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NOAA Technical Report NMFS Circular 413

Marine Flora and Fauna of
the Northeastern United States
Crustacea: Branchiura

Roger F. Cressey
May 1978

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

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NOAA Technical Report Circular 413

Marine Flora and Fauna of
the Northeastern United States.
Crustacea: Branchiura

Roger F. Cressey

May 1978

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U.S. DEPARTMENT OF COMMERCE

Juanita M. Kreps. Secretary

National Oceanic and Atmospheric Administration

Richard A. Frank, Administrator

National Marine Fisheries Service

For Sale by the Superintendent of Documents, U.S. Government Printing Office
Washington, D.C. 20402 Stock No. 003-0I 7-00419-8

FOREWORD

This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna of the
Northeastern United States." This subseries will consist of original, illustrated, modern
manuals on the identification, classification, and general biology of the estuarine and coastal
marine plants and animals of the northeastern United States. Manuals will be published at
irregular intervals on as many taxa of the region as there are specialists available to collaborate
in their preparation.

The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the
Woods Hole Region," edited by R. I. Smith, published in 1964, and produced under the auspices
of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. Instead
of revising the "Woods Hole Keys," the staff of the Systematics-Ecology Program decided to ex-
pand the geographic coverage and bathymetric range and produce the keys in an entirely new set
of expanded publications.

The "Marine Flora and Fauna of the Northeastern United States" is being prepared in
collaboration with systematic specialists in the United States and abroad. Each manual will be
based primarily on recent and ongoing revisionary systematic research and a fresh examination
of the plants and animals. Each major taxon, treated in a separate manual, will include an in-
troduction, illustrated glossary, uniform originally illustrated keys, annotated check list with in-
formation when available on distribution, habitat, life history, and related biology, references to
the major literature of the group, and a systematic index.

These manuals are intended for use by biology students, biologists, biological oceano-
graphers, informed laymen, and others wishing to identify coastal organisms for this region. In
many instances the manuals will serve as a guide to additional information about the species or
the group.

Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is
planned to include organisms from the headwaters of estuaries seaward to approximately the
200-m depth on the continental shelf from Maine to Virginia, but may vary somewhat with each
major taxon and the interests of collaborators. Whenever possible representative specimens
dealt with in the manuals will be deposited in the reference collections of major museums in the
region.

After a sufficient number of manuals of related taxonomic groups have been published, the
manuals will be revised, grouped, and issued as special volumes. These volumes will thus con-
sist of compilations of individual manuals within phyla such as the Cnidaria, Arthropoda, and
Mollusca, or of groups of phyla.

CONTENTS

Introduction I

Glossary 1

Key to the Branchiura of northeastern United States 3

Annotated systematic list 8

Selected bibliography 8

Systematic index 9

Acknowledgments 10

Coordinator's comments 10

The National Marine Fisheries Service (NMFS) does not approve, rec-
ommend or endorse any proprietary product or proprietary material
mentioned in this publication. No reference shall be made to NMFS, or
to this publication furnished by NMFS, in any advertising or sales pro-
motion 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.

in

Marine Flora and Fauna of the Northeastern
United States. Crustacea: Branchiura

ROGER F.CRESSEY'

ABSTRACT

Eleven species of Argulus are known from the northeastern United States. An illustrated key and
an annotated list of these species with notes on their hosts and distribution within and without the
study area are included. New host records are included.

INTRODUCTION

The class Branchiura, once considered a subgroup of
the class Copepoda, consists of a single order (Ar-
gulidea) and two families (Argulidae and Dipteropel-
tidae); the first with three genera (Argulus, Chonopeltis,
and Dolops) and the latter with two (Dipteropeltis and
Talaus). Argulus is cosmopolitan in distribution and the
only genus reported from North America and Europe.

Members of the class (commonly known as "fish lice")
are parasitic on freshwater and coastal marine fishes.
They are characterized by the presence of compound
eyes, prehensile antennae, and the second maxillae
transformed into sucker disks (except Dolops). The
feeding mechanism consists of a long preoral stylet or
"string" and a posterior mouth tube (Fig. 1). The
parasite repeatedly pierces the host tissue with the stylet
and probably uses the mouth tube to ingest fluids oozing
from the resultant wounds.

Eggs are attached to substrate objects (rocks, vegeta-
tion, etc.). Development is direct (newly hatched forms
resemble the adult). Tokiota (1936) described seven lar-
val stages and several subsequent molts for Argulus
japonicus, and he reported that sexual maturity was
achieved in about 30 days. The sexes can be distin-
guished in the first larval stage. Adult males are general-
ly smaller than females and can be distinguished from
them by the presence of a pair of elongate testes within
the abdomen along each side of the midline and by the
presence of sclerotized processes on the basal segments of
thoracic legs 2, 3, and 4. Females possess a pair of round-
ed spermathecae in the proximal third of the abdomen;
the basal portions of the legs are unmodified.

'Smithsonian Institution. U.S. National Museum of Natural History,
Washington. DC 20560.

Argulids are "loosely" associated with their hosts and
quickly drop off when the fish is removed from water.
Consequently, fish should be examined as soon after cap-
ture as possible. The parasites are found on the body sur-
face, in the gill chamber, and in the mouth of the host.
Argulids are often collected in plankton samples as well,
due to their transient attachment to their hosts. Collec-
tions should be preserved in 70% ethyl alcohol; 5% For-
malin is acceptable for short-term preservation. Where
Formalin is used, the material should be transferred to
alcohol as soon as possible as Formalin tends to decal-
cify materials over a long period of time. It may be neces-
sary to clear heavily pigmented specimens in a warm
solution of 10% KOH for 15 min or more in order to study
them microscopically. Small or less pigmented
specimens may be cleared in lactic acid for study. To
examine the respiratory area it may be necessary to
dissect one side of the carapace from the rest of the
body.

GLOSSARY

Carapace dorsolateral plate covering cephalothorax.
Respiratory areas two areas on each ventrolateral part

of the carapace devoid of sclerotization and open to the

overlaying tissues.
Scales short integumental processes with blunt or

broadly rounded tips occurring singly or in groups.
Spines short integumental processes with pointed tips

occurring singly or in groups.
Sucker modified second maxilla, seen ventrally, one on

each side of the oral area.
Sucker rod supporting structures composed of a series

of sclerotized processes radiating outwardly in the

membrane fringing the outer edge of the sucker.

preoral sting
antennae

sucker rod area
sucker

mouth tube
maxilla

respiratory areas

developing eggs

spermatheca
abdomen

Figure I .— Argulus alosae, female, ventral view, labeled.

KEY TO THE BRANCHIURA OF NORTHEASTERN UNITED STATES

Smaller respiratory area mostly or entirely anterior to larger (specimens "mostly anterior'
have sucker rods of 5-7 elements)

Figure 2. — Respiratory areas of: a. Argulus japonicus; b. A. chesapeakensis; c. A. laticauda; d. A. megalops; e. A. alosae;

f. A. funduli.

Smaller respiratory area mostly or entirely lateral to larger (specimens "mostly lateral" have

sucker rods of 2 elements) 7

a

Figure 3. — Respiratory areas of: a. Argulus versicolor; b. A. catastomi; c. A. appendiculosus; d. A. stizo-

8tethi; e. A. maculosus.

- i .' i Mouth tube with spines or scales at base .

2 (J) Mouth tube without spines or scales at base

W » j

a b c

Figure ■!.— Mouth tubes of: a. Argulus japonicus; b. A. alosae; c. A. megalops.

3(2) More than 15 elements in sucker rods Argulus funduli

Figure 5. — Sucker rods of Argulus funduli.

3 (2) Less than 15 elements in sucker rods 4

4 (3)

Mouth tube with spines at base
(see Fig. 4b). Sucker rods as in
Figure 6 Argulus alosae

Figure 6. — Sucker rods of Argulus alosae.

Figure 7.— Sucker rods of Argulus megalops.

4 (3) Mouth tube with scales at base (see Fig. 4c).

Sucker rods as Figure 7 Argulus megalops

1 1 I &

5(2) More than 5 elements in sucker rods . . . Argulus japonicus

Figure 8. — Sucker rods of Argulus japonicus.

5 (2) Less than 6 elements in sucker rods 6

6(5) Antennal spines nearly as wide as long. Sucker rods of 4-5 elements Argulus laticauda

Figure 9. — Argulus laticauda: a. antennae;
b. sucker rods.

6(5) Antennal spines much longer than wide. Sucker rods of 3-4 elements Argulus chesapeakensis

Figure 10. — Argulus chesapeakensis:
a. antennae; b. sucker rods.

7(1) More than 8 elements in sucker rods 8

7 (1) Less than 8 elements in sucker rods 9

8 (7) More than 12 elements in sucker rods Arbulus stizostithi

Figure 11. — Sucker rods of Argulus stizostethi.

(7) Less than 12 elements in sucker rods , ARgulus catostomi

Figure 12. — Sucker rods of Argulus catostomi.

9 (7) More than 3 elements in sucker rods

Argulus versicolor

Figure 13. — Sucker rods of Argulus versicolor.

9 ( 7) Less than 3 elements in sucker rods

10

10 (9) Basal element of sucker rod longer than terminal

Argulus maculosus

Figure 14. — Sucker rods of Argulus maculosus.

10 (9) Basal element of sucker rod
shorter than terminal . . .
Argulus appendiculosus

Figure 15. — Sucker rods of Argulus appendiculosus.

ANNOTATED SYSTEMATIC LIST

The 11 species of Argulus known from the north-
eastern United States are arranged alphabetically. Notes
on the distribution and hosts of each species within and
without the study area are included. Comments on
ecology and frequency are included wherever appro-
priate. An asterisk denotes new host record.

Argulus alosae Gould 1841. Distribution: east coast of
North America from Nova Scotia to the Gulf coast of
Texas. Hosts: Microgadus tomcod; Tautogolabrus
adspersus; Alosa pseudoharengus; Dorosoma
cepedianum; Clupea harengus; Cynoscion nebulosus;
Gasterosteus sp. [originally reported as G. bispinosus
from the Gulf of St. Lawrence (Wilson, 1902)];
Strongylura marina; Rhopilema verrilli*
(medusa). This species has been reported from
several species of inshore marine fish but its repeated
occurrence on clupeids (Alosa, Clupea, and Dorosoma)
may indicate a preference for that host family.

Argulus appendiculosus Wilson 1907. Distribution:
northern half of United States from Vermont to Vir-
ginia west to Wyoming and also Texas and Loui-
siana. Hosts: Catostomus catostomus; Ictiobus
cyprinellus, I. bubalus; Ictalurus punctatus, I. melas,
I. nebulosus; Amia calva; Micropterus salmoides;
Morbne (Roceus) chrysops; Perea flavescens; Cy-
prinus carpio; Stizostedion vitreum; Dorosoma
cepedianum; Lepisosteus osseus*. Argulus appen-
diculosus has so far only been reported from fresh-
water, but since one of its hosts (Dorosoma) is found in
coastal marine waters as well, the parasite may also be
eventually found there.

Argulus catostomi Dana and Herrick 1837. Distribu-
tion: northern United States from Massachusetts west
to Minnesota. Hosts: Catostomus commersoni, C.
catostomus; Hypentelium nigricans; Erimyzon sucet-
ta; Cyprinus carpio; Notemigonus crysoleucas. So far
this parasite has been found only in freshwater and on
members of the families Catostomidae and Cy-
prinidae.

Argulus chesapeakensis Cressey 1971. Distribution:
southeast coast of United States from Chesapeake Bay
south to Sapelo Island, Ga. Hosts: Opsanus tau;
Archosargus probatoeephalus*; Arius felis*; Mugil
cephalus*; Gobiosoma bosci*; Paralichthys dentatus;
Dasyatis americana*; Pteroplatea maclura*; Rachy-
centron canadum. This species has so far been col-
lected only from the southeast coast of the United
States. It is closely related to A. laticauda, found along
the northeast coast.

Argulus funduli Kroyer 1863. Distribution: east coast of
North America from New Brunswick, Canada, south
to the mouth of the Neches River, Texas, collected at
several localities between. Hosts: Menidia notata;
Lagodon rhomboides; Pseudopleuronectes ameri-
canus; Cyprinodon variegatus; Fundulus grandis, F.
hptcroclitus, F. majalis, F. ocellaris. This species is
common along the entire east coast of the United

States and is most often associated with marine
cyprinodontids.

Argulus japonicus Thiele 1900. Distribution: entire
United States. Hosts: Carassius auratus (goldfish).
This argulid is an introduced species brought into the
United 'States from the Orient on aquarium fish. It is
very common anywhere goldfish are found. No records
from marine waters to date.

Argulus laticauda Smith 1873. Distribution: northeast
coast of United States from New England south to
Long Island Sound. Hosts: Prionotus sp.; Anguilla
rostrata; Pseudopleuronectes americanus. Common
on the American eel and winter flounder of the north-
east coast.

Argulus maculosus Wilson 1902. Distribution: eastern
half of United States and southeastern Canada as far
west as Iowa and collected in northern and southern-
most states. Hosts: Ictalurus natilis, I. nebulosus;
Amia calva; Ambloplites rupestris; Erimyzon sucetta;
Esox sp.; Umbra limi; Lepisosteus osseus*. This common
argulid has so far been collected only in freshwater.

Argulus megalops Smith 1873. Distribution: east coast
of North America from New Brunswick, Canada, to
Florida. Hosts: a wide variety of inshore marine
fishes (recorded from 16 species) with no indications as
to host preferences.

Argulus stizostethi Kellicott 1880. Distribution: north-
eastern quarter of the United States from Iowa and
Minnesota east to New England and New Brunswick,
Canada. Hosts: Salvelinus fontinalis; Stizostedion
vitreum, S. canadense; Acipenser fulvescens; Esox
masquinongy; AIoso sapidissima; Gasterosteus sp.;
Notropis sp.; Coregonus sp.; Dorosoma cepedia-
num*. This parasite seems to be restricted to those
states near the Great Lakes and St. Lawrence River.
All collections are from freshwater except that from
Dorosoma from Woods Hole, Mass.

Argulus versicolor Wilson 1902. Distribution: scattered
throughout the eastern half of the United States.
Recorded in Massachusetts, Maryland, Indiana,
Georgia, and Texas. Hosts: Esox niger; Ambloplites
sp.; and "perch." So far there are no marine records
for this species, but four of the recorded states are
marine coastal.

SELECTED BIBLIOGRAPHY

CRESSEY. R.F.

1972. The genus Argulus (Crustacea: Branchiura) of the United
States. Bioat of Freshwater Ecosystems. Environmental Protec-
tion Agency, Identification Manual No. 2, 14 p.
MEEHAN, O. L.

1940. A review of the parasitic Crustacea of the genus Argulus in
the collections of the United States National Museum. Proc.
U.S. Nat. Mus. 88:459-522.
TOKIOKA. T.

1936. Larval development and metamorphosis of Argulus japoni-
cus. Mem. Coll. Sci. Kyoto Imp. Univ. ser. B, 12:93-114.
WILSON. C. B.

1903, North American parasitic copepods of the family Argulidae
with ;i bibliography of the group and a systematic review of all
known species. Proc. U.S. Nat. Mus. 25:635-742.

SYSTEMATICS INDEX

Acipenser fuluescens 8

Alosa 8

pseudoharengus 8

sapidissima 8

Ambloplites

rupestris 8

sp 8

Amia calva 8

Anguilla rostrata 8

Archosargus probatucephalus 8

Argulidae 1

Argulus

alosae 2, 3, 4, 8

appendiculosus 3, 7, 8

catostomi 3, 6, 8

chesapeakensis 3, 5, 8

funduli 3, 4, 8

japonicus 1, 3, 4, 5, 8

laticauda 3, 5, 8

maculosus 3, 7, 8

megalops 3, 4, 5, 8

stizostethi 3, 6, 8

versicolor 3, 7, 8

Arius felis 8

Carassius auratus 8

Catostomidae 8

Catostomus

catostomus 8

commersoni 8

Chonopeltis 1

Clupea 8

harengus 8

Coregonus sp 8

Cynoscion nebulosus 8

Cyprinidae 8

Cyprinodon variegatus 8

Cyprinus carpio 8

Dasyatis americana 8

Dipteropeltidae 1

Dipteropeltis 1

Dolops 1

Dorosoma 8

cepedianum 8

Erimxson sucetta 8

Esox

masquinongy 8

niger 8

sp 8

Fundulus

grandis 8

heteroclitis 8

majalis 8

ocellaris 8

Gasterosteus sp 8

Gobiosoma bosci 8

Hypentelium nigricans 8

Ictalurus

melas 8

natalis 8

nebulosus 8

punctatus 8

Ictiobus

bubalus 8

cyprinellus 8

Lagodon rhomboides 8

Lepisosteus osseus 8

Menidia notata 8

Microgadus torn cod 8

Micropterus salmoides 8

Morone chrysops 8

Mugil cephalus 8

Notemigonus crysoleucas 8

Notropis sp 8

Opsanus tau 8

Paralichthys dentatus 8

Perca flavescens 8

Prionotus sp 8

Pseudopleuronectes americanus 8

Pteroplatea maclura 8

Rachycentron canadum 8

Rhopilema verrilli 8

Salvelinus fontinalis 8

Stizostedion

canadense 8

vitreum 8

Strongylura marina 8

Talaus 1

Tautogolabrus adspersus 8

Umbra limi 8

ACKNOWLEDGMENTS

Preparation of the "Marine Flora and Fauna of the Northeastern
United States" is being coordinated by the following Board:

Coordinating Editor:

Melbourne R. Carriker. College of Marine Studies, Univer-
sity of Delaware. Lewes. DE 19958.

Editorial Advisers:

Marie B. Abbott. Marine Biological Laboratory, Woods
Hole, Mass.

Arthur G. Humes. Bostom University Marine Program,
Marine Biological Laboratory, Woods Hole, Mass.

Wesley N. Tiffney. Department of Biology. Boston Univer-
sity. Boston. Mass.

Ruth D. Turner. Museum of Comparative Zoology, Har-
vard University. Cambridge, Mass.

Roland L. Wigley, National Marine Fisheries Service
Northeast Fisheries Center, NOAA, Woods Hole, Mass.

Robert T. Wilce, Department of Botany, University of Mas-
sachusetts, Amherst, Mass.

The Board, which established the format for the "Marine
Flora and Fauna of the Northwestern Unites States," invites
systematists to collaborate in the preparation of manuals,
reviews manuscripts, and advises the Scientific Editor of the
National Marine Fisheries Service.

The illustrations were done by Hillary Boyle. The manu-
script was critically reviewed by Roland Wigley and Arthur
Humes. The materials and records embodied in this paper are
housed in the National Museum of Natural History, Washing-
ton, D.C.

COORDINATING EDITOR'S COMMENTS

Publication of the "Marine Flora and Fauna of the North-
eastern LTnited States" is most timely in view of the growing uni-
versal emphasis on environmental work and the urgent need for
more precise and complete identification of coastal organisms
than has been available. It is mandatory, whenever possible,
that organisms be identified accurately to species. Accurate
scientific names unlock the great quantities of biological infor-
mation stored in libraries, obviate duplication of research al-
ready done, and often make possible prediction of attributes of
organisms that have been inadequately studied.

Roger Cressey started working with parasitic copepods and
branchiurans as a graduate student at Boston University in
1958. After receiving his doctoral degree in 1965, he joined the
staff of the National Museum of Natural History, Smithsonian

Institution. His studies on parasitic crustaceans have resulted in
over 40 publications. He has conducted field work in the Indian
Ocean, Madagascar, southeastern Pacific Ocean, and Gulf of
Mexico off the west coast of Florida.

Preparation of this manual was supported in part by a grant
from the Environmental Protection Agency to the Editorial
Board of the "Marine Flora and Fauna of the Northeastern
United States." Work on the "Marine Flora and Fauna of the
Northeastern United States" by the Coordinating Editor is sup-
ported by the College of Marine Studies, University of Dela-
ware.

Manuals are available for purchase from the Superintendent
of Documents, U.S. Government Printing Office, Washington,
DC 20402. The manuals so far published in the series are listed
below:

Marine Flora and Fauna of the Northeastern United States:

Circular

COOK. DAVID G., AND RALPH 0. BRINKHURST. Annelida: Oligochaeta.

BORROR. ARTHUR C. Protozoa: Ciliophora.

MOUL, EDWIN T. Higher Plants of the Marine Fringe.

McCLOSKEY, LAWRENCE R. Pycnogonida.

MANNING, RAYMOND B. Crustacea: Stomatopoda.

WILLIAMS, AUSTIN B. Crustacea: Decapoda.

POLLOCK, LELAND W. Tardigrada.

LARSON, RONALD J. Cnidaria: Scyphozoa.

CAYALIERE, A. R. Higher Fungi: Ascomycetes, Deuteromycetes, and Basiomycetes.

COULL, BRUCE C. Copepoda: Harpacticoida.

CUTLER, EDWARD B. Sipuncula.

PAWSON, DAVID L. Echinodermata: Holothuroidea.

HO, JU-SHEY. Copepoda: Lernaeopodida.

HO, JU-SHEY. Copepoda: Cyclopoids Parasitic on Fishes.

( RKSSEY, ROGER F. Crustacea: Branchiura.

374
378
384
386
387
389
394
397
398
399
403
405
406

* U.S. GOVERNMENT PRINTING OFFICE: 1978—796-751/18 REGION 10

10

388. Proceedings of the first U.S. -Japan meeting on aquaculture at
Tokyo, .Japan, October 18-19, 1971. William N. Shaw (editor). (18
papers, 14 authors.) February 1974, iii + 133 p. For sale by the
Superintendent of Documents, U.S. Government Printing Office,
Washington, D.C. 20402.

389. Marine flora and fauna of the northeastern United States.
Crustacea: Decapoda. By Austin B. Williams. April 1974, iii + SO p., Ill
figs. For sale bv the Superintendent of Documents, U.S. Government
Printing Office,' Washington, D.C. 20402.

.190. Fishery publications, calendar year 197.1: Lists and indexes. By
Mary Ellen Engett and Lee C. Thorson. September 1974, iv + 14 p., 1 fig.
For sale by the Superintendent of Documents, U.S. Government Printing
Office, Washington, D.C. 20402.

391. Calanoid copepods of the genera Spinocalanus and Mimocalanus
from the central Arctic Ocean, with a review of the Spinocalanidae. By
David M. Damkaer. .June 1975, x + 88 p., 225 figs., 4 tables. For sale

by the Superintendent of Documents, U.S. Government Printing Office.
Washington, D.C. 20402.

392. Fishery publications, calendar year 1974: Lists and indexes. By
Lee C. Thorson and Mary Ellen Engett. June 1975, iv + 27 p., 1 tig.

393, Cooperative Gulf of Mexico estuarine inventory and study — Texas:
Area description. Bv Richard A. Diener. September 1975, vi + 129 p.,
55 figs., 26 tables.

194. Marine Flora and Fauna of the Northeastern United States. Tar-
digrada. Bv Leland W. Pollock. Mav 1976, iii + 25 p., tigs. For sale
In the Superintendent of Documents, U.S. Government Printing Office,
Washington. D.C. 20402.

395. Report of a colloquium on larval fish mortality studies and their
relation to fishery research, January 1975. By John R. Hunter. May
1976, iii + 5 p. For sale by the Superintendent of Documents, U.S.
Government Printing Office, Washington, D.C. 20402.

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