NOAA Technical Report NMFS 96 January 1991

Marine Flora and Fauna

of the Eastern United States

Copepoda, Cyclopoida:

Archinotodelphyidae, Notodelphyidae,

and Ascidicolidae

Patricia L. Dudley

Paul L. Illg

U.S. Department of Commerce
NOAA Technical Report NMFS _

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69. Environmental quality and aquaculture systems: Proceedings of the

thirteenth US.-Japan meeting on aquaculture, Mie, Japan, October 24-25,

1984, edited by Carl J. Sindermann. October 1988, 50 p.

70. New and innovative advances in biology/engineering with potential

for use in aquaculture: Proceedings of the fourteenth U .-Japan meeting

on aquaculture, Woods Hole, Massachusetts, October 16-17, 1985, edited

by Albert K. Sparks. November 1988, 69 p.

71. Greenland turbot ReiTlho.rdJius hippoglo55oides of the eastern Bering Sea

and Aleutian Islands region, by Miles S. Alton, Richard G. Bakkala, Gary

E. Walters, and Peter T. Munro. December 1988, 31 p.

72. Age determination methods for northwest Atlantic species, edited

by Judy Penttila and Louise M. Dery. December 1988, 135 p.

73. Marine flora and fauna of the Eastern United States. Mollusca:

Cephalopoda, by Michael Vecchione, Clyde F. E. Roper, and Michael

J. Sweeney. February 1989, 23 p.

74. Proximate composition and fatty acid and cholesterol content of 22

species of northwest Atlantic finfish, by Judith Krzynowek, Jenny Murphy, Richard S. Maney, and Laurie J. Panunzio. May 1989, 35 p.

75. Codend selection of winter flounder PseudfJpleuronectes americallus, by

David G. Simpson. March 1989, 10 p.

76. Analysis of fish diversion efficiency and survivorship in the fish return

system at San Onofre Nuclear Generating Station, by Milton S. Love,

Meenu Sandhu, Jeffrey Stein, Kevin T. Herbinson, Robert H. Moore,

Michael Mullin, and John S. Stephens Jr. April 1989, 16 p.

77. Illustrated key to the genera of free-living marine nematodes of the

order Enoplida, by Edwin J. Keppner and Armen C. Tarjan. July 1989,

26 p.

78. Survey of fishes and water properties of south San Francisco Bay,

California, 1973-82, by Donald E. Pearson. August 1989, 21 p.

79. Species composition, distribution, and relative abundance of fishes

in the coastal habitat off the southeastern United States, by Charles A.

Wenner and George R. Sedberry. July 1989, 49 p.

80. Laboratory guide to early life history stages of northeast Pacific fishes,

by Ann C. Matarese, Arthur W. Kendall Jr., Deborah M. Blood, and

Beverly M. Vinter. October 1989, 651 p.

81. Catch-per-unit-effort and biological parameters from the Massachu~

setts coastal lobster (Homarus ameruaTlUS) resource: Description and Trends,

by Bruce T. Estrella and Daniel J. M Kiernan. September 1989, 21 p.

82. Synopsis of biological data on the cobia RackyceTltroTl canadum (Pisces:

Rachycentridae), by Rosalie Vaught Shaffer and Eugene L. Nakamura.

December 1989, 21 p.

83. Celaphopods from the stomachs of sperm whales taken off California, by Clifford H. Fiscus, Dale W. Rice, and Allen A. Wolman. December 1989, 12 p.

84. Results of abundance surveys ofjuvenile Atlantic and Gulf menhaden, Brevoortia tyraTlTlus and B. patruTlus, by Dean W. Ahrenholz, James F.

Guthrie, and Charles W. Krouse. December 1989, 14 p.

85. Marine farming and enhancement: Proceedings of the Fifteenth

US.-Japan Meeting on Aquaculture, Kyoto, Japan, October 22-23, 1986,

edited by Albert K. Sparks. March 1990, 127 p.

86. Benthic macrofauna and habitat monitoring on the continental shelf

of the northeastern United States. 1. Biomass, by Frank Steimle. February 1990, 28 p.

87. Life history aspects of 19 rockfish species (Scorpaenidae: Sebastes) from

the Southern California Bight, by Milton S. Love, Pamela Morris, Merritt McCrae, and Robson Collins. February 1990, 38 p.

88. Early-life-history profiles, seasonal abundance, and distribution of

four species of clupeid larvae from the northern Gulf of Mexico, 1982 and

1983, by Richard F. Shaw and David L. Drullinger. April 1990, 60 p.

89. Early-life-history profiles, seasonal abundance, and distribution of

four species of carangid larvae off Louisiana, 1982 and 1983, by Richard

F. Shaw and David L. Drullinger. April 1990, 37 p.

90. Elasmobranchs as living resources: Advances in the bi logy, ecology,

systematics, and the status of the fisheries, edited by Harold L. PrattJr.,

Samuel H. Gruber, and Toru Taniuchi. July 1990, 518 p.

91. Marine nora and fauna of the northeastern United State, Echinodermata: Crinoidea, by Charles G. Messing and John H. Dearborn. August

1990, 30 p.
NOAA Technical Report NMFS 96

Marine Flora and Fauna

of the Eastern United States

Copepoda, Cyclopoida:

Archinotodelphyidae, Notodelphyidae,

and Ascidicolidae

Patricia L. Dudley

Paul L. Illg

January 1991

u.s. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

John A. Knauss, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

William W. Fox Jr., Assistant Administrator for Fisheries
Foreword _

This ~OAA Technical Report NMFS is part of the subseries "Marine Flora and Fauna

ofthe Eastern United States" (formerly "Marine Flora and Fauna ofthe Northeastern

United States"), which consists of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants

and animals of the eastern United States. The manuals are published at irregular intervals on as many taxa of the region as there are specialists available to collaborate

in their preparation. These manuals are intended for use by students, biologists,

biological oceanographers, informed laymen, and others wishing to identify coastal

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

speCies or groups.

The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates

of the Woods Hole Region," edited by R.I. Smith, and produced in 1964 under the

auspices of the Systematics Ecology Program, Marine Biological Laboratory, Woods

Hole, Massachusetts. Geographic coverage of the "Marine Flora and Fauna of the

Eastern United States" is planned to include organisms from the headwaters of estuaries

seaward to approximately the 200-m depth on the continental shelf from Maine to

Florida, but can vary somewhat with each major taxon and the interests of collaborators.

Whenever possible, representative specimens dealt with in the manuals are deposited

in the reference collections of major museums.

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

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

is based primarily on recent and ongoing revisionary systematic research and a fresh

examination of the plants and animals. Each manual, treating a separate major taxon,

includes an introduction, illustrated glossary, uniform originally illustrated keys, 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.

Manuals are available from the National Technical Information Service, United

States Department of Commerce, 5285 Port Royal Road, Springfield, VA 22161, either

as a paper copy or microfiche, for a charge. Manuals are not copyrighted, and so can

be photocopied from the NOAA Technical Report NMFS Circulars and Reports

available in most major libraries and listed at the end of this manual.

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

recommend or endorse any proprietary product or proprietary

material mentioned in this publication. No reference shall be made

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

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Text printed on recycled paper

ii
Marine Flora and Fauna of the Eastern United States

Copepoda, Cyclopoida: Archinotodelphyidae, Notodelphyidae,

and Ascidicolidae-Associates of Ascidians

PATRICIA L. DUDLEY

Department of Biological Sciences

Barnard College, Columbia University

New York, NY 10027

PAUL L. ILLG

Department of Zoology

University of Washington

Seattle, WA 98195

ABSTRACT

This manual includes an introduction to the general biology, a selected bibliography, and an

illustrated key to 11 genera and 17 species of copepods of the Crustacea, Subclass Copepoda,

Order Cyclopoida, Families Archinotodelphyidae, Notodelphyidae and Ascidicolidae, associated

with ascidians from the Atlantic Coast of the United States. Species distributed from the Gulf

of Maine to Long Island Sound are emphasized. An annotated systematic list, with statements -

of the world distribution and new records of association with hosts, and a systematic index are

also provided.

Introduction _

The 8000 to 10,000 species of Crustacea belonging to the

Subclass Copepoda occupy many niches in the ecosphere

(Bowman and Abele 1982). Copepods are small, often less

than 0.5 mm and rarely exceeding 10 mm in length

(Kaestner 1970). They lack compound eyes and a carapace,

although a cephalic head shield drawn out as an anteromedial rostrum is characteristic. Their segmented bodies

are usually divided into a broad forebody and a narrow

hindbody, separated by a major body articulation. In some,

however, the taper of the body is more gradual. The genital

apertures are on the last thoracic segment. There are 5 pairs

of cephalic appendages and up to 7 pairs of thoracic appendages, the structure and numbers serving as important

diagnostic features in classification. The postgenital abdominal segments of the hindbody never bear appendages,

but two caudal rami extend from the posteriormost

segment which includes the telson. The development of a

copepod includes a series of naupliar and copepodid

stages.

Most of the copepods are benthic, demersal, or planktonic free-living organisms in freshwater, estuarine or

marine habitats. Some are free-living in such semiterrestrial

habitats as wet moss, moist soils or leaflitter (Reid 1986).

Many species have developed the capability of living as

ectosymbionts or endosymbionts of other marine invertebrates or freshwater and marine fish (Gotto 1979;

Kabata 1979). As might be expected, the bodies and appendages of copepods have undergone extensive adaptive

morphological changes in concert with their radiation into

so many different habitats. Symbiotic species, for example,

with their needs to emphasize modes of attachment to a

host and enhanced reproduction, may lack many of the

defining characters of the Copepoda given above and their

affinities with the group may only be determined by studying their developmental stages (Kaestner 1970). Gotto

(1979), Kabata (1979) and Marcotte (1982, 1986) discuss

the evolution ofthe diversity of body forms in the Copepoda

and speculate on the different modes of feeding, locomotion, reproduction, and development required for their

diverse life styles.
2 NOAA Technical Report NMFS 96 _

Cephalosome

Metasome

Urosome

s

S

LQ

o

8

Figure 1

Pararchinotodelphys gumeyi, female; (A) habitus, dorsal view, showing divisions of body; (B) urosome, ventral view, showing segmental

composition, caudal rami and fifth legs; (C) antenna with terminal prehensile hook; (D) maxilliped; (E) first leg with components

labelled. As = anal segment; Al = first antenna (antennule); Cr = caudal ramus; Gc = genital complex; L5 = fifth leg; Ph =

prehensile hook.
__________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 3

The order Cyclopoida is one of the 8 orders of the

subclass Copepoda (Kabata 1979; Marcotte 1982) and contains about 450 species (Bowman and Abele 1982). In this

classification, still controversial, the other orders are the

orders Calanoida, Harpacticoida, Misophrioida, Mormonilloida, Monstrilloida, Poecilostomatoida, and Siphonostomatoida. The copepods in the orders Cyclopoida and

Poecilostomatoida differ from those in the other orders by

universally possessing uniramous antennae (the second pair

of appendages ofthe cephalon) rather than biramous antennae. While some copepods in the order Siphonostomatoida

also have uniramous antennae, others have biramous

antennae with extremely reduced exopodites. Cyclopoida

and Poecilostomatoida differ from all Siphonostomatoida

by having an open buccal cavity, rather than a siphonlike

extension, containing buccal stylets and formed by fusions

of the labrum and labium. The major differentiating characters between the orders Cyclopoida and Poecilostomatoida relate to the structure and function of the mandibles;

the cyclopoids are basically gnathostomous (having biting

or chewing mandibles) rather than poecilostomous (having falcate nonbiting mandibles or none) (Kabata 1979).

In the following brief diagnostic discussions, significant

anatomical terms used in the keys are in boldface type and

are explained in the text; an index to these is provided on

p. 38.

The order Cyclopoida is an assemblage of free-living

benthic, demersal or planktonic freshwater and marine

species and many symbiotic species. Three of the families

in the Cyclopoida, the families Archinotodelphyidae, Notodelphyidae, and Ascidicolidae, are marine symbionts in

ascidians (there are a few records of occurrences in other

invertebrate hosts) and are the subjects of this manual. The

other three families which we include in the Cyclopoida

are free-living species: the family Cyclopinidae, benthic or

demersal in marine littoral zones; the family Cyclopidae,

benthic, demersal or planktonic in freshwater or, less commonly, in the marine littoral zone; and the family Oithonidae, planktonic in the marine littoral and oceanic pelagic

zones.

In general, an adult female cyclopoid copepod has its

body segments organized as in most Crustacea into explicit

functional regions, the tagmata. The term segment is used

here to denote a major section of the body distinguished

by clear articulations or by the presence of one pair of

bilaterally arranged appendages. In the least modified

cyclopoid females, as in females of the family Cyclopinidae

or Archinotodelphyidae (Fig. lA), the tagmata consist of

a cephalosome bearing paired appendages consisting, from

anterior to posterior, of the pre-oral cephalic antennules

and antennae, the post-oral cephalic mandibles, maxillules

and maxillae, and the thoracic maxillipeds, a metasome

(Fig. lA) of four free thoracic segments, each with a pair

of biramous swimming legs, and a five-segmented urosome

(Fig. 1, A and B). The urosome, set off from the metasome by the major body articulation, is composed of the

segment ofthe reduced fifth legs ( = the sixth thoracic segment), a genital complex ( = the seventh thoracic segment

fused with the first abdominal segment) which has a ventral gonopore and lateral or dorsolateral oviducal apertures,

from which ovisacs are suspended at times of reproduction, and three additional abdominal segments. The least

modified males of the order Cyclopoida are distinguished

from females in their body segmentation only by having

six segments in the urosome rather than five. Their genital

segment ( = the seventh thoracic segment), which bears

small sixth legs not present in females, remains free of the

first abdominal segment. In both sexes, the terminal abdominal element ( = the anal segment) includes the telson

with its pair of posterior caudal rami and dorsally directed

anus. The caudal rami are flattened protrusions, each of

which has four terminal, plumose setae as well as one small

dorsal and one small lateral seta.

The forebody, consisting of the cephalosome and the

metasome, is often referred to as the prosome (Gooding

1957) and is wider than the urosome. This arrangement

in cyclopoids, whereby the major body articulation falls

in front of the segment of the reduced fifth legs, is known

as the podoplean condition. This contrasts with the gymnoplean condition of copepods of the order Calanoida in

which the major body articulation falls posterior to the segment of the fifth leg and the urosome, therefore, bears no

appendages (Giesbrecht 1892; Marcotte 1982).

As might be expected from the long evolutionary history

of the Copepoda, none of the extant species of the order

Cyclopoida can be considered archetypical with respect to

its appendages. However, many investigators (Schellenberg 1922; Lang 1946, 1949; Lindberg 1952; Illg 1955,

1958; Dudley 1966) have recognized that species of the

family Cyclopinidae have more appendicular features that

might be referable to ancestral cyclopoids (plesiomorphic

characters) than representatives of any other family in the

order Cyclopoida. The most basic features ofthe cyclopoid

appendages which are listed below, therefore, draw upon

those characters which might be found in some species of

cyclopinid. It is recognized, though, (Illg 1955; Dudley

1966) that extant species of the Cyclopinidae actually show

mixtures of generalized and specialized characters.

In cyclopinids, the antennule is uniramous and has 10

to 26 segments. It is geniculate in males and has locking

distal articulations used in holding the female during

copulation. The uniramous antenna is 4-segmented, consisting of a 2-segmented protopodite and a 2-segmented

endopodite, armed apically with setae only. The buccal

cavity is open. There is a well-developed labrum anchored

anterior to the mouth, but only a small labium, possibly

represented in part by paired lobes called paragnaths

posterior to the mouth. The biramous mandible has a

2-segmented protopodite and a biramous palp with a

2-segmented endopodite and an exopodite of up to 4
4 NOAA Technical Report NMFS 96 _

A

Figure 2

Lateral views of representative females and males of Notodelphyidae; (A) Notodelphys monoseta, female; (B) Doropygopsis longicauda,

female; (C) Doropygus laticornis, female; (D) Pachypygus macer, female; (E) Gunenotophorus curvipes, female; (F) Doropygopsis longicauda,

male; (G) Doropygus laticornis, anamorphic male; (H) Doropygus laticornis, metamorphic male. Bp = broodpouch; GAl = geniculate

antennule; L5, L6 = fifth and sixth legs; Ss = spermatophoral sac.
__________________ Dudley and Illg: Copepoda, Cydopoida of the Eastern United States 5

segments. The mandible is gnathostomous with a medial,

coxopodal, gnathal plate which can work under the labrum.

The biramous maxillule has a 2-segmented protopodite

with medial setose endites as well as a lateral epipodite and

a 1-segmented endopodite and exopodite. The uniramous

maxilla is 6-segmented. The basal 3 segments have setose

endites and there is a claw, as well as setae, on the endite

of the third segment. The 3 small, distal segments, armed

with setae, are considered by some investigators to be an

endopodite, although there is no ontogenetic reason to consider that this appendage is biramous. The maxilliped of

the first thoracic segment of the body is uniramous and

has up to 7 segments and up to 18 setae apportioned to

the segments. The four pairs of thoracic, biramous swimming legs are similar in the Cyclopinidae and the Archinotodelphyidae (Fig. 2E). These appendages have flattened

protopodites, each with a clearly indicated coxopodite

and basipodite, the latter supporting the two rami, the

3-segmented exopodite and endopodite. The endopodite

is armed only with setae while the exopodite has lateral

and apicolateral spines and medial and apicomedial setae.

There is a quadrate plate, called the copula or intercoxal

plate, which links the contralateral legs on each metasomal

segment and allows the paired legs to serve as a single unit

during swimming. The uniramous, 2- to 3-segmented fifth

legs on the first urosomal segment have up to 7 setae. The

sixth legs, found only in males, consist of apposed subtriangular plates on the ventral surface of the second

urosomal segment. Each is armed with 3 setae.

Diagnoses and Relationships of Families

of Cyclopoid Ascidicolous Copepods _

Family Archinotodelphyidae

The family Archinotodelphyidae is the smallest family of

ascidicolous copepods, consisting of only three genera and

six species. These live in simple ascidians ofseveral families

(Hansen 1923; Lang 1949; Illg 1955; Monniot 1968, 1987),

and, paradoxically, one species, Nearchinotodelphys indicus

Ummerkutty (1960), in a boring bivalve mollusc. The

family is of interest with respect to the phylogeny of

Copepoda because of its suggested position evolutionarily

between the many marine representatives of the cyc1opoid

family Cyclopinidae and the now substantially numerous

species ofthe symbiotic family Notodelphyidae (Illg 1955;

Dudley 1966).

A major common feature of the family Archinotodelphyidae and the family Cyclopinidae is the generalized

cyc1opoid habitus or general aspect of bodily organization

of the females. In the northeastern United States, the

female of Pararchinotodelphys gurneyi Illg (1955) from the

ascidian Styela partita is a representative example in regard

to its habitus (Fig. 1, A and B). The urosome may, however, be variable among archinotodelphyid genera. Lang

(1949) reported the urosome offemale Archinotodelphys typicus

as having 6 segments. There is no broodpouch and the embryos are carried in external ovisacs.

The appendages of archinotodelphyids differ from those

of cyclopinids in the following ways: the antennule has 15

to 17 segments rather than 10 to 26 as in cyclopininds; the

antenna (Fig. 1C), although 4-segmented as in cyclopinids,

has fewer setae and an apical, articulated, prehensile hook,

which is absent in cyclopinids. The prehensile hook is the

only clear appendicular modification of archinotodelphyids

related to their symbiotic existence in ascidians. The maxillule, peculiarly, may be more specialized in cyc10pinids

than in archinotodelphyids by having fewer endites on the

protopodite and, in some cases, by lacking the epipodite

possessed by all archinotodelphyids. The maxilliped (Fig.

1D) has 3 to 5 segments and fewer setae than do cyclopinids. In no cyclopinid does the maxilliped have less than

5 segments and, in some species, may have as many as

7 segments.

Similarities in appendages between archinotodelphyids

and cyclopinids exist in the structure of the mandible, maxilla, and the metasomallegs (Fig. 1E), which conform in

all details of segmentation and have a similar armature.

In addition, the males, known only in Archinotodelphys

profundus Monniot and Nearchinotodelphys indicus Ummerkutty, have geniculate antennules as do cyclopinid males.

Family Notodelphyidae

The Copepoda of the family Notodelphyidae have been

almost universally described as inhabitants of ascidians

although some descriptions of species have given as their

provenance washings or mud from unspecified marine collections or collections from other colonial marine invertebrates that could be confused with compound ascidians.

One major criterion defines all females of the species of

the family Notodelphyidae: the formation of an internal

broodpouch (Fig. 2, A through E) within which the embryos develop until they are released for the free-living

phases ofthe life cycle. Another feature defines both males

and females ofthe family: the presence of a prehensile hook

on the apex of each uniramous antenna (Fig. 3, B and C).

Both of these are doubtlessly basic adaptations to help

adults or their developmental stages avoid the filter-feeding

entrapment mechanisms and the powerful flushing and

back-flushing which occur in the ascidian's branchial basket. These defining characteristics of the Notodelphyidae,

however, persist even in those highly modified species

which occupy loci in the ascidian host where they or their

embryos are not subject to filter-feeding entrapment.

The vividly colored eggs or embryos in the broodpouches

of females are easily seen when the ascidian hosts are

dissected and, largely because of this, adult females are

much better known than are the smaller, white or yellowish
6 NOAA Technical Report NMFS 96 _

males. Features of identification of notodelphyid genera

are usually based on females and the characters used in

the key in this manual are so restricted. The habitus is one

of the most important features for the identification of the

genus of a notodelphyid female. The presence ofthe broodpouch causes the bodies (Fig. 2, A through E) to deviate

to varying degrees from the habitus of the generalized

cyclopoid female. The least modified females of Notodelphyidae, living in the branchial baskets of solitary ascidians, are those of species of Notodelphys (Fig. 2A), which

are very active and can leave the ascidian under adverse

conditions. The broodpouch does show some bulging

laterally and dorsally and occupies both the fifth and sixth

thoracic segments ( = segments of the fourth and fifth legs).

Because the segment of the fifth leg is incorporated in the

pouch and the major body articulation falls behind it, the

urosome carries no legs and a secondary gymnopleaD condition exists, a peculiarity in the Cyclopoida.

As other examples, species of Doropygopsis (Fig. 2B),

Doropygus (Fig. 2C), and Pachypygus (Fig. 2D) are laterally

compressed and have large gibbous broodpouche8 that

occupy only the last metasomal segment (= segment of the

fourth legs). Because the fifth legs are found on the first

urosomal segment, species of these genera show the

podoplean condition. The development ofthis large pouch

causes the prosome in these females to tilt ventrally relative

to the urosome and, in all of these genera, a progression

of restrictions in motility accompanies its formation. No

adult females of species in these genera are known to leave

their hosts. In species in other genera, also found in branchial chambers of solitary ascidians, the broodpouch may

extend more anteriorly into the metasomal segments or the

body may be grossly inflated and a lobed broodpouch may

superficially cover much of the fused metasome as in

Gunenotophorus (Fig. 2E). Fifth legs are absent in species

of this genus.

Adult females of species which occupy cysts have the

most profound modifications of body form among notodelphyid associates of solitary ascidians. In females ofsuch

species as Scolecodes huntsmani, living in cysts in the subendostylar blood vessels ofits hosts (Illg 1958; Dudley 1968)

and Scolecimorpha joubini, living in cysts in the tunic wall

(Illg and Dudley 1961), the bodies are vermiform and show

much fusion ofsegments. Broodpouches are very capacious

and long, occupying only an extended sixth thoracic segment in S. huntsmani and the whole lengthened metasome

in S. joubini. But however much the bodies of notodelphyids

in cysts in solitary ascidians are modified, it is in the

notodelphyid associates of compound ascidians where

specialization reaches extremes. All of these species from

compound ascidians have bodies that are so changed

toward a vermiform or globular habitus that it is sometimes

difficult even to recognize that they are copepods. Among

these, the species that occupy the branchial baskets of

zooids are less modified than those that are found in the

common cloaca, postabdomens of zooids, or the matrix of

the colony (Illg and Dudley 1961,1965). Representatives

of these extremely modified notodelphyids from cysts in

solitary ascidians or from compound ascidians have not

as yet been collected from the Atlantic coast of North

America.

Although we have insufficient material to use males of

the Notodelphyidae in the key, we have collected a male

of Doropygopsis longicauda (Fig. 2F) and males of two species

of Doropygus, D. laticornis (Fig. 2, G and H) and D. curvatus. A few generalities are given here in the expectation

that other males in the family will be found. Additional

information on males in this family can be found in Dudley

(1966, 1986) and in the discussion of Male Dimorphism

below. A caveat must be given that the mere physical

presence of both males and females in a single specimen

of a host is not a sufficient criterion for a common specific

identity because concurrent infestation of a single ascidian

by more than one species of notodelphyid can occur.

The males of species of such genera as Doropygopsis

and Notodelphys are found in the branchial baskets of their

hosts but have been seen by us to leave their hosts and

can swim actively. They have a generalized cyclopoid

habitus, very similar to that of males of cyclopinids or

archinotodelphyids. They are about 1/2 to 2/3 the lengths

of their corresponding females but differ from them by

having 6 urosomal segments and by having sixth legs

(Fig. 2F) on the ventral surface of the second urosomal

segment. Two spermatophoral sacs (seminal vesicles) are

visible within the second urosomal segment and the sixth

legs overlie the two spermiducal openings from these sacs.

Otherwise, except for their geniculate grasping antennules (Fig. 2F), these males conform closely in their

appendicular structure to the females, and can, in fact, be

identified by using the characters of the appendages of the

females.

In species of Doropygus, there are anamorphic, creeping

males, which, like the female, have lost considerable

motility and which are only about 1/2 the length of the

females (Fig. 2G). Their appendages are very similar to

those of the females, but, unlike the males of Notodelphys,

anamorphic males of Doropygus do not have geniculate

antennules. Such males are found in the branchial baskets

of their hosts or, even more commonly, on the peribranchial wall in the atrium along with various immature

copepodid stages. Neither they, nor the late copepodid

stages, leave the host. However, in addition to the anamorphic male, a second kind of male has been found in

D. laticornis, appearing by metamorphosis from fifth

copepodid males placed in cultures. These males (Fig. 2H)

have a cyclopoid habitus and legs adapted for swimming,

and can indeed swim actively, but their mouthparts are

so reduced that it is unlikely that they can feed. The

development of male dimorphism in this and other species

of Notodelphyidae is discussed below.
__________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 7

In addition to using the habitus and the segmental

modifications of the body as described above, the features

ofthe cephalosomic appendages (Fig. 3, A through G) are

primarily invoked in identifying the females of notodelphyids to genus and species. The identification may also

depend on the structure of the first through fifth legs (Fig.

4, A through E) and the caudal rami (Fig. 4, D, F, and G).

In making these determinations, the posture, contours,

segmentation ofthe appendages and their features of armature and ornamentation furnish useful characters. Segmentation concerns the numbers of constituent articulated

elements of appendages, which we refer to as segments,

but which could also be called articles or podomeres.

Armature consists of spines (shorter, stouter elements with

elaboration, if any, consisting of a row or rows of spinules

or tooth-like projections [Fig. 4A] and setae [Figs. 3A, 4A],

longer and more flexible and slender, with elaborations,

if any, consisting of fine hair-like processes in marginal

rows). Such a fully elaborated seta is referred to as plumose

(Fig. 4A); aesthetascs, localized on the antennules, and

thought to be chemoreceptors, are flattened, blunt-ended,

transparent elements (Fig. 3A); and hooks or claws, more

massive, are suggestive of prehensile function in most cases

(Fig. 3, B and C). Characteristic placements ofspines and

setae on the various appendages serve as diagnostic characters. Ornamentation consists of specializations of any

cuticular surface, such as hairs (Fig. 4A), spinules (Fig.

4E), flanges, pits, and pores.

There is an extremely wide range of structural variation in appendages through the family Notodelphyidae.

Appendicular characteristics that differ little from those of

the generalized cyclopoid, as well as highly specialized

features, can be found in the same species. However,

notodelphyids that have very intimate relationships with

their host ascidians, such as those living in cysts in solitary

ascidians and all of those occupying compound ascidians,

have such specialized appendages that most similarities to

the generalized cyclopoid have been lost. In adult females

of many of these, one or more pairs of appendages may

even be missing and it is often difficult to determine

homologies of the persistent appendages or their components. Only developmental evidence and the retention

of the defining familial characteristics of prehensile antenna and broodpouch allow the placement of these species

in the family Notodelphyidae.

In the least modified Notodelphyidae, found in the branchial baskets of solitary ascidians, the appendages have a

number of points of similarity to the appendages of the

species of the family Archinotodelphyidae but are somewhat further removed from those of the family Cyclopinidae by reductions in segmentation and setation. Evolutionarily, the Archinotodelphyidae appears to lie between

the other two families. This relationship between the three

families has been discussed in detail by Illg (1955, 1958)

and Dudley (1966).

In comparing the appendages of the notodelphyids with

those of the archinotodelphyids, we use only the features

of species of the least modified genera Notodelphys, Doropygopsis, Notodelphyopsis, and Doropygus. As an aid to understanding the structure of appendages and caudal rami we

present illustrations for Doropygus demissus (Figs. 3, A

through G; 4, A through G) as representative of the genus

Doropygus whose species are the most commonly encountered notodelphyids in the eastern United States.

Similarities in the appendages in the most generalized

notodelphyids and archinotodelphyids lie in the presence

of the articulated, prehensile hook on the apical segment

of a uniramous antenna (Fig. 3, B and C); concurrence

in the overall construction of the biramous mandible (Fig.

3D) and the biramous maxillule (Fig. 3E), although there

are differences in the two families in numbers of setae on

protopodites and rami; the 4 pairs of metasomallegs conform to the general cyclopoid type of segmentation and

armature. In Doropygus (Fig. 4, A through C), however,

the proportions of the rami of the legs differ from those

of cyclopinids and archinotodelphyids. In legs 2-4 there

can be consolidations of the apical and middle segments

of the endopodites, and the elements of armature on the

lateral margins of exopodites are more like setae than

spines. The caudal rami of Notodelphys, Doropygopsis, and

Notodelphyopsis have long terminal setae and shorter separate

dorsal and lateral setae, as in archinotodelphyids, but, in

Doropygus (Fig. 4, D, F, and G), the terminal setae are

greatly reduced and the rami are somewhat curved and

tapered.

There are a number of differences in appendages of the

most generalized notodelphyids and those of archinotodelphyids. These are as follows: There is a trend in notodelphyids to reduction in the segmentation of the uniramous

antennule (Fig. 3A), the least-modified notodelphyid

females have 9 to 15 segments compared with 15 to 17

segments in archinotodelphyids; the antennule in male

notodelphyids is not geniculate except in species of Notodelphys, Doropygopsis, and Notodelphyopsis; the antenna (Fig.

3, B and C) has fewer setae and the number of segments

is reduced from 4 to 3, although Doropygopsis does have an

indication of a partially fused fourth apical segment; and

a maxilla (Fig. 3F) which is 5-segmented rather than

6-segmented as in archinotodelphyids because of a fusion

of the 2 basal segments, indicated by a nonfunctional

articulation in Doropygopsis. There are still 3 small apical

segments in the maxilla as in archinotodelphyids, a portion of the appendage that is occasionally referred to as an

"endopodite." In some species of notodelphyids, as in

those of some species in the genus Doropygus, the characteristic claw on the second segment becomes seta-like; the

maxilliped (Fig. 3G) in notodelphyids never has more than

3 segments and can even be 2- or I-segmented, thus

overlapping the range of 3-5 segments in archinotodelphyids. There are fewer setae on the terminal segment of
8 NOAA Technical Report NMFS 96 _

Figure 3

Appendages of cephalosome, Doropygus demissus, female; (A) antennule; (B) antenna; (C) apex of distal segment of antenna; (D) mandible; (E) maxillule; (F) maxilla; (G) maxilliped. Aes = aesthetasc; Al = first antenna (antennule); A2 = second antenna (antenna);

Basp = basipodite; Coxp = coxopodite; Edt = endite; End = endopodite; Ep = epipodite; Ex = exopodite; Cn = gnathobase,

Mxl = first maxilla (maxillule); Mx2 = second maxilla (maxilla); Mxp = maxilliped; Ph = prehensile hook; Rs = rostrum; Se = seta.
__________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 9

Urosome

L5

E

L3

Cr

Figure 4

Appendages of metasome and appendages of urosome, DOTOPygUS demissus, female; (A through C) first, third, and fourth legs; (D) urosome,

ventral view; (E) fifth leg; (F) anal segment ofurosome with caudal rami; (G) caudal ramus. As = anal segment; Basp = basipodite;

Cop = copula; Coxp = coxopodite; Cr = caudal ramus; End = endopodite; Ex = exopodite; Go = genital opening; Hr = hair;

Lt, L3, L4-, L5 = first, third, fourth, and fifth leg; Se = seta; Sp = spine; Spl = spinule.
10 NOAA Technical Report NMFS 96 _

the maxilliped in notodelphyids; the fifth legs (Fig. 4, D

and E), when present in notodelphyids, are uniramous and

2-segmented as in archinotodelphyids but never bear more

than 1 lateral seta on the basal segment and 2 setae on the

distal segment, compared with 1 seta on the basal segment

and 4 on the distal segment in archinotodelphyids.

Family Ascidicolidae

The family Ascidicolidae includes six subfamilies of ascidicolous copepods: Ascidicolinae, Botryllophilinae, Buprorinae, Enterocolinae, Enteropsinae, and Haplostominae

(Illg and Dudley 1980). A seventh subfamily, Enterognathinae, includes copepods that are associated with echinoderms and hemichordates but which show many similarities

to the subfamilies associated with ascidians. The family

Ascidicolidae, unlike the family Notodelphyidae, has no

all-encompassing features which define it unequivocally.

However, the familial concept (Illg and Dudley 1980)

derives from the recognition of interlocking evolutionary

trends, most particularly as they relate to the many degrees

of adaptation of the females to a symbiotic existence.

Drawing upon the information in Illg and Dudley

(1980), Ooishi (1980) and Ooishi and Illg (1977), we can

present comparisons of the structure ofthe ascidicolid subfamilies. No female ascidicolid has a generalized cyclopoid

habitus. Only females in the subfamily Buprorinae (Fig.

5A) have broodpouches like those in the Notodelphyidae;

all of the other female ascidicolids carry external ovisacs.

The bodies of female ascidicolids range from those with

well-articulated body segments and heavily setiferous or

spiniferous appendages, somewhat referable to the cyclopoid type except for the extensive development of the

fifth legs (subfamily Ascidicolinae), to those whose metasomes show some degree ofsegmental fusion and inflation

and whose appendages show more reductions in segmentation and armature (subfamily Botryllophilinae; Fig. 5,

B through D) to those with extensive fusions of body

segments, an overall grub-like appearance with a trend

toward elongation and enlargment ofthe body, some losses

of cephalosomic appendages and specialization of the appendages which remain (subfamilies Enterocolinae, Enterognathinae, Enteropsinae, and Haplostominae; Fig. 5,

E and F).

The fact that different clusters ofthese subfamilies share

particular characteristics of the appendages in an interwoven pattern justifies the concept of an overall cohesiveness of the family Ascidicolidae. Thus, females in the subfamilies Ascidicolinae, Buprorinae, and Botryllophilinae

have antennules of 3-7 segments which are provided with

many setae of moderate length, but, in the other subfamilies, the antennule is an unsegmented to weakly

2-segmented tapered lobe with a reduced number of small

setae. The antennule is missing in species of LequerrlJa in

the Enterocolinae. The antenna is uniramous and 2- to

3-segmented in all of the subfamilies, but the armature is

generically distinct. Only the females of Ascidicolinae have

an apical articulated prehensile hook on the antenna which

resembles that of the Notodelphyidae. Females of Haplostomides and Haplostoma species in the Haplostominae do

have apical antennal spines that are moderately hooked,

but they appear to be different from hooks in notodelphyids. The mandible in the subfamilies Ascidicolinae,

Botryllophilinae, Buprorinae, and Enterognathinae shows

the basic gnathostomous derivation of the family by

possessing a medial coxopodal gnathal plate. In the first

two subfamilies, the mandible has a uniramous 1- to

2-segmented setiferous palp but the palp is reduced to a

single setiform process in the latter two subfamilies. The

mandible is absent in the Enterocolinae and Enteropsinae

and is represented by a very small unsegmented to

4-segmented lobe, usually with a few setae, in the

Haplostominae. The maxillule is lacking in females of

species of Haplostoma, Haplostomella, and Haplosaccus in the

Haplostominae, but in Haplostomides in this subfamily and

in all other subfamilies the maxillule consists of a basal piece

that bears setae medially and an unsegmented palp with

apical setae. The numbers ofsetae vary according to genus.

The basal component of the maxillule resembles the coxopodite and endite of the basic notodelphyid maxillule but

the palp is specialized. The maxilla, lost or reduced to a

very small setiferous lobe in the Haplostominae, shows

many similarities between Ascidicola in the Ascidicolinae

and all species in the Buprorinae, Enterocolinae, and

Enterognathinae. In these, there is a large basal segment

with a medial setiferous or spinelike endite and 1 or 2 distal

segments, with the apical segment drawn out as a hook.

The maxilla of Enteropsinae is 2-segmented and the endite on the basal segment is lacking, but otherwise it has

many similarities to the maxilla in the other subfamilies

listed above. This prehensile maxilla differs considerably

from the maxillae of female Styelicola in the Ascidicolinae

and all species in the Botryllophilinae, in which the basal

segment has several endites bearing long setae and where

there are up to 3 distal segments, also armed with setae.

This more closely resembles the appendage of the

Notodelphyidae. The maxilliped is not present in

Enterocolinae, Enteropsinae, and Enterognathinae but is

found in the other subfamilies and can have one of two

very different forms: 1) in Buprorinae and Ascidicolinae

it is an unsegmented lobe with medial and apical setae,

reminiscent of the maxilliped in some of the more modified Notodelphyidae; 2) in the Haplostominae and

Botryllophilinae it is a massive prehensile appendage

of 3 segments with a prehensile hook-like process on

its distal segment and is unlike any maxilliped in the

Notodelphyidae.

The 4 pairs of legs of the second through fifth thoracic

segments show patterns of segmentation and armature

which are generically distinct. None of these legs is
__________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 11

Figure 5

Lateral views of representative females of Ascidicolidae: (A) Buprorus loveni; (B) Botryllophilus norvegicus; (C) Botryllophilus sp. from Aplidium

glabrum; (D) Schizoproctus inflatus; (E) Haplostomides amarouci; (F) Haplosaccus elongatus.

A B

L5
12 NOAA Technical Report NMFS 96 _

missing in an ascidicolid female, unlike the situation in the

most modified notodelphyids which usually lack one or

more pairs of legs. The first through fourth legs are closest

in their structure to the basic cyclopoid pattern in females

of Ascidicola in having a 2-segmented protopodite and in

being biramous, but their exopodites are only 2-segrnented

and bear spines only and the 3-segmented endopodites have

both spines and extraordinarily long setae on their distal

segments, thus differing from the pattern in free-swimming

cyclopoids. Within the family Ascidicolidae, there is a

range of specialization from the ascidicolin type to the

very modified uniramous, spiniferous legs of some

Haplostominae to those of the Enteropsinae with bulky,

uniramous, 2-segmented legs which are armed distally with

a retractile hook.

Except in the subfamily Enteropsinae, where the fifth

legs are absent in females, and in Haplosaccus females

(Haplostominae) where fifth legs are represented by a 3ingle

seta, there is a trend within the Ascidicolidae to a displacement of the fifth legs to a more dorsolateral position than

in the Notodelphyidae and a tendency for these appendages

to become expanded by incorporating part of the body wall

(Fig. 5, B through E). In most ascidicolid females, the fifth

legs participate in stabilizing the bases of the external

ovisacs or, if lamellate, overlie and shelter entire, shortened, flattened globose ovisacs. There is a considerable

variety of caudal rami within the family. Caudal rami can

be similar to those of cyclopinids or archinotodelphyids

(Ascidicolinae), or they can be clawed (Botryllophilinae),

or tiny pointed processes (Enteropsinae), or barely indicated lobes with very small setules or spinules or lacking

armature (Enterocolinae, Enterognathinae, Haplostominae), or they are not set off from the end of the ur030me

at all (Buprorinae).

With the exception of the subfamily Buprorinae, males

of at least one species in each of the subfamilies have been

described (Canu 1892; Giesbrecht 1892; Monniot 1965;

Illg and Dudley 1980; Ooishi 1980; Gotto et al. 1984;

Ooishi and Illg 1977, 1986). The males in two species of

Haplostoma are described as being of the creeping type

(Ooishi and Illg 1977), but all ofthe other males ofspecies

of Haplostominae are swimmers. Virtually all ofthe males

in the other subfamilies are also thought to be capable of

swimming, but some have been described only from preserved specimens and have not been studied in life. Iftrue,

the ability of most ascidicolid males to swim actively could

explain their rarity in the ascidians. We have not been able

to find any males in our study of the Ascidicolidae on the

Atlantic coast.

Except for a presumed male of Schizoproctus, illustrated

by Sars (1921), which shows characteristics of segmental

fusion and inflation in its body similar to the modified

females, all of the other males described to date have a

cyclopoid habitus with podoplean tagmosis, reduced fifth

and sixth legs on the venter ofthe first and second urosomal

segments, respectively, and setiferous caudal rami. Conservative features of the appendages in the swimming and

creeping males include the 4-9 well-articulated segments

in the antennules and the segmentation in legs 1-4 which

approaches the cyclopoid pattern. However, the antennules

are nongeniculate and are therefore more reminiscent of

males of modified notodelphyid species than those of males

of the Archinotodelphyidae or Cyclopinidae and there are

some modifications of the legs, particularly the first legs,

which are generically characteristic and specialized. The

other appendages of male ascidicolids show subfamilial or

generic characters. If an appendage is absent in the female

of a given species, it almost always will be lacking in the

male, too. One exception is seen in males of some species

of Haplostoma which possess a small lobed setiferous maxilla, even though it is lacking in corresponding females

(Ooishi and Illg 1977).

The males ofAscidicola rosea are reported as having head

appendages very similar to those of the females (Monniot

1965), but sexual dimorphism is characteristic in the males

from other subfamilies. The dimorphic features in males,

compared with those in females, include: the larger numbers of segments in the antennule, its more profuse setal

complement and, in the Haplostominae and Botryllophilinae, the many aesthetes; the lengthening and narrowing

of the segments ofthe antenna and the presence of a distal

antennary prehensile hook in males, but not females, of

species ofEnterocola; the loss or great reduction of the coxopodal gnathal plate of the mandible in males of species

of Botryllophilus; the attenuation of the residual mandibular

palp in Botryllophilus and species of Haplostoma and Haplostomides; the loss of the basal endite of the maxillule in the

Botryllophilinae, Enteropsinae, Enterocolinae, and Haplostomides in the Haplostominae and the lengthening of the

remnant of the palp or its setae; the great reduction of the

maxilla in the males of Enterocolinae, Enteropsinae, and

Botryllophilinae to simple lobes with one to several setae,

very different from the appendage in the females; and the

proportionate lengthening of the maxilliped and the

enlargement of its apical hook in the Botryllophilinae and

Haplostominae; the reduction of the fifth legs to, at most,

a minute protrusion whose basal segment bears 1 lateral

seta and a distal segment with 1 or 2 apical setae. In the

male of Mychophilus roseus, the male has fifth legs even

though fifth legs are absent in females (Gotto et al. 1984).

A critical comparison of males in different subfamilies

of ascidicolids can lead to new ideas of their relationship.

Such a study by Ooishi and Illg (1986) llil.s suggested a

much closer relationship between the subfamilies Botryllophilinae and Haplostominae, based on similarities in the

armature and segmentation of the antennules, mandibular

remnants, and maxillipeds in males, than could be determined by studying the modified adult females alone.

Critical analyses of structure in the males also reinforce

our ideas about the close relationship ofthe Notodelphyidae
__________________ Dudley and lUg: Copepoda, Cyclopoida of the Eastern United States 13

and the Ascidicolidae. For example, the prehensile hook

on the antennae ofthe males ofspecies ofEnteroeola is very

similar to that of Notodelphyidae, as are the fifth legs and

other conservative features ofsegmentation and armature

of the swimming legs.

The ascidicolids have become more specialized than

notodelphyids in their adaptation of appendages for prehensile attachment. Little is known of their biology because

they often live in such small or opaque ascidians that studying them in place is most difficult. We assume that they

use the prehensile appendages to attach to their hosts or,

in the case of males, to hold the females during copulation. Although most ascidicolids (except Styelieola females

and Enteroeola males) do not have a prehensile articulated

hook on their antennae, they have other appendages which

may serve the same function. In males and females of the

Haplostominae and males of Botryllophilinae, the maxilliped is the most massive prehensile organ but, in females

of Enterocolinae and Enteropsinae in which the maxillipeds

are absent, the maxilla is a large, similarly developed attachment organ. On generic levels, legs may show various

prehensile modifications and, in the Botryllophilinae, the

caudal rami are clawed. There is no clear modification of

the appendages of the subfamily Buprorinae for prehension, but only the females are known.

Biology _

Associations With Ascidian Hosts

In addition to understanding the systematics and biology

of the ascidicolous cyclopoid copepods, the researcher on

these animals faces an additional challenge in needing to

be cognizant of the systematics and biology of the hosts

as well. The extent of host specificity varies among notodelphyids and ascidicolids and, as Gotto (1979) remarked,

"few hard and fast rules can be applied." Although most

species of ascidicolous cyclopoids occupy a single species

of ascidian host (phylum Chordata, subphylum Urochordata [ = TunicataJ, class Ascidiacea), there are records of

a single copepod species associating with two or more

related species of ascidian hosts in the same genus, or

associating with ascidian species in two or more genera of

the same family, or occupying two or more unrelated

species of ascidian hosts in different families or even orders

(Illg and Dudley 1961,1965; Ooishi and Illg 1977). Some

of the rosters of unrelated hosts include older records and

may stem from misidentifications of the hosts or the

copepod associates (see Annotated Systematic List in this

manual). Others, however, are real and may either result

from the copepods' ability to sense required anatomical

or physiological features even in ascidians that are not taxonomically close or they may just be cases of expediency

if preferred hosts are not as accessible as substitute hosts

at the times of infection.

There is no proven record of any ascidicolous cyclopoid

species occupying both solitary and compound ascidians.

However, in some genera, there are examples of species

that associate with solitary ascidians even though their congeners associate only with compound ascidians. Examples

from Ooishi and Illg (1977) are Haplostoma gibberum,

Haplostoma cruea, Haplostomella australensis, and Haplostomella

haloeynthiae and from Illg and Dudley (1980; this report):

Enteroeola latieeps, Botryllophilus norvegieus, and a Botryllophilus

species, all from solitary ascidians. Other species in these

genera live in compound ascidians.

Within solitary ascidians, the most common locus for

copepods is the branchial basket ( = pharynx). This site is

characteristic for Archinotodelphyidae and most females

and males of Notodelphyidae, although creeping males in

Doropygus are often found on the outside of the branchial

wall and can move back and forth through the stigmata

from the atrium to the branchial basket. Two species of

notodelphyids occupy cysts in blood vessels ofsolitary ascidians, Seoleeodes huntsmani, in the subendostylar blood vessel,

and Seoleeimorphajoubini, in blood vessels ofthe tunic. Some

representatives of the Ascidicolidae can also be found in

the branchial basket of their solitary hosts, but other sites

where representatives of this family are found are the

esophagus, stomach, intestine, or atrium. Females of the

ascidicolids Aseidieola rosea and Enteropsis ehattoni spend most

of their time at the bottom of the branchial sac near the

esophagus or within the esophagus but make periodic trips

into the stomach to deposit their ovisacs (Gotto 1957,

1961).

In compound ascidians, female Notodelphyidae can be

found in the branchial baskets or postabdomens of zooids

or in the common cloaca of the colony but their most common locus is in the matrix, either free or in cysts formed

by the ascidian colony. Males of these notodelphyids are

not known. Ascidicolidae in compound ascidians show a

wider choice of habitats. They have been found in the branchial basket; stomach; intestine; postabdomen; peribranchial, perivisceral, or epicardial cavities; common cloaca

of zooids; or in the common matrix. Both males and

females show a considerable peregrination into the matrix

(Ooishi and Illg 1977). The paths ofmovement of ovigerous

females are often marked by dropped ovisacs in the matrix

or the zooids. Chatton and Harant (1924) described how

the ascidicolid Enteroeolides eeaudatus travelled from deep in

the matrix to the surface ofthe compound ascidian in order

to lay its eggs.

The knowledge of the nutritional biology of the ascidicolous copepods is very incomplete. We adopt the neutral

term "associate" (Gooding 1957) in referring to these

copepods because the trophic nature of the symbiotic

association is usually only inferred. For most cases, we

would probably be correct in using the term "commensalism" because the interaction is obligate for the associates

but no harm appears to devolve to the host. For copepods
14 NOAA Technical Report NMFS 96 _

living in the branchial baskets, the ascidian appears to

supply a nutritional bonanza in the form of plankton or

detritus carried in sea water through the branchial siphon

and filtered by sheets of mucus on the walls of the branchial basket. The sea water, largely depleted of food, then

passes through the modified gill slits (= stigmata) into the

atrium and is expelled by the atrial siphon. The sheets of

mucus, produced by glands of the endostyle, are moved

by cilia along the branchial walls to the dorsal lamina where

a more concentrated food string is fashioned and is

ultimately passed to the esophagus and stomach. Because

of the opacity of most ascidian hosts, it has been difficult

to study the behavior of ascidicolous copepods in situ, but

Gotto (1957, 1979) succeeded in observing the feeding process of female Ascidicola rosea living in transparent Corella

parallelogramma, and female Pachypygus gibber living in transparent Ciona intestinalis. He found that both females attach

to the food string as it passes from the dorsal lamina into

the esophagus and feed on particles from the concentrated

string. He referred to this mode of feeding as "larder

feeding" and suggested that species of Notodelphys possibly

feed in a somewhat similar manner, although their agility

might permit them to nibble on the food captured by the

moving mucous sheets on the branchial walls rather than

having to attach to the food string.

Species of ascidicolid copepods that occupy the intestine,

stomach, or atrium of their hosts would be bathed in partially digested food or feces and might fit Gotto's (1979)

category of "debris feeders." The notodelphyid Scolecodes

huntsmani is known to contain only ascidian blood cells in

its gut (Dudley 1968) and, for this grade of interaction,

"parasitism" would be more appropriate description than

"commensalism." Nothing is known about the actual food

or feeding methods of the copepods that live in the tunic

of solitary ascidians or in the postabdomen or matrix of

compound ascidians.

Ascidicolous cyclopoids are not the only copepods known

to be associated with ascidians. In the literature on Copepoda of the world, there are many cases of association of

copepods of the order Poecilostomatoida, family Lichomolgidae, with a long roster of ascidian hosts (Hume., and

Stock 1973) and a single species, Henicoxiphium redactum Illg

and Humes (1971), has been described from Florida and

North Carolina. In our experience, members of the order

Harpacticoida are also frequently encountered in association with ascidians, but it is not certain whether there are

true symbiotic relationships. Recorded as associates of

species of compound ascidians of the genus Amaroucium

(now Aplidium) in the vicinity of Woods Hole, Massachusetts (SeiweIl1928; Wilson 1932; Coull 1977) are the harpacticoids, Tisbe gracilis (Scott) (Synonym: Tisbe wilsoni

Seiwell, 1928) and Paramphiascella commensalis (Seiwell),

1928. Because of the many microhabitats within an ascidian, one can also find a rich variety of other organisms

living in or on these animals. These other associates include algae, protozoans, hydroid coelenterates, flatworms,

nemertean worms, nematode worms, polychaete annelids,

burrowing and nestling bivalve molluscs, and crustacean

arthropods such as amphipods and decapods. Little is

known about the ecology ofthis microcosm, but predatorprey and competitive interactions and the partitioning of

food resources must be very complex.

Life Cycles

Although nothing is known of the development of the

Archinotodelphyidae, we have considerable information on

the life cycles of some copepods of the family Notodelphyidae (Canu 1892; Dudley 1966; Hipeau-Jacquotte

1978a) and some of the subfamilies of the Ascidicolidae

(Canu 1892; Gotto 1957; Dudley 1966; Anderson and

Rossiter 1969; Illg and Dudley 1980; Ooishi 1980). These

investigators have used cultures in sea water to determine

that the life cycle incorporates early free-swimming lecithotrophic naupliar and copepodid stages in both the notodelphyids and the ascidicolids. Only the last few copepodid

stages are symbiotic and these have been discovered by

careful examinations of ascidians. It is important to emphasize that, because the free stages are lecithotrophic and

do not feed during this free existence, they really have an

ultimate nutritional dependency on the host.

The nauplii have their antennules, antennae, and mandibles developed as swimming appendages with no feeding

functions. Characteristically, the development of the postmandibular appendages is cryptic in nauplii, and only the

distal margins of these internally developing appendages

are evident on the naupliar exuviae. The yolk in the developing gut ofthe nauplius, upon which it subsists, is usually

brightly colored in shades of red, yellow, green, blue or

purple, or mixtures of these, although in some species the

yolk is white. Contrasting color patches or spots of orange,

yellow, red, brown, or green pigment are often bilaterally

arranged in the naupliar body. A single naupliar eye, not

always present, is medially placed near the anterior end

and ranges from a brilliant red to pink to white. Tapetal

crystals impart a shiny appearance to the eye. After a

number of naupliar stadia ranging from as few as one, as

in Haplostomella australiensis (Anderson and Rossiter 1969),

to three in some ascidicolids of the Haplostominae (Ooishi

1980), to four in Ascidicola rosea (Gotto 1957), to five in most

notodelphyids (Dudley 1966) and other Haplostominae

(Ooishi 1980), to six in the notodelphyid Pachypygus gibber

(Hipeau-Jacquotte 1978a), a molt to the first copepodid

stage occurs. The number of naupliar stadia and the total

length of the naupliar period is dependent not only on the

species but is often temperature dependent as well in

Notodelphyidae (Dudley 1966; Dudley, unpub!. observ.).

In addition, if the embryos have a particularly long incubation time in the female's broodpouch, there may be a

smaller number of free-swimming stages after they hatch.
__________________ Dudley and Illg: Copepoda, Cydopoida of the Eastern United States 15

The first copepodids, indeterminable to gender, have a

cyclopoid body form with all of the appendages of the

cephalosome characteristic of the species present in at least

rudimentary form, a metasome of two or three segments

bearing swimming legs that are not yet of the definitive

form, and a urosome whose most anterior segment bears

the rudiments oflegs. These copepodids swim actively. At

the molt to the second copepodid stage, the exopodite of

the antenna is lost in all of the species of ascidicolous

copepods studied and other reductions can occur if these

are characteristic of the adult of the species. The second

copepodid is the infective stage in all notodelphyids studied

(Dudley 1966, 1969; Hipeau-Jacquotte 1978a, 1984),

becoming photonegative and sinking in the water to enter

a benthic ascidian. A similar situation obtains in the

ascidicolid haplostomins whose development is known

(Ooishi 1980). In notodelphyid second copepodids, proportionately large aesthetes on the antennules and a

cephalic sensory organ that reaches its maximal development at this stage and later declines (Dudley 1972), may

help in sensing an appropriate host. In the symbiotic stages,

with each subsequent molt, legs and cephalosomic appendages progress in their development toward the definitive

form ofthe adult: there are additions of new legs posteriorly

up to the characteristic number and a refinement of the

segmentation and setation of other appendages. Thus,

among copepods extracted from ascidians there is a fair

possibility of encountering developmental stages, from the

second through fifth copepodids, as well as the adults. In

the least modified species, sexual dimorphism is first apparent in the fifth copepodid stages, but, in those species

in which sexual dimorphism is profound, the divergence

in form between males and females may be more gradual,

appearing first in the fourth copepodids.

Male Dimorphism

Two types ofmales have been reported in the notodelphyids

Doropygus seclusus from the U.S. Pacific coast (Dudley 1966),

Doropygus laticornis from the U.S. Atlantic coast (Dudley

1986), and the European Pachypygus gibber (HipeauJacquotte 1978 band c; 1980 a and b; 1984; 1988). These

two types of males are referred to as "anamorphic" and

"metamorphic" by Dudley and as "typical" and "atypical", respectively, by Hipeau-Jacquotte.

In these notodelphyids, some male fifth copepodids molt

to anamorphic (typical) adult males (Fig. 2G), which like

the adult females are adapted for life within ascidian hosts.

Their cephalosomic appendages are like those of the females; the body segments show fusions; the setae on the

appendages are short; and the legs are not adapted for

swimming. In Doropygus spp., these males move by creeping

on the branchial and/or peribranchial wall ofthe ascidian.

Other male fifth copepodids undergo a profound metamorphosis to become actively free-swimming metamorphic

(atypical) adult males with such extreme reductions in the

gnathal portions of their mouthparts that it is unlikely that

they can feed. However, the augmentation of the setation

of their antennules and swimming legs and the streamlining and more functional articulations of the segments of

their bodies than in the anamorphic males, makes the

metamorphic males very proficient swimmers.

Before the occurrence of such metamorphic males was

appreciated, some males collected in the absence of female

ascidicoles were diagnosed as notodelphyids, and a genus

Agnathaner was erected for the reception ofsuch males (Canu

1891). This genus has received a number of species, none

pertinent to the fauna of our considerations, but in the main

they have been reallocated as the corresponding females

have been determined.

With the exception of two males found inside the branchial basket of ascidians, Dudley (1966, 1986) has only obtained the metamorphic males in species of Doropygus when

their male fifth copepodids were removed from the ascidian and studied in cultures. The anamorphic males are

common inhabitants of the host ascidians. Based on results

of infection experiments, Hipeau-Jacquotte (1984, 1988)

proposed that environmental factors exert an epigenetic

effect on the development of the two kinds of males in

Pachypygus gibber. She has shown (1988) that the development of atypical swimming males in P. gibber occurs only

within very young representatives of the host Ciona intestinalis which she infected with third copepodids of Pachypygus. Conversely, the development oftypical (anamorphic)

males occurs only in older hosts. The causative environmental factors for the development of the typical males

appear to be the presence of a female and a sufficient

amount of food in the older ascidian hosts, while the

reduced amount of food in very young Ciona intestinalis

evokes the development of atypical swimming males. After

the terminal molt of the atypical fifth copepodid within the

young ascidian, the atypical swimming male of Pachypygus

gibber leaves the small ascidian, within which a female cannot develop, and enters into the second pelagic phase in

its life cycle. In order to search out a host containing a

mature female, the swimming male would undoubtedly require chemical signals and would need effective sensory

organs to receive these. Hipeau-Jacquotte (1984, 1986,

1987) has described sensory/glandular complexes, hundreds of which are present on the ventrolateral, pleural

surfaces of the cephalosome of atypical swimming males,

but absent in the typical males and females. Each functional unit, possibly chemosensory, consists of 2 bipolar

sensory cells whose dendrites have basal 9 + 0 cilia, 1

envelope cell, and 1 apical cell which forms a canal through

the cuticle. The long dendrites from the sensory cells pass

through the cuticular canals and are exposed externally on

the margins of the cephalosome. These long end organs

lack a cuticular covering and become paraciliary distally,

containing hundreds of microtubules. Secretions from
16 NOAA Technical Report NMFS 96 _

adjacent exocrine dermal glands are released through pores

and spread over the cephalosomic cuticle and the sensory

dendrites (Hipeau-Jacquotte 1987). These secretions, according to this author, may serve to amplify transduction

by absorbing chemostimulatory compounds or they may

serve a protective function for the long, naked dendritic

processes. Gotto et al. (1984), using SEM, have described

structures which project from the lateral surfaces of the

cephalosome in the swimming males of the enteropsin

Mychophilus TOseus that appear to correspond to the external dendrites in P. gibber. Nishida (1986), using both TEM

and SEM, has also described similar presumable sensory

organs on cephalosomes of males of free-living planktonic

species of Oithona, thus showing that they are not exclusive

to swimming males of ascidicolous cyclopoids.

Hipeau-Jacquotte (1989) has also shown that spermatogenesis, when nonflagellate, but motile spermatozoa are

formed and encased in spermatophores are formed, is identical in the atypical and typical males of Pachypygus gibber.

Both males can mate with mature females, using their

prehensile antennae to hold onto the females' appendages

in slightly different ways, and can attach spermatophores

to the females' gonopore. The atypical male, at least under

the culture conditions used, dies after extruding only one

spermatophore from each of its two gonopores because the

process destroys its genital segment. The typical males,

however, can discharge at least two sets of two spermatophores and maybe more.

All of the other species of notodelphyids and ascidicolids

in which males are known have only one kind of male,

either a creeping male or, more commonly, a swimming

male (Dudley 1966, 1986; Ooishi and Illg 1977, 1986).

Creeping males in some species of notodelphyids, found

only in the ascidians, have appendages that are similiar

to those of the females. Swimming males in notodelphyids

may be one of two types: 1) males of the least modified

species (for example, those in Notodelphys spp. that are about

1/2 to 2/3 the lengths of the females, are very active, can

swim, can commonly be found in the ascidians, and have

appendages that are very similar to those of the females

and probably feed in the same way) and 2) males of greatly modified notodelphyid species such as those of Gunenotophorus and Scolecodes spp. that are very much smaller than

the females and have appendages that are very different.

These males show losses of gnathal portions of the mouthparts and attenuation of the residual components and

probably are unable to feed in the same ways as their

corresponding females. In the Ascidicolidae, there is less

difference in the appendicular structure of creeping and

swimming males in the various species ofHaplostoma. Both

males show modifications in their mouthparts which are

reminiscent of the changes shown by type-2 swimming

males ofthe notodelphyids (Ooishi and Illg 1977). Similar

modifications are seen in all other swimming males in the

Ascidicolidae, with the possible exception of the male of

Ascidicola rosea, which has mouthparts like those of the

females (Monniot 1965) and is, therefore, more like type-l

swimming males in the notodelphyids.

Materials and Methods _

Collections of Ascidian Hosts

The problems of collecting ascidicolous copepods are in the

main those of obtaining quantities of ascidians, the host

organisms. These are entirely marine, very few tolerating

even an approach to brackish conditions. Availability of

free-flowing water is a prime requisite for abundance of

ascidians, notable encrusting organisms of piers and floats.

However, our experience indicates that the most favorable

source for infested ascidians would be bottom-dwelling

beds, usually below the level of tidal fluctuation. Dredging or SCUBA operations, therefore, offer the recommended procedures for collecting the hosts. Our experience in

dealing with the various developmental stages of copepods

has demonstrated a very striking reversal of phototactic

locomotion as the copepods reach the infesting stages and

seek the hosts. This situation, of course, explains the predominance of successful infestations in bottom-dwelling

hosts.

Assisted by colleagues and the staff of the SystematicsEcology Program, Marine Biological Laboratory at Woods

Hole, we dredged from the SS Verrill, collected specimens

from Boston whalers in waters of the Cape Cod region,

collected specimens in the intertidal zone in Maine, and

were able to obtain most ofthe live ascidians used as hosts

in this study. We also had access to preserved ascidian

species from USFW expeditions to the Gulf of Maine and

from the Gray Museum of the Marine Biological Laboratory. Ascidians from this northeastern region represent

both the Atlantic boreal population north of the Cape and

in its deep outer banks and the Atlantic temperate population south ofthe Cape, as well as cosmopolitan species (Van

Name 1945; Plough 1978). We also made intertidal collections in Florida, assisted by the staff of the Smithsonian

Marine Station at Link Port, Fort Pierce, FL and were,

therefore, able to sample ascidians from the West Indian

Region (Van Name 1945; Plough 1978). Collections ofthe

National Museum of Natural History, Smithsonian Institution, and the American Museum of Natural History were

drawn upon for confirmatory specimens of previously

recorded collections and for additional specimens extracted

by us from identified ascidians in their collections. We did

not collect live specimens from localities north of Maine

or south of Fort Pierce, FL but we have expanded our

list with copepods we extracted from preserved ascidians

and with the meager published records of material from

along the Atlantic Coast of North America from Labrador

to Key Biscayne, FL, which we corroborated wherever

possible by examination of voucher specimens of the
__________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 17

Table 1

New records from eastern North America of ascidian hosts

and their associated symbiotic copepods of the families

Ascidicolidae and Notodelphyidae. Names of ascidian

species follow Van Name (1945).

copepods. Ofthe 25 species of preserved or newly collected

ascidians examined by us, representatives of 11 species

were found to possess copepod associates. An additional

roster of7 ascidian host species is based on earlier published

reports. The publications consulted were those of Blake

(1929, 1933), Gray (1938), Illg (1955, 1958), Pearse (1947),

Prefontaine (1936), Prefontaine and Brunei (1962), and

Wilson (1921, 1932).

The following tables present, alphabetically, new records

of associations (Table 1) and the confirmed and unconfirmed reports from the literature (Table 2).

Methods of Obtaining

and Studying Copepods

Relatively few copepods inhabit compound ascidians of our

region; so far finds have been made only in association with

species of Aplidium ( = Amaroucium). The few that so occur

are sometimes observable in the living host but a serious

Ascidian host

Aplidium glabrum (Verrill)

Ascidia callosa Stimpson

Ascidia curvata (Traustedt)

Ascidia obliqua Alder

Ascidia prunum Muller

Boltenia echinata (Linnaeus)

Boltenia ovifera (Linnaeus)

Molgula arenata Stimpson

Styela partita (Stimpson)

Copepod associates

Botryllophilus sp.

Haplosaccus elongatus Ooishi and Illg

Buprorus loveni Thorell

Schizoproctus injlatus Aurivillius

Notodelphys monoseta Pearse

Buprorus loveni Thorell

Doropygopsis longicauda (Aurivillius)

Buprorus loveni Thorell

Doropygus dernissus Aurivillius

Schizoproctus inJlatus Aurivillius

Doropygus laticornis Wilson

Botryllophilus norvegicus Schellenberg

Table 2

Published records from eastern North America of ascidian hosts and their associated symbiotic copepods of the families

Archinotodelphyidae, Ascidicolidae and Notodelphyidae. Names of ascidian species follow Van Name (1945).

Ascidian host

Confirmed published records

Aplidium glabrum (Verrill)

Ascidia callosa Stimpson

Ascidia interrupta Heller

Ascidia prunum Muller

Boltenia ovifera (Linnaeus)

"Cynthia carnea" Verrill

Dendrodoa carnea (Agassiz) 1932 or

Styela coriacea (Alder and Hancock)

Molgula complanata Alder and Hancock

Molgula manhattensis DeKay

Styela coriacea (Alder and Hancock)

Styela partita (Stimpson)

Styela plicata Lesueur

Unconfirmed published records

Ascidia callosa Stimpson

Ascidia obliqua Alder

Corella parallelogramma Muller

Didernnum albidum (Verrill)

Polycarpa fibrosa (Stimpson)

Styela mollis (Stimpson)

Copepod associate

Haplostomities amarouci (Blake),

Doropygopsis longicauda (Aurivillius)'

Notodelphys monoseta Pearseb

Doropygus dernissus Aurivilliusb

Doropygopsis longicauda (Aurivillius)" b

Doropygus dernissus Aurivillius" b

Botryllophilus norvegicus Schellenbert

Doropygus pulex ThoreW

Doropygus laticornis Wilson" b

Gunenotophorus curvipes IIIg'" b

Doropygus curvatus Gray"

Pararchinotodelphys gurneyi Illg'" b

Doropygus pulex Thorellb

Pachypygus macer Illg'" b

Botryllophilus sp.

Botryllophilus sp.

Notadelphys agilis Thorell

Haplostomides amarouci (Blake)

Botryllophilus norvegicus Schellenberg

Botryllophilus norvegicus Schellenberg

References

Blake 1929

Blake 1933

Pearse 1947; Illg 1958

Illg 1958

Illg 1958

Illg 1958; Prefontaine and Brunei 1962

Wilson 1921,1932

Wilson 1932

Wilson 1932

Illg 1958

Gray 1938

Illg 1955

Illg 1958

Illg 1958

Blake 1933

Wilson 1932

Wilson 1932

Blake 1929

Prefontaine 1936; Prefontaine and Brunei 1962

Prefontaine 1936; Prefontaine and Brunei 1962

'New collections confirm reported occurrences.

bVerification of identity based on examinations of voucher copepod specimens.
18 NOAA Technical Report NMFS 96 _

canvass for associated copepods would require processing

the ascidians. Teasing out zooids is necessary for capture

offorms in branchial cavities. Sometimes hosts left in stale

water will extrude the zooids, or the stagnation effects may

induce copepods of some motility to leave the host. The

most favorable procedure involves systematic slicing and

teasing of the colony to expose associates embedded in the

matrix and inhabiting the zooids or associated channels and

cavities in the matrix.

The majority of forms will be found within the bodies

of simple, or solitary, ascidians. These may be tumbled

out by wholesale slicing of quantities of ascidians. Preferably, a systematic search may be made by dissection of the

host, which procedure assures the preservation of the ascidian specimen for taxonomic identification or vOl:.cher

preservation and yields the optimum in details of biological

relationships of host and associated copepod. For dissection, it is necessary to orient the ascidian. If the siphons

are obvious, the tunic may be slit well away from them

and peeled like a rind from the enclosed body. If there is

no surface landmark for orientation, the best procedure

is cautious peeling to expose the body and establish the

location of the siphons. The exposed body should then be

oriented and the dissection commenced by cutting off the

siphons. The body may then be opened by a cut between

the siphons, this proceeding ventrally along one side ofthe

body so that the latter can be opened like a book. Alternatively, the body can be sectioned by a deep cut around

the whole periphery just to one side of the midline and proceeding from one siphon around to the other, leaving the

two halves ofthe body united by the intersiphonal portion.

In individuals with thin or transparent tests this procedure

can be accomplished by cutting directly through test and

all. The sectioned body can be spread open and pinned

down, preferably under water. Conspicuous commensals

will often be detected lying in the branchial cavity. A more

thorough search, preferably under magnification, should

proceed by exploring the plications and other recesses of

the branchial basket or pharynx. The branchial basket

should then be separated from the mantle by lifting it at

the edges and by severing the many small vascular connecting strands. Care should be taken to avoid undue

damage to these. The gut and gonads will be among the

organs exposed in the epicardial cavity. The wall of the

stomach should be slit and the lumen investigated for

copepods. Adult males, developmental stages, and, rarely, females may be found in the atrial cavity clinging to

the atrial side of the branchial wall.

Living copepods may be placed in 15 to 25 mL of sea

water in embryologically clean covered Stender dishes

which should be maintained at about normal seawater

temperature. Gravid females so maintained often release

nauplii which can be further studied to establish the sequence of developmental stages. Unfiltered seawater of

reasonable quality is adequate for such maintenance of the

organisms. Contact of the seawater with metals such as

copper, aluminum, and zinc should be avoided. The procedure might be referred to as "culturing" in the sense

that clean dishes and daily or more frequent changes

of water should be maintained. No specially prepared

medium is required because the naupliar stages lack a

patent gut and subsist on their abundant yolk. Study of

such developmental stages can proceed by maintaining

large numbers in culture dishes, which should be examined

regularly for the detection of exuviae. Alternatively, a single

nauplius may be maintained and the exuviae collected en

masse after the last stage attainable in culture has been

reached. Such collected exuviae should be mounted directly

in glycerine and maintained or studied immediately as wet

mounts in seawater on slides. Microscopic examination of

living nauplii, using drop slides or other pressure-protected

mounts, will reveal the development of the postmandibular

appendages.

Adult individuals are readily fixed and preserved by the

standard fixatives and preservatives. Very contractile specimens may be induced to die in a relaxed state by floating

a drop or two of clove oil in a small amount of water in

which they are contained. Direct fixation in 95 % ethanol

is satisfactory, as is fixation in dilute formalin, Bouin's fixative, Helly's mixture or other combinations. Transfer to

70 % ethanol, preferably containing a small amount of glycerine, is suitable for long-range storage ofthe specimens.

For taxonomic determination it is necessary to prepare

microscopic slide mounts of the copepods. For best preservation on slides, we have found green-tinted Euparal to

be the best mountant. Dissection of the copepod in this

medium with fine needles requires some practice, however,

because the solvent Euparal Essence must constantly be

added to keep the mountant fluid. Temporary slide mounts

can be readily made in glycerine or lactic acid, preferably

tinged lightly with Methyl Blue, Lignin Pink, Chlorazol

Black or other appropriate stain. Dissection is greatly

facilitated by softening the copepod in a drop or two of

tinted lactic acid. A few minutes will usually prepare the

specimen for dissection which may then be carried on under

magnification with fine-tipped needles. Such dissections

can be studied directly in the dissecting medium, or a

slightly longer lasting preparation can be prepared by

transferring the appendages dissected in lactic acid to

polyvinyllactophenol or to various media for mounting insects, such as Faure's, Reyne's and Hoyer's formulae.

Mounts in the latter media can be made somewhat longer

lasting by ringing with cements such as Murrayite or with

transparent fingernail polish. Pressure-protected and nondistorted mounts should of course be sought for purposes

of adequate study and taxonomic determination. A detailed

account of methods of anatomical preparation is presented

in a paper by Humes and Gooding (1964).
_________________ Dudley and lUg: Copepoda, Cydopoida of the Eastern United States 19

Key to the Copepoda, Cyclopoida: Archinotodelphyidae,

Notodelphyidae, and Ascidiocolidae of the Eastern United States _

1 Body of typical cylopoid form (Fig. 1, A through E) (see description p. 5) Pararchinotodelphys gurneyi

1 Body not of typical cyclopoid form; modified in some appreciable way as by formation of internal

broodpouch or fusion of body segments (Figs. 2, A through E; 5, A through F; 8; 9) 2

2(1) Mature female with internal broodpouch, enclosed dorsally and laterally within one or more thoracic

segments (Figs. 2, A through E; 5A; 8; 9) 3

2(1) Mature female lacking internal broodpouch but possessing external ovisacs dependent from genital

segment at times of reproduction (Fig. 5, C, E, and F) 13

3(2) Antenna (= second antenna) uniramous, bearing one terminal articulated prehensile hook and up to

6 short setae on apical margin of distal segment (Figs. 6, 18, 19) 4

Figure 6

Antenna, Doropygus curvatus.

Figure 7

Antenna, Buprorus loveni.

3(2) Antenna (= second antenna) uniramous, lacking articulated, prehensile hook, but with 5 stiffened

setae on apical margin of distal segment (Fig. 7) BuproTUs loveni (Fig. 5A)
20 NOAA Technical Report NMFS 96 _

4(3) Body dorsoventrally depressed; broodpouch in thoracic segments of 4th and 5th legs bulging dorsally

and laterally; urosome continuing straight from forebody (Figs. 8, 2A) 5

Figure 9

DOTOPygUS CUTvatus female, lateral view.

Figure 8

Notorklphys agilis female, dorsal view.

4(3) Body laterally compressed; broodpouch in one or more thoracic segments bulging laterally and dorsally;

body curved, urosome held at angle to forebody (Figs. 9; 2, B through E) 6

5(4) Terminal segment of fifth leg with a seta and a spine (Fig. 10). Lateral seta set at approximate middle

of caudal ramus (Fig. 12) Notodelphys agilis (Fig. 8)

Figure 11

Fifth leg, Notodelphys monoseta.

Figure 10

Fifth leg, Notodelphys agilis.
_________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 21

Figure 12

Anal segment and caudal rami, Notodelphys agilis.

Figure 13

Anal segment and caudal rami,

Notodelphys monoseta.

5 (4) Terminal segment of fifth leg with single seta (Fig. 11). Lateral seta set at approximate distal fifth of

caudal ramus (Fig. 13) Notodelphys monoseta (Fig. 2A)

6(4) Broodpouch occupying mainly thoracic segment of fourth legs (= fifth thoracic segment). Body segments

anterior to fifth thoracic segment not markedly inflated (Figs. 2, B through D; 9). Rami of all legs

straight, setiferous (Fig. 4, A through C) 7

6 (4) Broodpouch occupying segments ofsecond through fourth legs (= third through fifth thoracic segments).

Body segments anterior to fifth thoracic segment markedly inflated (Fig. 2E). Rami of legs 2-4 curved,

sprawling apart. Exopodites and endopodites of legs 2-4 contorted and lacking normal setiform and

spiniform armature GunenotophoTUS curvipes (Fig. 2E)
22 NOAA Technical Report NMFS 96 _

7 (6) Maxilliped slender, of 3 segments; basal segment longest with 8 or 9 setae; second segment with single

long seta; apical segment a truncated cone with 4 setae (Figs. 14, 15) 8

Figure 14

Maxilliped, DOTOPygopsis longicauda.

Figure 15

Maxilliped, Pachypygus macer.

7 (6) Maxilliped flattened, squat, basically of 2 segments; basal segment bearing 9 setae; apical segment very

small, bearing 2 plumose setae (Fig. 16); apical segment sometimes only obscurely articulated with basal

segment (Fig. 17) 9

Figure 16

Maxilliped, DOTOPygUS demissus.

Figure 17

Maxilliped, DOTOPygUS CUTvatus.
__________________ Dudley and Illg: Copepoda, Cydopoida of the Eastern United States 23

8(7) Distalmost seta of maxilliped long and plumose (Fig. 14); antenna with basal segment bearing one

long, plumose seta and one very short seta (Fig. 18) Doropygopsis longicauda (Fig. 2B)

Figure 18

Antenna, Doropygopsis longicaudo..

., Figure 19

Antenna, Pachypygus macer; (A) entire appendage; (B) 2 basal segments with small

seta in articulative membrane. Se = seta.

8(7) Distalmost seta of maxilliped very short (Fig. 15); antenna with basal segment lacking setae (Fig. 19A)

or with one or two very minute setae on articulative membrane between basal and second segments

(Fig. 19B) Pachypygus macer (Fig. 2D)

9(7) Maxillular endopodite with 2 setae (Figs. 20, 21) 10

Figure 20

Maxillule, Doropygus pulex. End

= endopodite; Ex = exopodite.

Figure 21

Maxillule, Doropygus schellenbergi. End = endopodite;

Ex = exopodite.
24 NOAA Technical Report NMFS 96 _

9(7) Maxillular endopodite with 3 setae (Figs. 22, 23) 11

Figure 23

Maxillule, Doropygus curvatus.

End = endopodite; Ex =

exopodite.

Figure 22

Maxillule, Doropygus laticornis. End

endopodite; Ex = exopodite.

10(9) Maxillular exopodite with 4 setae (Fig. 20) Doropygus pulex

10(9) Maxillular exopodite with 3 setae (Fig. 21) Doropygus schellenbergi

11 (9) Mandibular exopodite with 4 setae (Fig. 24) Doropygus laticornis (Fig. 2C)

Figure 24

Mandible, Doropygus laticomis.

Ex = exopodite.

Figure 25

Mandible, Doropygus curvatus.

Ex = exopodite.
_________________ Dudley and IIIg: Copepoda, Cyclopoida of the Eastern United States 25

11 (9) Mandibular exopodite with 5 setae (Fig. 25) 12

12(11) Maxilliped clearly 2-segmented; distal segment bearing 2 setae and clearly articulated with basal

segment (Fig. 16) Doropygus demissus

12(11) Maxilliped with distal segment indicated only as lobe bearing 2 setae; no clear articulation with basal

segment (Fig. 17) Doropygus curvatus (Fig. 9)

13 (2) Legs 1-4 biramous, each with 1-segmented exopodite and endopodite (Figs. 26, 28) or with 1-segmented

exopodite and obscurely 2-segmented endopodite (Fig. 27); caudal rami with apical, prehensile,

hooked claws (Figs. 29, 30) 14

Figure 26

Right first leg, Botryllophilus sp. from Aplidium glabrum. End

= endopodite.

Figure 30

Caudal ramus, Schizoproctus

inflatus.

Figure 27

Right first leg, Botryllophilus norvegicus. End = endopodite.

Figure 29

Caudal ramus, Botryllophilus

endopodite. sp. from Aplidium glabrum.

Figure 28

First leg, Schizoproctus inflatus. End
26 NOAA Technical Report NMFS 96 _

13(2) Legs 1-4 uniramous; single ramus (exopodite) of one segment, not clearly articulated with protopodite

(Figs. 31, 32); caudal rami without prehensile claws, armed only with setiform or spiniform elements

(Figs. 33, 34) 16

Figure 31

First leg, Haplostomides amarouci.

Figure 32

First leg, Haplosaccus elongatus.

Figure 33

Caudal ramus, Haplostomides amarouci.

Figure 34

Caudal ramus, Haplosaccus elongatus.

14(13) Fifth legs lamellate (Fig. 35); endopodites of legs 1-4 armed only with spines (Figs. 36, 28) .....

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizoproctus inflatus (Fig. 5D)

Figure 36

Fourth leg, Schizoproctus inflatus. End =

endopodite; Sp = spine.

Figure 35

Fifth leg, Schizoproctus inflatus.
_________________ Dudley and lUg: Copepoda, Cyclopoida of the Eastern United States 27

14(13) Fifth legs not lamellate, otherwise (Figs. 37, 38); endopodites of legs 1-4 armed with setae (Figs.

39, 26, 27) 15

Figure 37

Fifth leg, Botryllophilus norvegicus.

Figure 38

Fifth leg, Botryllophilus sp. from Aplidium

glabrum.

Figure 39

Right fourth leg, Botryllophilus sp. from

Aplidium glabrum. End = endopodite;

Se = seta.

15(14) Distal segment of antenna with 5 spiniform elements of armature (Fig. 40) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Botryllophilus norvegicus (Fig. 5B)

15(14) Distal segment of antenna with 7 setiform elements of armature (Fig. 41) .... Botryllophilus sp. (Fig. 5C)

Figure 40

Right antenna, Botryllophilus

norveglcus.

Figure 41

Right antenna, Botryllophilus sp.

from Aplidium glabrum.
28 NOAA Technical Report NMFS 96 _

16(13) Mandibles, maxillules and maxillae reduced; all setiferous (Figs. 42 through 45) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplostomides amarouci (Fig. 5E)

Figure 42

Appendages of cephalosome, Haplostomides amarouci, ventral view. Al = first antenna (antennule); A2 =

second antenna (antenna). Dp = dorsal cuticular plate; Ie = internal cuticular support; Lab = labrum;

Md = mandible; Mxl = first maxilla (maxillule); Mx2 = second maxilla (maxilla); Mxp = maxilliped; Rs = rostrum.

Figure 43

Mandible, Md, Haplostomides amarouci.

Figure 44

Maxillule, Mxl, Haplostomides amarouci.

Figure 45

Maxilla, Mx2, Haplostomides

amarOUCI.
________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 29

16(13) Mandibles, maxillules and maxillae reduced lobes without normal setae (Figs. 46, 47) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplosaccus elongatus (Fig. 5F)

Figure 46

Appendages of cephalosome, Haplosaccus elongalus, ventral view. Ai = first antenna (antennule); A2 = second antenna (antenna). Dp = dorsal cuticular plate;

Ie = internal cuticular support; Lab labrum; Mxl

= first maxilla (maxillule); Mx2 = second maxilla

(maxilla); Mxp = maxilliped; P = post-oral plate; Rs

= rostrum.

Figure 47

Enlargements of mouthparts of

Haplosaccus elongalus. Md = mandible; Mxl = maxillule; Mx2 =

maxilla.
30 NOAA Technical Report NMFS 96 _

The following list of ascidicolous copepods of the Cyclopoida (17 species) is arranged systematically in families

after Illg (1955, 1958) and Illg and Dudley (1980), according to a range of morphological adaptations ofthe copepods

to life in host ascidians. Within the family Notodelphyidae,

we list the type genus first and then present the other genera

alphabetically. In the genus Doropygus, the type species is

given first, and the other species follow alphabetically. Subfamilies of the family Ascidicolidae are also in alphabetical

order. Data on collections in the area covered here are

given as well as an indication of the world distribution of

species which are not endemic to the eastern United States.

Wherever possible, museum catalog numbers are given for

preserved collections. Additional voucher specimens of

newly collected material, as seemed appropriate to us, have

been deposited in the U.S. National Museum of Namral

History, Smithsonian Institution.

Family ARCHINOTODELPHYIDAE

Lang, 1949

Pararchinotodelphys gurneyi Illg, 1955-Found in the

branchial cavity of a specimen of Styela partita (Stimpson),

United States National Museum (USNM) No. 3181, collected about 85 miles southeast of Martha's Vineyard,

Mass., lat. 39°54'N, long. 69°51'30"W; USFC R/V Fish

Hawk Sta. 940, 4 Aug. 1881, 134 fathoms[?J, black sand;

4 females (Holotype, USNM No. 97608, Paratypes USNM

No. 92536). The species was described from preserved

specimens (Illg 1955) and has never been seen alive. The

inclusion here of this species is tentative. Normally the

seaward limit of distribution of forms in the MFFEUS

manuals is about 200 m. The record of collection of this

species from the continental slope at a depth of 134 fathoms

would seem disqualifying. The identification of the host

as Styela partita introduces a complication. The authority

for the identification is not known. The species is cited by

Plough (1978) as an inshore, shallow-water species. Ifthe

host identification is correct it is possible that P. gurneyi will

be encountered again in collections from our area. However, the tendency of archinotodelphyid species to inhabit

deep waters, as far as records to date would indicate, may

be the deciding factor to eliminate the species from the

present list.

Family NOTODELPHYIDAE Dana, 1853

Notodelphys agilis Thorell, 1859-Wilson (1932) recorded

3 females from Ascidia (now Corella) parallelograma Muller

at Woods Hole, Mass., 6 fathoms, in 1925. His distribution records also refer to an occurrence at the Bay of Fundy,

Maine (USNM 54090). We have examined the latter

specimens and determined them to be Doropygopsis longicauda (Aurivillius). Illg (1958), with reference to the North

American fauna and, particularly concerning published

reports of this species at Woods Hole, notes" some of the

reported occurrences have been checked and the specimens

upon which they were based have been found to be

representatives of other species." We have not been able

to confirm the occurrence of this species on the North

Atlantic coast of North America. Many published records

place this species in the European fauna. Thorell (1859)

recorded the color of the ova and embryos as "fuscovirentia. "

Notodelphys monoseta Pearse, 1947-This was described

from Ascidia interrupta Heller at Beaufort, North Carolina.

Illg (1958) redescribed the species from the type lot. We

have made new collections of this species, the only collections made since the original description: 1 female, 1 male,

1 female fifth copepodid from Ascidia curvata (Traustedt),

on shell, intertidal on - 0.8' tide, Little Jim Island, Indian River, Florida, 3/7/77; 3 females, 1 immature adult

female, 1 female fifth copepodid, 1 fourth copepodid from

Ascidia curvata (Traustedt), on rocks, intertidal on - 0.8'

tide, Sebastian Inlet, Indian River, Florida, 3/8/77. According to Van Name (1945), both of the ascidian hosts

are southern species. A. interrupta ranges from North

Carolina to the southern coast of Brazil. A. curvata ranges

from Florida to West Indies and Puerto Rico.

Doropygopsis longicauda (AuriviIlius) 1882 (Synonyms:

Doropygus longicauda Aurivillius, 1882; Doropygopsis

longicauda Sars, 1921)-We removed one male from a

specimen ofAscidia obliqua Alder, an ascidian supplied and

identified by H. Plough and collected USFW, Alb. IV-64,

off Portsmouth, N.H., 43°03'N, 700 24'W., 98 m, 1/17/64.

Other specimens examined include 11 females, 3 males and

1 fifth copepodid from Ascidia prunum Muller, intertidal,

Eastport, Maine, collectors M.P. Morse and N. Riser,

4/26/76. Specimens from Ascidiopsis complanata Verrill (now

A. prunum Muller), identified by C.B. Wilson as Notodelphys

agilis, from Todd's Head, Bay of Fundy, Maine, 1872,

25-30 fm, have been re-identified by us as the present

species. Illg's (1958) records include females from Ascidia

sp. from off Grand Manan Island, New Brunswick and

from the entrance to Goose Bay, Lake Melville, Labrador,

one female from A. prunum Muller from Lake Melville,

Labrador and males and females from unidentified asci~

dians from off Labrador. Blake (1933) listed both sexes of

Doropygopsis longicauda as associates ofAscidia callosa Stimpson from the Mt. Desert Island region, Maine. This species

is circumboreal, reported in Phallusia (now Ascidia) obliqua,

A. callosa and unidentified species of Ascidia from off

Sweden, Norway, Iceland, Greenland (Davis Strait) and

it is an associate ofA. callosa and A. paratropa (Huntsman)
__________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 31

in the Pacific Ocean off Washington State and British Columbia. Van Name (1945) reported that A. callosa is an

inhabitant of shallow water to moderate depths, usually

found in water shallower than 30 fathoms but occasionally extending as deep as 80 fathoms. It is circumboreal,

as is A. prunum, but the latter species lives in water from

a few fathoms below low water to 100 fathoms, while

A. obliqua is principally a species of deep water, with

American records ranging from 33 to 289 fathoms. While

we have not collected any specimens ofthis copepod in the

Cape Cod area, its collection could be expected on the

banks offCape Cod since A. callosa, A. prunum, and A. obliqua all extend there (Van Name 1945; Plough 1978).

Because all of the specimens collected for this report were

in preserved ascidians, we were unable to verify that the

colors of the copepod conform to those of the Pacific

specimens. The latter females are whitish, transparent,

have a bright red eye and rose-lavender embryos in their

broodpouches. The males are colorless and exhibit an agility comparable to those of species of Notodelphys.

DOTOPygUS pulex Thorell, 1859-Two females of this

species were obtained by Wilson (1932) in a Molgula

papillosa Verrill (now M. complanata Alder and Hancock)

dredged in 6 fathoms off Martha's Vineyard on a stony

bottom, 25 July 1924. We have confirmed the identification of these as D. pulex (USNM 56573) but we have not

been able to establish an exact attribution ofthe host since

there is no record of its preservation. As Illg (1958) points

out, the recorded distribution of this copepod species is virtually worldwide. This is the most extensive record compiled for any notodelphyid and the roster of its hosts is the

most diverse. We have discussed the broad concept applied

to this species by previous investigators, our recognition

of the degree of variation of representatives from different

hosts, and our belief that D. pulex s. lat. represents a complex of populations with morphological distinctions correlated with occurrences in particular hosts (Illg and Dudley

1961, 1965). It may be that when the variant populations,

including the population in M. complanata in the northeastern United States are studied in detail, they may prove

to be separate species rather than conspecific. We have proposed that the type host for Thorell's species be fixed as

a member of the family Ascidiidae (Illg and Dudley 1961,

1965). Because we have seen no living specimens of this

copepod in the Woods Hole region, we have no information on its biology or color. Records of occurrence of this

species from Boltenia ovifera (Linnaeus) from the St. Lawrence Estuary were published by PrHontaine and BruneI

(1962), based on collections made in 1933 by PrHontaine

and determined by C.B. Wilson. Another collection from

B. ovifera of PrHontaine, 1929, identified by Wilson, was

re-identified by Illg (1958) as Doropygus demissus and so

published by PrHontaine and Brunei (1962). Coming from

identical hosts, these two occurrences would seem to be

conspecific but the material first listed has not been found

for verification. We find this host association dubious and

thus doubt the record. We have specimens confirming the

distribution of this species in the southeastern United

States. Illg's (1958) record from ?Styela plicata (Lesueur)

from Key Biscayne, Florida, is confirmed by specimens

from the same lot we have in hand. We have not so far

been able to have the identification of the host ascidian

authoritatively established.

DOTOPygUS curvatus Gray, 1938-Described by Gray in

1938 as an associate of Styela partita (Stimpson) from the

Woods Hole, Mass. area, this species has not been cited

since its original description. Our collections in S. partita

have shown that D. curvatus is the commonest ascidicolous

copepod in the SW Cape Cod region and islands in terms

of the proportion of ascidians infected. Of 240 S. partita

inspected, 37 (15 %) had one or more adult female copepods in their branchial baskets and 56 (23 %) had either

a female copepod associate and/or males and immature

stages in the branchial baskets or atrial walls. Males were

unknown prior to these collections. D. curvatus can coexist with Botryllophilus norvegicus in the branchial basket of

_~s. partita. D. curvatus is endemic to the Woods Hole

region. Gray (1938) described the female of this species

as follows: "Main body ofthe animals is the usual whitish

yellow, the ovaries being darker and with a greenish tint.

Eye ruby red ... Eggs in the brood pouch very dark olive

green, indistinguishable from black by artificial light. " Our

examination of the embryos and nauplii under transmitted light discloses that the embryos have a mixture of

yellow-green and purple yolk and there are scattered, more

superficial spots of orange and yellow. Collections ofD. curvatus were made in S. partita from offJob's Neck, Naushon

Island, 70', 6120/69; Vineyard Haven, piles near Lagoon

Pond Bridge, Martha's Vineyard Island, Mass., 6', large

whaler, 7/1/69; 1 mile from Nobska Beach, Woods Hole,

41°32'48"N, 700 38'03''W, 60'-30', SS Verrill, 7/3/69; off

Falmouth Heights, 41°30'51"N, 700 34'55''W, 25', SS

Verrill, 7/3/69; off Falmouth Beach, trawl, 30', Summers

collector, 7/7/69; off Edgartown, Martha's Vineyard

Island, Mass., 21'-28', SS Verrill, 7/9/69; Steamship

Authority Dock, Martha's Vineyard Island, Mass.,

8/27/63; Nantucket Sound off Coatue Beach, north side

of Nantucket Island, Mass., on a line with Nantucket

standpipe and Nantucket breakwater beacon, Gray

Museum specimen SEP 371, P. Schwamb collector,

8/7/64; Vineyard Sound off Falmouth Harbor, Mass.,

Gray Museum specimen SEP 1608, 5/26/65.

DOTOPygUS demissus Aurivillius, 1885-We obtained

specimens from preserved ascidians in the Gray Museum

at the Marine Biological Laboratory, Woods Hole, Mass.,

in old Columbia University demonstration specimens or

in USFW specimens supplied by H. Plough and obtained
32 NOAA Technical Report NMFS 96 _

on the Albatross cruise of 8/68 in the Gulf of Maine. Males

are unknown and all of the specimens were females removed from the branchial baskets oftwo species of Boltenia:

records from B. ovifera (Linnaeus) are Gray Museum

specimen SEP 1422, off Nausett Beach, Cape Cod,

V. Zullo collector; Sta. 9, Albatross 68-12, E. of Orion

Shoal, NE of Nantucket Island, Mass., 38 m, 41026'N,

69°30'W, sandy gravel; Sta. 177, Albatross 68-19, S. off

Penobscot Bay, Maine, 60 m, 43°44'N, 68°56'W, till; Sta.

111, Albatross 68-17, south of Seal Island, SW of Cape

Sable, Nova Scotia, 43°14'N, 66°00'W, 40 m, shelly sand;

Sta. 94, NE edge of Brown's Bank, off south end of Nova

Scotia, 82 m, 42°46'N, 66°01'W; One female was removed from the branchial basket of a Columbia University specimen ofB. echinata (Linnaeus) labelled only "Campobello" (New Brunswick, Canada). Illg (1958) examined

female specimens from B. ovifera from the St. Lawrence

Estuary, Canada, and this record was also published by

Prefontaine and Brunel (1962); he recorded 8 D. demissus

from A. prunum from Maine, 42°25'N; 600 08'35''W, 12

fm, 8/31/1883, Albatross Sta. 2064; and saw material from

B. echinata from Plover Bay, Bering Sea, Siberia and from

off Pt. Barrow, Alaska. The type locality was off Pitlekaj,

Siberia, in Cynthia (now Boltenia) echinata. This species of

copepod is a circumboreal arctic species, with the Nantucket collection being the southernmost record in the

Atlantic Ocean.

Doropygus laticornis Wilson, 1932-This species was discovered by Wilson in July 1924 as an associate of Molgula

manhattensis DeKay at Woods Hole. Illg (1958) examined

the Wilson holotypic female (USNM 56570) and 5 paratypic females (USNM 56571) and redescribed the species.

We have collected many living animals from the Cape Cod

region, all from M. manhattensis attached to eel grass in

Sengekontacket Pond (= Long Name Pond) and piles near

Lagoon Pond Bridge, Martha's Vineyard and Bass River,

Cape Cod. We also removed specimens of this copepod

from a Gray Museum specimen of Molgula arenata Stimpson SEP 862 from 35 miles east of Sapelo Island, Georgia,

30 m.

This is a southern species, with Cape Cod being the

northernmost limit of its distribution. Specimens reported

by Illg (1958) from Styela partita from the Woods Hole

region have been reexamined and determined to be D. curvatus Gray. Records already published by Illg show that

D. laticomis extends from Massachusetts to the Gulf of Mexico coast in Florida. Our specimens from the Woods Hole

region had yellow bodies, yellow-green ova in the oviducts

and yellow-green to very dark green embryos in their

broodpouches. Wilson (1932) described the female specimens as having a yellow body and darker yellow eggs and

illustrated one in color. Wilson (1932) described a presumed male of the species, without illustrating it. His

specimens of males are unavailable. Our material has

provided the first record of male dimorphism in this species.

Illustrations of the two types of males are provided (Fig.

2, G and H) to demonstrate the striking differences between them.

Doropygus schellenbergi Illg, 1958-We corroborate the

identity from a specimen from the type lot, 11 specimens

from an unidentified ascidian dredged off Georgia,

32°03'N; 79°49.5'W, 14 fm, 13 Feb. 1940, USFW Vessel

Pelican, Sta. 181-13. No subsequent collections are known.

Gunenotophorus curvipes Illg, 1958-Identification of this

species, known from only a few specimens, was based on

material long-preserved at the American Museum of

Natural History. At the time of the original description

some ambiguities in the records of the Museum and also

in many manuscripts in the C.B. Wilson archives in the

Smithsonian Institution caused concern. Publication ofthe

new species had the effect of bringing forth additional information. On the basis of this, we have been able to

unravel more of the record in the archives and now conclude that the type locality originally cited for the species,

Bering Strait, Alaska, was wrongly attributed because of

the confusion in the old records. We are sure that the type

host, Styela coriacea (Alder and Hancock), was correctly attributed, and doubtlessly the collector was Captain R.A.

Bartlett. The exact year is a bit unsure but the probable

locality is somewhere off northeastern North America. We

cannot confirm the occurrence of the species in the Arctic

or the Pacific Oceans, although the host is a circumboreal

arctic form. The Bartlett material is supported by very confused records. From what we can make out, the material

was collected in 1924. Other museum specimens exist and

these derive from collections made in the St. Lawrence

Estuary. Some of these specimens were misidentified and

some of these mistaken occurrences, cited as other species,

were published by G. PrHontaine (1936). In these accounts

the records for Bonnierilla arcuata Brement were based on

specimens re-identified later by Illg as G. curvipes. PrHontaine specimen No. 3503, from S. coriacea, was catalogued

as USNM 69892, with locality given as "unknown" and

collection date 1930, as a type. Through the kindness of

Dr. P. BruneI in 1961, at that time Biologist at Station de

Biologie Marine, Grande-Riviere, Quebec, access was

gained to the PrHontaine records and further information

is now available. USNM 69892 is from the branchial sac

of S. coriacea (Specimen 2743 AMNH, determined by W.G.

Van Name)-dredging No. 11, St. Lawrence Estuary49°16'N; 69°08'W, 27-30 m-21 July 1932. PrHontaine

specimen 3505 is from S. coriacea (AMNH No. 2726)-

dredging No. 60-48°08'49"N; 69°37'30"W, 50 m, 21

Aug. 1933. These corrections were published by PrHontaine and Brunel (1962). We have seen another specimen,

collected by PrHontaine, 1934, deposited unidentified as

AMNH 1080. Material provided us in 1960 by D.P.
__________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 33

Abbott from S. coriacea, from St. Peter's Bank, off Newfoundland, USFC Albatross sta. 2466, 45°29'N; 55°24'W,

67 fm, 1885, first established for us definitively that we were

dealing with a North Atlantic species.

Pachypygus macer Illg, 1958-This species was first recorded, misidentified as P. gibber, by C.B. Wilson, 1927,

from Microcosmus exasperatus Heller, collected at Spanish

Water, Cura~ao. Illg (1958) restudied Wilson's specimens,

and with a holotype from Styela plicata (Lesueur), collected

at Hurricane Harbor, Key Biscayne, Florida, and additional specimens from the Gulf of Mexico, 1 from an

unidentified ascidian from Sarasota, Florida, and 2 from

M. exasperatus from Sanibel Island, Florida, proposed the

new species. Later collections have added a further record

from Cura~ao (Stock 1970).

Family ASCIDICOLIDAE Thorell, 1859

Subfamily BOTRYLLOPHILINAE

Sars Nomen Conservandum

Botryllophilus norvegicus Schellenberg, 1921 (Synonyms:

Blakeanus corniger Wilson, 1921; Blakeanus groenlandicus Hansen, 1923)-We have not been able to establish

the exact publication date of the paper of Schellenberg

(1921), in which he described B. norvegicus. Inquiries at the

Trondheim Museum produced for us the information,

through Mr. Tor Strlbmgren, that the day and month of

publication are not cited in the records of the Museum,

nor are they available in the files of the printer for the

Museum. Wilson described the same species in a new

genus as Blakeanus corniger on 10 June 1921. We are using

Schellenberg's binomial for the species. Wilson's Blakeanus

is undoubtedly a synonym of Botryllophilus, so the dating

here is irrelevant. Schellenberg provided the correct generic

attribution and accepting his binomial avoids for the present the formation of a new combination. However, evidence not now available to us may demonstrate the priority

of Wilson's specific epithet corniger, producing a new combination for the species. Published records show that this

species is found on both sides of the Atlantic in ascidians

of the family Styelidae. It was recorded as an associate of

Pelonaia corrugata Goodsir in Norway (Schellenberg 1921),

from Cynthia (now Styela) rustica (Linnaeus) at West Greenland (Hansen 1923) and from "Cynthia carnea" (which

could be either Dendrodoa carnea [Agassiz] or Styela coriacea

[Alder and Hancock] in today's taxonomy) from Long

Island Sound (Wilson 1921, 1932). While we have examined the Wilson type specimen and have verified its

identity, we are unable to corroborate the host's identity

because it was apparently not preserved. Records published

by Prefontaine and Brunel (1962) list this copepod (determined as B. groenlandicus Hansen by C.B. Wilson) from

Polycarpa fibrosa (Stimpson), determined by W.G. Van

Name, and from Styela mollis (Stimpson), determined by

W.G. Van Name, from the St. Lawrence Estuary. Unpublished records ofIllg show this species to be an associate

of S. coriacea from off Pt. Barrow, Alaska and from Dendrodoa aggregata (Rathke) at Bristol Bay, Alaska. Thus, this

species is circumboreal arctic in its distribution; the Wilson

(1921, 1932) record from Long Island Sound is the

southernmost. All of our collections in the Woods Hole

region were from S. partita, as follows: from Vineyard

Haven, Martha's Vineyard Island, Mass., 10', 6/12/69;

from offJob's Neck, Naushon Island, Mass., 70',6/20/69;

trawled off Falmouth Beach, Cape Cod, Mass., 30', Summers collector, 7/9/69; Gray Museum specimen SEP 906,

from Vineyard Sound off Falmouth, Mass. Harbor, 500

yds south of Bell Buoy 16, 25'-35', V. Zullo collector,

5/26/65. Three ofthe 4 specimens found in the Woods Hole

region shared the branchial basket of their ascidian host

with females of Doropygus curvatus. B. norvegicus is a very

colorful copepod. The body is yellowish, transparent, with

a brick red midgut and a yellowish orange hindgut or the

entire gut is brick red. White crystalline patches are symmetrically arranged under the cuticle of the cephalosome,

metasome, and urosome and in the rami ofthe mouthparts

and first through fifth legs. Ova in the oviducts are yellow,

while embryos in ovisacs are transparent and have a greyish

white yolk and more superficial, symmetrical orange pigment patches. Neither adults nor immature stages have an

eye. Males are unknown.

Botryllophilus sp.-We have removed many specimens of

females of a small (average 1.3 mm) Botryllophilus from the

branchial baskets of zooids and from the matrix ofthe compound ascidian Aplidium ( = Amaroucium) glabrum (Verrill)

in the Cape Cod area and in Maine, as follows: Crab Ledge

off Chatham, Mass., 41°37.4'N, 69°47.5'W, 100', SS

Verrill, 7/16/69; 41°39.8'N, 69°48.l'W, 111' SS Verrill,

7/16/69; 40 0 38.8'N, 69°47.8'W, 109', 8/28/69;

400 38.8'N, 69°47.8'W, 115', 8/28/69; Crowe Neck,

Cobscook Bay State Park, Maine, - 2.8' tide, 7/31/69;

Chamberlain, Maine, rocky beach off State Rt. 22, Gray

Museum specimen SEP 1890,J. Reinhart collector, 8/1/69.

This animal has a deep red gut, pinkish red eye and green

ova in its oviducts and green embryos in its ovisacs.

Detached ovisacs containing embryos are sometimes found

in the ascidian's matrix. The only other records of species

ofBotryllophilus from the Massachusetts and Maine coasts,

other than B. norvegicus, are those of Wilson (1932) and

Blake (1933). Wilson (1932) reported that his specimens

(from Fish Hawk Station 237), which he identified as

B. brevipes Sars, 1921, were obtained from the branchial

basket of a solitary ascidian Phallusia (now Ascidia) obliqua.

Blake (1933) made note of an unidentified Botryllophilus

which he also found in a solitary ascidian, A. callosa, in the

Mt. Desert Island region of Maine. There are many points

of resemblance between B. brevipes of Wilson, of which we

have seen a dissected specimen (USNM 60499), and the
34 NOAA Technical Report NMFS 96 _

description of this species by Sars (1921) and with the

description ofB. bergensis by Schellenberg (1921). However,

both of these Norwegian species were collected in compound ascidians (from Botryllus sp. and Leptoclinidesfaeroensis

Bjerkan, respectively) rather than from solitary ascidians.

The deficiencies of the published descriptions make it

impossible for us fully to differentiate them from our

considerable suite of Botryllophilus sp. (from the compound

ascidian A. glabrum) and from Wilson's specimen. Such

differentiation must await more adequate compa:ative

material from Scandinavia. Should the Wilson, Sars and

Schellenberg organisms, as well as the Botryllophilus sp. from

A. glabrum of the northeastern United States, prove to be

conspecific, it should be noted that B. brevipes Sars, 1921

is a junior homonym of the valid Mediterranean species

B. brevipes Brement, 1909.

Schizoproctus inflatus Aurivillius, l885-All of the specimens we obtained were females, males, and immature

stages from branchial baskets and atria of preserved specimens of Ascidia callosa Stimpson, supplied and identified

by H. Plough: USFW 675, Alb. IV-66, Sta. 79, east of

Pollock Rip off Cape Cod, Mass., 41°38'N, 69°28'W,

99 m, gravelly sand, 2/10/66. This is the southernmost

record for this species. There is no published information

on its color. Aurivillius (1885) described the female from

the ascidian Phallusia sp. from Spitzbergen, Norway and

recorded a single female of the species from this ascidian

from the coast of Finmark, Norway. Sars (1921) also

recorded males and females from Phallusia (now Ascidia)

obliqua from 60 fm off the east coast of Finmark, Norway.

Later published records have shown that this species ranges

widely in the Arctic Ocean and it can be considered a circumboreal arctic species. It has been reported from arctic

Boltenia ovijera (Linneaus) and Molgula retortijormis Verrill,

as well as from Ascidia dijmphniana (Traustedt) and A. obliqua. Specimens in the USNM include a suite collected off

the coast of Labrador. Those for which a host has been

identified were from B. ovijera.

Subfamily BUPRORINAE Thorell, 1859

Buprorus loveni Thorell, l859-All of the specimens we

obtained were females from the branchial baskets of preserved specimens of species of Ascidia supplied and identified by H. Plough: from A. prunum Muller: USFW 635,

Delaware 61-19, Sta. 76, east ofCape Cod Light, 42°03'N,

600 04'W, 128 m, 11/4/61; from A. callosa Stimpson:

USFW 675, Alb. IV-66, Sta. 79, east of Pollock Rip, off

Cape Cod, Mass., 41°38'N, 69°28'W, 99 m, gravelly

stone, 2/10/66; fromA. obliquaAlder: USFWAlb. IV-64-1,

Sta. 11, off Portsmouth, N.H., 98 m, silt clay, 1/17/64;

USFW Alb. IV-63-7, Sta. 16, Fippinies Ledge, Gulf of

Maine, 42°56'N, 69°26'W, 157m, glacial till, 11/15/53;

ca. 25 mi east ofUniv. of Georgia Marine Institute, 12 m,

sandy, 31 °35'N, 800 45'W, USFW, Plough collector. This

is a boreal copepod on both sides of the Atlantic, known

from Ascidiidae from Norway, Sweden and the Faeroes.

Sars (1921) described it as "of an uniform whitish colour,

with the rather large ova, contained within the incubatory

cavity, of the same colour, but more opaque." "It may ...

easily escape attention, on account of its small size and inconspicuous colour. The mobility of the animal is almost

wholly lost, the only token of life perceptible is a slight

fumbling movement ofthe antennae and legs." Males are

unknown.

Subfamily HAPLOSTOMINAE

Chatton and Harant, 1924

Haplostomides amarouci (Blake) 1929-0ur collections of

females of this copepod are from the branchial baskets of

zooids or, less frequently, from the matrix of Aplidium

glabrum Verrill from Crab Ledge off Chatham, Mass.,

41°37.4'N, 69°47.s'W, SS Verrill, 7/16/69; 40 0 38.8'N,

69°47.8'W, 109', SS Verrill; and from Crowe Neck, Cobscook Bay State Park, Maine - 2.8' tide, 7/31/69. These

are the first reports of this species since its initial description by Blake (1929) as Cryptopodus amarouci, with the holotype from Amaroucium glabrum (now Aplidium glabrum) and

paratypes from Tetradidemnum (now Didemnum) albidum

(Verrill). Blake (1929) described the females as "pinkishwhite, the body rather opaque. The eggs are borne in long

strings and are pale-violet." Our specimens are pure white,

opaque, with a light pinkish-white shiny eye. Ovisacs, containing white embryos, may be attached to females or free

in the branchial baskets of zooids or in the matrix of the

colony. Males are not known.

Haplosaccus elongatus Ooishi and Illg, 1977-Females of

this species were collected in Aplidium glabrum (Verrill) at

Crab Ledge offChatham, Mass., 41 °39.9'N, 69°48.l'W,

111', SS Verrill, 7/16/69 and from a rocky beach near

Chamberlain, Maine, offRt. 22, Gray Museum specimen

SEP 1890, Reinhart collector, 8/1/69. The females are

found in excavations in the matrix of the ascidian colony

near its base. The female often lies on her anteriorly

directed ovisacs. Detached ovisacs were also found lying

in the basal matrix, either in the excavation occupied by

a female or separately. The color ofthe animal is yellowishwhite and it has a red eye and orange gut. Ova in the

oviducts are light pinkish or lavender. Embryos in the

ovisacs are white and have a very faint lavender-pink tint.

This species was described from the coast of British Columbia in A. glabrum (Verrill) and in Washington State in

A. arenatum (Van Name) by Ooishi and Illg (1977). Males

are unknown.
__________________ Dudley and Illg: Copepoda, Cydopoida of the Eastern United States 35

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Systematic Index

Copepoda

Archinotodelphyidae 19, 30

Ascidicolidae 19, 30, 33
_________________ Dudley and Illg: Copepoda, Cyclopoida of the Eastern United States 37

Blakeanus 33

corrzger 33

groenlandicus " 33

Bonnierilla

arcuata , 32

Botryllophilinae 33

Botryllophilus 33

bergensis 34

brevipes 33, 34

norvegicus 25, 27,31, 33

sp 25, 27, 33, 34

Buprorinae 34

Buprorus

loveni 19, 34

Cryptopodus

amarouci 34

Cyc1opoida 19, 30

Doropygopsis

longicauda 22, 33, 30

Doropygus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 30

curvatus 19, 20, 22, 24, 25, 31, 32

demissus 22, 25, 31, 32

laticornis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24, 32

longicauda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

pulex 23, 24, 31

schellenbergi 23, 24, 32

Gunenotophorus

curvipes 21, 32

Haplosaccus

elongatus 26, 29, 34

Haplostomides

amarOUCl 26, 28, 34

Haplostominae 34

NotodeIphyidae 19, 30

Notodelphys

agilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20, 21, 30

monoseta 20, 21, 30

sp 31

Pachypygus

gibber 33

macer ; 22, 23, 33

Pararchinotodelphys

gurneYl 19, 30

Schizoproctus

inflatus 25, 26, 34

Ascidiacea

Amaroucium

arenatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

glabrum 33, 34

Aplidium

glabrum 25, 27, 33, 34

Ascidia

callosa 30, 31, 33, 34

curvata 30

db·mphniana 34

interrupta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 30

obliqua 30, 31, 33, 34

parallelograma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

paratropa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

prunum 30,31,32,34

sp 30

Ascidiidae 31, 34

Ascidiopsis

complanata 30

Boltenia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 32

echinata 32

ovifera 31, 32, 34

Botryllus

sp 34

Corella

parallelograma 30

Cynthia

carnea 33

echinata 32

rustica 33

Dendrodoa

aggregata 33

carnea 33

Didemnum

albidum 34

Leptoclinides

faeroensis 34

Microcosmus

exasperatus 33

Molgula

arenata 32

complanata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

manhattensis 32

papillosa .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

retortiformis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Pelonaia

corrugata 33

Phallusia

obliqua 30, 33, 34

sp 34

Polycarpa

fibrosa 33

Stylelidae 33

Styela

coriacea 32, 33

mollis 33

partita 30,31,32,33

plicata 31, 33

rustica 33

Tetradidemnum

albidum 34
38 NOAA Technical Report NMFS 96 _

Index of Anatomical Terms _

The terms in this index will be found boldface in the Introduction and the Diagnosis and Relationships of Families

sections, and the text explains their meanings.

aesthetascs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

anal segment 3

armature 7

basipodite 5

caudal rami . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

cephalosome . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3

claws 7

copula 5

coxopodite .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5

endopodite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

exopodite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

geniculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

habitus 5

hooks 7

intercoxal plate 5

metasome 3

ornamentation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

plumose 7

prosome 3

protopodite 5

rami 5

segment .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 7

segmentation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7

setae 7

sixth legs 6

spines 7

tagmata 3

urosome 3

Acknowledgments _

Preparation ofthis 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" through the College of Marine Studies,

University of Delaware.

We gratefully acknowledge support and assistance from

Melbourne Carriker, Director of the Systematics-Ecology

Program, Marine Biological Laboratory, Woods Hole,

during our tenure there; Johanna Reinhart who was

curator at the Gray Museum of the Marine Biological

Laboratory in 1969; and other colleagues, students, and

personnel whose sampled specimens proved to be hosts for

ascidicolous copepods. We are grateful to the personnel of

the research vessel SS A.E. Verrill for their special help;

the late Dr. Harold Plough who lent his great expertise

to help us in our identification of ascidian hosts, and who,

with Dr. Roland L. Wigley of the National Marine Fisheries Laboratories at Woods Hole, supplied specimens of

ascidians from the Albatross IV expedition to the Gulf of

Maine. We are also indebted to Dr. Mary Rice, Director,

and to Julianne Piraino and Douglas Putnam ofthe Smithsonian Marine Station at Link Port, Fort Pierce, Florida,

for their hospitality and help in collecting and photographing specimens of copepods from the Indian River. To the

staff of the Harbor Branch Foundation Laboratory at Link

Port, we give our appreciation. Museum specimens and

information have been made available to us from the

National Museum of Natural History; Smithsonian Institution, mainly through Dr. Roger Cressey; American

Museum of Natural History, through Mr. Harold Feinberg; and the Vitenskapmuseet, Zoologisk Avdeling, Universitet i Trondheim, Norway, through Mr. Tor Str¢>mgren. We thank also Dr. Shigeko Ooishi, Mie University,

Mie, Japan, for information and discussion of great benefit

to us in interpreting our findings. Lastly, we thank Dr.

A.O.D. Willows, Director of the Friday Harbor Laboratories, University of Washington, for use of library and

laboratory facilities.

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, Gettysburg, PA 17325.

Arthur G. Humes, Boston University Marine Program,

Marine Biological Laboratory, Woods Hole, MA 02543.

David L. Pawson, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Kenneth P. Sebens, Marine Science and Maritime Studies

Center, Northeastern University, East Point, Nahant,

MA 01908.

Ruth D. Turner, Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138.

Robert T. Wike, Department of Botany, University of

Massachusetts, Amherst, MA 01002.

In addition to establishing the format for the "Marine

Flora and Fauna of the Eastern United States," the Board

invites systematists to collaborate in the preparation of

manuals, reviews manuscripts, and adivses 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
__________________ Dudley and lUg: Copepoda, Cyclopoida of the Eastern United States 39

universal emphasis on work in the marine environment and

the crucial need for precise and complete identification of

organisms related to this work. It is essential, if at all possible, 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.

Patricia L. Dudley first studied ascidicolous copepods

as a Research Assistant to Paul Illg when she was a doctoral student in the Department of Zoology at the University of Washington in 1953. She chose to work on the

life cycles of copepods of the family Notodelphyidae for

her Ph.D. thesis (1957). For two post-doctoral years she

travelled to European marine labs and the Hawaii Marine

Laboratory, collecting symbiotic copepods ofthe Ascidicolidae and Notodelphyidae and studying their development.

In 1959, she became associated with the Department of

Biological Sciences, Barnard College, Columbia University, where she still teaches as a Professor ofBiology. During her tenure at this college and numerous summers at

the Friday Harbor Laboratories ofthe University ofWashington, she has continued to study ascidicolous copepods,

adding electron microscopy of internal structures of copepods, other crustaceans and Mesozoa as subjects ofinterest.

During the summers of 1969 and 1970 she was associated

with the Systematics-Ecology Program, Marine Biological

Laboratory at Woods Hole and collected many of the

ascidicolous copepods upon which this manual is based.

Paul L. Illg first encountered ascidicolous copepods

among the rich fauna at the Friday Harbor Laboratories

during the summer of 1939, while he was a graduate student at the University ofCalifornia, Berkeley, California.

Some of the material was presented in a thesis offered for

the M.A. degree, 1941. During tenure as an associate

curator, U.S. National Museum, he resumed studies on

the material and was able to augment the collections by

a stay at Friday Harbor during the summer of 1950.

Material from the east coast was also incorporated in his

monographic studies. The completed study, offered as a

thesis to qualify for the Ph.D. degree in 1952 at the George

Washington University, was subsequently published by the

U.S. National Museum (1958).

During his tenure at the Museum he acquired a fairly

wide acquaintance with copepods, free-living and symbiotic. In 1952, acceptance of an offer from the University of Washington returned him to the Northwest where,

with a succession ofstudents, he continued studies on taxonomy, life history and biology of copepods, retiring as

Professor Emeritus in 1982. An invitation from the Boston

University Marine Program and the Systematics-Ecology

Program of the Marine Biological Laboratory in 1969

brought him to Woods Hole where a substantial amount

of his research time was devoted to the copepods which

are the subject of this manual.
40 NOAA Technical Report NMFS 96 _

Published Manuals

Marine Flora and Fauna of the Northeastern United States

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 Ho, Ju-Shey

Copepoda: Cyclopoids Parasitic on Fishes Ho, Ju-Shey

Crustacea: Branchiura Cressey, Roger F.

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

Crustacea: Cumacea Watling, Les

Artbropoda: 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., and John H. Dearborn

Marine Flora and Fauna of the Eastern United States

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

Copepoda, Cyclopoida: Archinotodelphyidae, Notodephyidae, and

Ascidicolidae Dudley, Patricia L., and Paul L. Illg

NOAA Tech Rep.

NMFS Circular no.

374

378

384

386

387

389

394

397

398

399

403

405

406

409

413

419

423

425

438

439

440

446

NMFS no.

33

91

73

96

NTIS no.

COM 73 50670

73 50888

74 50019

74 50014

7450487

74 51194

PB 257 987

261 839

268036

268 714

273 062

274 999

280040

281 969

222 923

285 538

296460

297 676

124 520

225 053

219 387

124 735

PC A03/MF A01

PB 86 156395

PB 89 189 583