Hemitripterus americanus . . . . . . . . . . . . . . . . . . . .. 13, 18

Hippoglossoides platessoides . . . . . . . . . . . . . . . . . .. 13, 18

Hippoglossus hippoglossus. . . . . . . . . . . . . . . . . . . . . . .. 13

Humpback whale. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Icelinus Jilamentosus. . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Ictalurus furcatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

[ctalurus nebulosus . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Jordanella Jloridae. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Kyphosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12

Kyphosus sectatrix 12, 13

Labrisomus nuchipinnis . . . . . . . . . . . . . . . . . . . . . . . .. 14

Laqodon rhomboides . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Larimusfasciatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Larus argentatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Larus argentatus smithsonianus. . . . . . . . . . . . . . . . . .. 19

Leiostomus xanthurus . . . . . . . . . . . . . . . . . . . . . . .. 13, 16

Lepomis cyanellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Leptocottus armatus . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Limosa limosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Liparis atlanticus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Lobotes surinamensis. . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Lucania parva . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Lutjanus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Lutjanus griseus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15
26 NOAA Technical Report NMFS 135

Lutjanusjocu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Lycodes cortezianus. . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Macrourus berglax . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Macrozoarces americanus . . . . . . . . . . . . . . . . . . . . . . .. 18

Mallotus villosus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Melanitta deglandi 17, 18, 19

Melanitta fusca. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Melanitta niger. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Melanitta nigra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Melanitta perspicillata 18, 19

Melanogrammus aeglefinus 13, 18

Menidia beryllina 18, 19

Menticirrhus americanus . . . . . . . . . . . . . . . . . . . . . . .. 14

Mergus merganser. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Mergus serrator . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Merluccius bilinearis. . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Merluccius productus . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Mesoplodon bidens .. . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Microgadus proximus . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Microgadus tomcod 12, 13, 14, 16

Micropogonias undulatus 13, 16

Micropterus salmoides . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Monacanthus hispidus . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Morone amencanus. . . . . . . . . . . . . . . . . . .. 13, 14, 16, 17

Morone saxatilis 13, 14, 15

Mugil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Mugil cephalus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Mugil curema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Mulloidichthys martinicus . . . . . . . . . . . . . . . . . . . . . .. 14

Mycteroperca venenosa .. . . . . . . . . . . . . . . . . . . . . . . .. 15

Myoxocephalus octodecemspinosus. . . . . . . . . . . . . .. 13, 18

Myoxocephalus polycanthocephalus. . . . . . . . . . . . . . . .. 13

Myoxocephalus quadricornis . . . . . . . . . . . . . . . . . . . . .. 13

Myoxocephalus scorpius. . . . . . . . . . . . . . . . . . . . . . . . .. 18

Nemichthys scolopaceus . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Nezumia bairdi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

North Atlantic bottle nose whale. . . . . . . . . . . . . . .. 17

North Atlantic right whale. . . . . . . . . . . . . . . . . . . .. 17

Notropis sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Nycticoraxnycticorax 17,18,19

Nycticorax n. nycticorax . . . . . . . . . . . . . . . . . . . . . . . .. 19

Oidemia americana. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Oncorhynchus gorbuscha. . . . . . . . . . . . . . . . . . . . . . . .. 13

Oncorhynchus keta . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Oncorhynchus kisutch . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Oncorhynchus nerka . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Oncorhynchus tshawytscha . . . . . . . . . . . . . . . . . . . . . .. 13

Orthopristis chrysoptera 13, 15, 16

Osmerus mordax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Oxyura leucocephala . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Pandion haliaetus carolinensis . . . . . . . . . . . . . . . . . . .. 18

Paralabrax humeralis , 15

Paralichthys albigutta . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Paralichthys dentatus 13,15,16,19

Paralichthys lethostigma . . . . . . . . . . . . . . . . . . . . .. 16, 19

Pelecanus erythrorhynchos . . . . . . . . . . . . . . . . . . . . . . .. 18

Pelecanus occidentalis . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Pelicanus occidentalis carolinensis . . . . . . . . . . . . . . . .. 17

Phalacrocorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Phalacrocorax auritus . . . . . . . . . . . . . . . . . . . . . . .. 17, 19

Phalacrocorax auritusfloridanus . . . . . . . . . . . . . . . . .. 17

Phalacrocorax carbo .. . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Phalacrocorax pelagicus . . . . . . . . . . . . . . . . . . . . . . . .. 17

Phalaropus tricolor . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Phoca fasciata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Phoca hispida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Phoca richardi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Phoca vitulina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Phoenicopterus ruber . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Pholis gunnellus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Platichthys stellatus. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Platophrys ocellatus . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Pleuronectes americanus . . . . . . . . . . . . . . . . . . . . .. 13, 18

Pleuronectesferrugineus . . . . . . . . . . . . . . . . . . . . .. 13, 18

Pleuronectes putnami . . . . . . . . . . . . . . . . . . . . . . .. 13, 15

Podiceps grisegena. . . . . . . . . . . . . . . . . . . . . . . . . .. 17, 18

Podothecus acipenserinus . . . . . . . . . . . . . . . . . . . . . .. 12

Pollachius virens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Polvdactylus octonemus 13, 14

Pomatomus saltatrix . . . . . . . . . . . . . . . . . . . . . . . .. 16, 19

Prionotus tribulus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16

Pylodictus olivaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Rachycentron canadum. . . . . . . . . . . . . . . . . . . . . . . . .. 16

Raja radiata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Recurvirostrata americana . . . . . . . . . . . . . . . . . . . . . .. 17

Reinhardtius hippoglossoides. . . . . . . . . . . . . . . . . . . . .. 13

Roccus lineatus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Salmo fario . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Salmo salar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Salvelinus alpinus. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Salvelinus fontinalis . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Salvelinus malma . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Salvelinus namaycush. . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Scophthalmus aquosus ..... . . . . . . . . . . . . . . . . .. 13, 18

Sebastes aleutianus . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes alutus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes babcocki . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes borealis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes caurinus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes diplorora . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastesflavidus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes helvomaculatus . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes proriger. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes ruberrimus . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sebastes zacentrus . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Sei whale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 17

Seriola lalandi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Somateria dresseri . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Somateria mollissima. . . . . . . . . . . . . . . . . . . . . . . . . . .. 18

Stenotomus chrysops .. . . . . . . . . . . . . . . . . . . . . . . . . .. 16
________________________ Amin: Acanthocephala of the Eastern United States 27

Strongylura marina. . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Synodusfoetens 15,16

Tautogolabrus adspersus. . . . . . . . . . . . . . . . . . . . . . . .. 13

Theragra chalcogramma . . . . . . . . . . . . . . . . . . . . . . . .. 13

Trachinotus carolinus. . . . . . . . . . . . . . . . . . . . . . . . . .. 14

Trachinotus falcatus . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15

Tylosurus acus .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Tylosurus carribaeus . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Urophycis chuss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Urophycis tenuis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

Acknowledgments _

Prepararion 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."

Deep appreciation is expressed to Dr. Wilbur L. Bullock, a friend and a scholar, who generously shared

much of his professional expertise and knowledge, specimens and print resources, insights, and opinions. Dr.

Bullock also made available the original Linton manuscript (unpublished) and critically reviewed a rough

draft of the text. Dr. Bullock's help was indispensable

to the completion ofthis project. Drs. Mary H. Pritchard

and J. R. Lichtenfels kindly provided valuable assistance through their respective museums resources. Finally, the contributions of two anonymus reviewers and

editors are acknowledged.

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 of Marine Studies,

University of Delaware, Lewes, DE 19958.

Editorial Advisers _

A. Ralph Cavaliere, Department of Biology, Gettysburg

College, Gettvsburg, 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 ofComparative 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 ofmanuals, reviews manuscripts, and advises the

Scientific Editor of the National Marine Fisheries Service.

Coordinating Editor's Comments

Publication of the "Marine Flora and Fauna of the

Eastern United States" is most timely in view of the

growing universal emphasis on work in the marine,

environment and the critical need for precise and complete identification of organisms related to this work. It

is essential, if at all possible. that organisms be identified accurately 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 possible

prediction of attributes of organisms that have been

inadequately studied.

Omar M. Amin received his B.S. in Biology (1959),

M.S. in Medical Entomology (1963) from Cairo University, and Ph.D. in Parasitology (1968) from Arizona

State University (ASU). He worked at Old Dominion

University (1967-69), the Centers for Disease Control

(1969-70), and the University of Wisconsin as a Professor of Parasitology and Epidemiology from 1971 until

he established the Institute of Parasitic Diseases (IPD) in

1992, currently in Arizona, providing diagnostic and educational services to the medical community for human

parasitic diseases. He also is a Research Professor ofParasitology at ASU, pursuing his research activities in wildlife

parasitology, particularly in acanthocephalans offish.
28 NOAA Technical Report NMFS 135

Published Manuals

NOAA Tech Rep.

NMFS Circular no.

Marine Flora and Fauna of the Northeastern United States

NTIS no.

Ho,ju-Shey

Ho,ju-Shey

Annelida: Oligochaeta Cook, David G., and Ralph O. Brinkhurst

Protozoa: Ciliophora Borror, Arthur C.

Higher Plants of the Marine Fringe Moul, Edwin T.

Pycnogonida McCloskey, Lawrence R.

Crustacea: Stomatopoda Manning, Raymond B.

Crustacea: Decapoda Williams, Austin. B.

Tardigrada Pollock, Leland W

Cnidaria: Scyphozoa Larson, Ronald j.

Higher Fungi: Ascomycetes, Deuteromycetes, and Basidiomycetes Cavaliere, A. R.

Copepoda: Harpacticoida Coull, Bruce C.

Sipuncula Cutler, Edward B.

Echinodermata: Holothuroidea Pawson, David L.

Copepoda: Lernaeopodidae and Sphyriidae

Copepoda: Cyclopoids Parasitic on Fishes

Crustacea: Branchiura Cressey, RogerF.

Protozoa: Sarcodina: Amoebae Bovee, Eugene c., and Thomas K. Sawyer

Crustacea: Cumacea Watling, Les

Arthropoda: Cirripedia Zullo, Victor A.

Cnidaria: Scleractinia Cairns, Stephen D.

Protozoa: Sarcodina: Benthic Foraminifera Todd, Ruth, and Doris Low

Turbellaria: Acoela and Nemertodermatida Bush, Louise F.

Lichens (Ascomycetes) of the Intertidal Region Taylor, Ronald M.

Echinodermata: Echinoidea Serafy, D. Keith, and F. julian Fell

Echinodermata: Crinoidea Messing, Charles G., andJohn H. Dearborn

Erect Bryozoa Ryland, john S., and Peterj. Hayward

Marine Flora and Fauna of the Eastern United States

Cephalopoda Vecchione, Michael, Clyde F. E. Roper, and Michaelj. Sweeney

Copepoda, Cyclopoida: Archinotodelphyidae, Notodephyidae, and Ascidicolidae

Dudley, Patricia, and Paul L. lllg

Dicyemida Short, Robert B.

Platyhelminthes: Monogenea Hendrix, Sherman S.

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

COM 73 50670

COM 73 50888

COM 74 50019

COM 74 50014

COM 74 50487

COM 7451194

PB 257987

PB 261 839

PB 268 036

PB 268 714

BP 273 062

BP 274999

BP 280 040

BP 281969

BP 222 923

BP 285538

BP 296 460

BP 297 676

BP 124 520

BP 225 053

BP 219 387

PB124735

PC A03/MF AOI

PB 86156395

PB91173013

PB 89189583

PB 91 154179

PB 92118884

PB 95136222