NOAA Technical Report Circular 409 Marine Flora and Fauna of the Northeastern United States. Copepoda: Cyclopoids Parasitic on Fishes

Ju-Shey Ho

February 1978

U.S. DEPARTMENT OF COMMERCE Juanita M. Kreps. Secretary National Oceanic and Atmospheric Administration Richard A. Frank. Administrator National Marine Fisheries Service

Iw

FOREWORD

This issue of the "Circulars" is part ot subseries entitled "Marine Flora and Fauna of the Northeastern United States." This subseries will CODsist of original, illustrated, madem meDuals on the identification, dassifieatlon, and general biology of the estuarine and cMsla1 marine plants and animals of the Nortb­ eastern United States. Manuals will be published at irregular intervals on as many taxa ofthe region as thcre are specialists aVBiloblc to collaborate in their preparation. The rnanuals are an outgrowth of the widely used "Keys to Marine Invertebrates of tbe Wood. Hole Region," edited by R. I. Smith, published in 1964. and produeed under the auspieces of the Systematie.­ Ecology Program, Marine Biological Laboratory. Woods Hole, Mass. Instead of revising the "Woods Hole Key.... the .tatT ot the Sy.tematic.-Ecology Program decided to expand the geogr"phic coveroge and bathy­ metric range and produce the keys in an entirely new set of expanded publication•• The "Marine Flora and Fauna of the Northeaslern United States" is being prepared in collaboration with systematic specialist. in the Uniled States and abroad. Each manual will be ba.ed primarily on recent and ongoing revisionary s:rstematic research and a fresh examination of the plants and animals. Each major :.axon. treated in a separate manual, will include an introduction, illustrated glossary, uniform originally il­ lustrated keys, annotated check list with information when available on distribution, habitat, life history, ~nd related biology, references 10 the major literature of the group, and a systematic index. These manuals are intended for u.~ by biology students. biologists. biological oceanographers, informed laymen, and others wishing to identify coastal organisms tor this region. In many instances the manuals will serve as a guide to additional intormation about the species or the group. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is pla.med to include organisms from the headwaters of estuaries seaward to approximately the 200-01 depth on the eon­ tinentalsheIr from Maine to Virginia, but may vary somewhat with each major taxon and the interests ~f col­ laborators. Whenever possible representative specimens dealt with in the manual. will be deposited in the reference collections of major museums in the region. Arter a sufficient number of manuals of related taxonomic groups have been published, the manuals will be revised. grouped, and issued as special volumes. These vcJumes will thus consist of compilations of in .. dividual manuals within phyla such as the Cnidaria. Arthropoda, and Mollusca, or of groups of phyla.

ii CONTENTS

Page

Introduction ...... ,...... 1 Key to the marine cyclopoid copepods parasitic on fishes of the northeastern United States 2 Annotated systematic list 9 Literature cited . 10 Systematic index 11 Acknowledgments 12 Coordinating Editor's comments 12

The National Marine Fisheries Service (NMFS) does not approve, rec­ ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro­ motion which would indicate or imply that N~IFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication.

iii Marine Flora and Fauna of the Northeastern United States. Copepoda: Cyclopoids Parasitic on Fishes

JU-SHEYHO'

ABSTRACT

This manual includes an introduction on the ifen~ral biology, an illustruted key, an annotated sys­ temati" list. a selected hihliogrnph}', and nn index to the 19 speeies of eye!opoid copep.ds parasitic on marine fishes or the northeastern United States.

INTRODUCTION modified cephalothorax (LemE.eidae); or by boring into and lying free inside the cephalic canal of the host's gill The "Order Cyclopidea" as defined by Yamaguti operculum and head (Philicilthyidae). (1963) consists of four families of copepods parasitic on Our knowledge of the biology of this group of parasites both freshwater and marine fishes. They are Bomo­ is still scanty. A complete life cycle is knovm of only some Iichidae, Ergasilidae, Grandiunguidae, aud Tucciclae. freshwater species. Some sporadic reports on the larval However. according to Sproston et al. (1950), Ho (-Hl67b, stages are available for a few marine species, However, 1970, 1971a), Izawa (1973), and Cressey (1975), the Order based on the existing information. it seems that the should include six more families: Lernaeidae, Tel­ parasitic cyclopoids do not require an intermediate host. sidae, Chondracanthidae, Pharodidae, Philichthyidae, The nauplius hatched f!om the egg develops into the and Shiinoidae; and the family Bomolichidae should be copepodid larva, which is the infective stage and separated into the BOlD.clochidae and Taeniacanthidae. searches for the new host. Usually, there are five The family Grandiuneuidae was created by Yamaguti copepodid stages and mating takes place right after the (1963) to accommodate a single species of copepod. female finishes its last molt in the larval stage. The male Grandiungus promicrops Pearse 1952. I have checked the dies after mating, but occasionally it is found holding to type-specimen of this species deposited in the National the genital segment of the ovigerous female (Bomolochi­ Museum of Natural History in Washington, D.C., and dae, Shiinoidae, Taeniacanthidae, Telsidae) or lying in have discovered that it is merely a chalimus larva of a the host's cephalic canal with the maturl: female lernaeocerid copepod detached from its frontal filament. (Philichthyidae). However, in the family Chondra­ Therefor.', the family Grandiunguidae together with its canthidae, the male is characteristically a symbiont of monotypic genus and species, Grandiungus promicrops, the female; it is a dwarf (Fig. 1) and holds on to one of the should be discarded. Consequently, the "Order Cy­ two vermiform processes on the female trunk near the c1opidea" as it now stands consists of ten families, name­ genital somite. Therefore, adult cyc1opoids found at· ly, Bomolochidae, Chondrllcanthidae, Ergasilidae. Ler­ tached to fishes are all female. The mature females of the naeidae, Phar()didae, Philichthyidae, Shiinoidae, families Chondrscanthidae, Lemaeidae, Pharodidae, Taeniacanthidae, Telsidae, and Tuccidae. The lemaeids Philichthyidae, Shiinoidae, and Tuccidae are highly are exclusively parasitic on freshwater fishes and the transformed and do not appear as a typical cyclopoici ergasilids are predominently parasites of freshwater (Fig. 2), In these modified copepods an extensive but fishes, except El few species that are found on fishes of gradual modification of the body occurs in the process of brackish or coastal waters. The members of the other maturation after the last molt in the copepodid stage, eight families are strictly marine parasites. While all the cephalic and oral appendages are retain­ Cyclopoid copepods on fishes are found on the body ed with certain degree of modification in the adult surface or in the oral cavity, gill cavity, nasal cavity, and female, the thoracic appendages of the parasitic cy­ cephalic caM!. They attach to the host by hooking their c1opoids exhibit a wide spectrum of variation. While the modified, prehensile second antennae (Ergasilidae, maxillipeds are lacking in some families (Ergasilidae, Chondracanthidae, Pharoclidae, Tuccidae), m'lxillae Philichthyidae. Shiinoidae), the remaining thoracic (Telsidae),or maxillipeds (Bomolochidae, Shiinoidae, appendages vary from the typical unmodified cyclopoid Taeniacanthidael; by burrowing their anchorl;ke thoracopods (Ergasilidae, Lernaeidae) through the par­ tially (Bomolochidae, Taeniacanthidae, Telsidae, The­ cidae) or completely (Chondracanthidae, Pharodidae, 'Deportment of Biology. Conf.rnla State Univen;ity. Long Beach. CA Philichthyidae) modified legs to the absence of the legs 903-l0. entirely (Chondracanthidae, Philichthyidae).

-. ANTENNULE --

HEAD

LEG

LEG

Figure L-Acanthochondria cornuta; dorsal view, showing structures and terms used in key. TRUNK

MALE

POSTERIOR } GENITO­ PROCESS ABDOMEN

There are about 380 species of cyclopoids parasitic on must be very carefcl not to damage the attachment ap­ marine fishes. However, only 19 species belonging to six paratus of the parasite, for, in many CaSf1S, this modified. families are so far known to occur within the range appendage carries important taxonomic information. covered by this manual (from Maine to Virginia). Since Removal of the parasite from the host is preferably done only a small fraction of fishes occurring off the coast of under a dissecting microscope, but, if the removal must northeastern United States have ever been examined for be done in the field, a generous amount of the host tissue the copepod parasites, certainly many more species of must be taken together with the parasite. The parasite is parasitic cyclopoids are yet to be discovered. subsequently removed from the collected host's tissue It is well known that a freshwater fish heavily infested under a dissecting microscope in the laboratory. The with cyclopoids, particularly with those of the genera collecled specimens are preserved and stored in 70% al· Ergasilus and Lemaea, often dies. Although no such cohol. The study of their external anatomy always re­ reports have ever been made on marine fishes, a fish kill quires dissection of the appendages. This can be done due to heavy infestation of marine cyclopoid copepods first by soaking the specimen in lactic acid for several cannot be ruled out, because loss of weight and retarda­ hours and then removing the appendages under a dis­ tion ofthe rale ofgrowth of the fish under the influence of secting microscope with a pair of sharp needles. The parasitism is known. appendages removed are studied under a compound In collecting specimens of parasitic cyclopoids, one microscope.

KEY TO THE MARINE CYCLOPOID COPEPODS PARASITIC ON FISHES OF THE NORTHEASTERN UNITED STATES

The following key is constructed for the female cyclo­ from Maine to Virginia. A separate key to the male is not poids that have been reported parasitic on marine fishes given because some ofthem are unknown and the others, 2

.f 1 ANTENNULE---

PROSOME

LEG

LEG 2 .-.----b\.•.

LEG 3 ---/m'~~~Offl LEG 4---'''-

LE G 5 ------

GENITAL UROSOME SEGMENT--..../

CAUDAL RAMUS --+ J

Figure 2.-Holobomolochus albidus; dorsal "iew, showing structures and terms used in key.

when they occur, are always in close association with Bomolochidae, two species of Ergasilidae, and one their female partners. species ofTaeniacanthidae is not provided because these Although 19 species of cyclopoids are known from the families are relatively large and very poorly known from northeastern United States, the following key is prepared the region. If a key to the few known species is provided, for identification of the 11 species of Chondracanthidae, ii might lead a user to misidentify an unreported Rpecies one species of Philichthyidae, and one species of Tuc­ as one listed in the key. cidae. Identification of the remaining three species of

1 Body typical copepod form (Fig. 2) .. , 2

1 Body modified, not copepod form (Fig. 1) 4 3

.,~ ...__.:.l.:;j," 2 (1) Antenna with single terminal claw (Fig. 3) .Ergasiliclae

2 (I) Antenna bearing several terminal setae and claws (Fig. 4) ...... 3

Fil(Ure 3.-ErgacillUJ labracia; antenna.

Fhrure 4.-Tucca impreSBUS; antenna.

~~~ '~ ..;,:.:,;~ ....", .... 1... _

3 (2) Maxillary hook present (Fig. 5) .Taeniacanthidae

3 (2) Maxillary hook absent ...... Bomolochidae

Figure 5.-Anchialrolos occidenlaliB; cephalothorax in ventral view with antennules and antennae removed. 4 4 (1) Without attachment apparatus. living loosely in cephalic canal of fish (fig. 6) Philichthys xiphiae

l'igure 6.-PhilichthYB "iphiae; remnle.

4 (1) With attachment apparatus of modified antennae or maxillipeds. firmly attached to host tissue 5

5 (7) Antenna bearing several termin:l.l setae and claws (Fig. 4) Tucca impressus

5 (7) Antenna transformed into a tripartite process (Fig. 7) or a recurved hook (Fig. 8) 6

Figure 7.-Blias prionotu.s; antenna.

Figure 8.-Pseudoehondraeanthus dieeraUB; head, ventral view. 5 6 (5) Trunk region without any form ofoutgrowths; antenna tripartite (Fig. 7) Blias priorwti

6 (5) Trunk region with outgrowths in form of processes, protrusions, or knobs; antennae uncinate (Fig. 8) .... 7

7 (6) Trunk region with only one pair of processes at the posterior end ofthe body (Fig. 1) . 8

7 (6) Trunk region with more than one pair of processes ...... 13

8 (7) One pair of lobate legs present: head bearing a pairoflateral processes (Fig. 8) Pseudochondracanthus diceraus 8 (7) Two pairs of lobate legs present; head without lateral processes ...... 9

9 (8) Legs unilobate (Fig. 9); parasite of elasmobranchs ...... Acanthochondrites annulatus

Figure 9.-Acan/hochondrit•• annum/us: leg.

9 (8) Legs bilobate (Fig. 10); parasite ofteleosts ...... 10

Figure IO.-Acan/hochorulria phycidiB: leg.

6 10 (9) Rami of legs short and broad (Fig. 10) ...... •4canthochondria phycidis

10 (9) Rami of legs long and conical (Fig. 11) .11

Figure ll.-Acanthochondria gO/Brita; leg.

11 (10) Antennule small (relative to head) and filiform (Fig. 12) Acanthochondria exilipes

Fisure 12.-Acanthochondria "",;UpeB; head.

11 (10) Antennule large (relative to heRd) and l0bate (Fig. 1) 12

12 (11) Rami oflegs pointed (Fig. 11) ...... A canthochondria galerita

12 (11) Rami oflegs not pointed (Fig. 13) . Acanthochondria cornuta

Figure 13.-Acanthochondria comuta; leg.

7 13 (7) Legs partially modified, with sadike protopod and rodlike rami (Fig. 14) ..... Chondracanthodea deflexus

-PROTOPOD

ENDOPOD-

Figure 14,-ChondracanthodeB deflexUB; leg,

13 (7) Legs completely modified, either bilobate (Fig. 13) or trilobate (Fig. 15) 14

Figure 15.-Chondracanthus nodOBUB; leg,

14 (13) Legs trilobate (Fig. 15) Chondracanthus nodosus

14 (1.:1) Legs bilobate (Fig, 13) . ., .... , ... , ..15 8 15 (14) Head with a pair of cephalic horns; antennule smaII (relative to head) (Fig. 16) .. Chondracanthus merluccii

Figure 16.-Chondr.canthw merluccif; head.

-CEPHALIC HORN

I':; \ 14) Head without cephalic horns; antennule large (relative to head) (Fig. 17) ..... Chondracanthus cottunculi

Figure 17.-Chondracanthus cottunculi; head.

ANNOTATED SYSTEMATIC LIST Florida; striped mullet, Mugil cephalus, off Texas (Pillai 1965). This list is arranged alphabetically in families, genera Nothobomolochus saetiger (Wilson 1911). In gill cavity under their family, and species under their genus. Notes of tropical two wing flyingfish, Exocoetus volitans, off on host and distribution are given. When more than one Woods Hole, Mass.; Cypselurus callopterus from Gala­ host is known, their names are arranged alphabetically in pagos Islands (Wilson 1911). genera and species under their genus. If the common name of the host is known, it is given immediately prec­ Family CHONDRACANTHIDAE eding its scientific name. Reference to important works are cited at the end of each species. Acanthochondria cornuta (Muller 1776). In gill and oral cavities of largescaled tonguesole, Cynoglossus Family BOMOLOCHIDAE macrolepidotus, off Ceylon; witch flounder, Glyp­ locephalus cynogl08SUS, in Massachusetts Bay and Holobomolochus albidus (Wilson 1932). In gill cavity of eastern North Atlantic; American plaice, Hippoglos­ angler, Lophius piscatorius, off Woods Hole, Mass. soides platesoides, in North Sea, off Iceland and (Vervoort 1969). Greenland, and from Maine to Massachusetts; Atlan­ Nothobomolochus teres (Wilson 1911). In gill cav:ty of tic halibut, Hippoglossus hippoglossus, off Midleton Gulf menhaden, Brevoortia patronus, off Texas; At­ Island, Alaska; whiff, Lepidorhumbus whiffiagonis, in lantic menhaden, B. tyrannus, from Massachusetts to eastern North Atlantic; dab, Limanda limanda, in 9

._-~~: Irish Sea and off Iceland; flounder, Platichthys flesus, Ergasilus manicatus Wilson 1911. On gill gilaments of in North Sea; European plaice, Pleuronectes platessa, sheepshead minnow, Cyprinodon variegatus, from in eastern Nonh Atlantic; Alaska, plaice, P. quad­ Beaufort, N.C.; mummichog, Fundulus heteroclitus, rituberculatus off Pallas, Alaska; turbot, PsettG max­ from Cape Ann, Mass.; striped killifish, F. majolis, ima, off Sweden; winter flounder, Pseudopleuro­ from Beaufort, N.C.; blackspotted stickleback, Gas­ nectes americanus, from Maine to Massachusetts (Ho terosteus wheatlandi, from Cape Ann, Mass.; rain­ 1970). water killifish, Lucania parva, from Beaufort, N.C.; Acanthochondria exilipes Wilson 1932. On gills of tile­ Atlantic silverside, Menidia menidia, from Woods fish, Lopholatilus chameleonticeps, off Martha's Vine­ Hole, Cape Ann, and Barnstable, Mass.; rainbow yard, Nantucket, and Woods Hole, Mass. (Ho 1970). smelt, Osmerus mordax, from St. Andrews, New Acanthochondria galerita (Rathbun 1886). In oral Brunswick (Roberts 1970). cavity of Gulf flounder, Paralichthys albigutta. off Florida; summer flounder, P. dentatus, off Woods Family PHILICHTHYIDAE Hole, Mass. (Ho 1971b). Acanthochondria phycidis (Rathbun 1886). On gills of Philichthys xiphiae Steenstrup 1861. In cephalic canals Chilean hake, Merluccius gayi, off Chile; spotted hake, of swordfish, Xiphias gladius, in Mediterranea... , nff Urophycis reguis, off Georgia and Florida; white hake, European and American coasts off North Atlantic, U. tenuis, off Martha's Vineyard, Mass.; unknown New Zealand, and Japan (Delamare-Duboutteville host from Falkland Islands (Ho 1971b). 1962). Acanthochondrites annulatus (Olsson 1868). On gills of flapper skate, Raja batis, off Norway; spiny ra~p skate, FamilyTAEN!ACANTHIDAE R. kenojei, in Sea of Japan; barndoor skate, R. laevis off South Harpswell, Maine (Ho 1970). Anchistrotos occidentalis Wilson 1924. On gills of BliCUi prionoti Krglyer 1863. On gills of black drum, Po­ striped burrfish, Chilomycterus schoepfi, from Woods gonias cromis, off Mississippi; northern searobin, Hole, Mass. (Humes and Rosenfield 1960). Prionotus carolinus, off North Carolina; soldier sea­ robin, P. miles pectoralis, off Texas; leopard searobin, Family TUCCIDAE P. scitulus, off Florida; striped searobin, P. evolans, off Woods Hole, Mass.; bighead searobin, P. tribulus, off Tucca impressus Kr6yer 1837. On fins of bridled burr­ Florida (Ho 1970). fish, Chilomycterus antennatus, from Jamaica; striped Chondracanthodes deilexus Wilson 1932. On gills of burfish, C. schoepfi, from Massachusetts to Louisiana; marlin-spike, Nezumia bairdi, off Woods Hole, Mass.; porcupinefish. Diodon hystrix, from Jamaica; northern Macrurus sp. from Galapagos Islands; Coryphae­ puffer, Sphoeroides maculatus, from Woods Hole, noides abyssorum off California (Ho 1970). Mass.; spinyback puffer, S. marmoratus, from Chondrocanthus cottunculi Rathbun 1886. In gill cavity Jamaica (Ho 1967a). ofarctic sculpin, Cottunculus microps, and pallid scul­ pin, C. thompsonii, off New England coast (Ho 1971b). Chondracanthus merluccii (Holten 1802). In oral and LITERATURE CITED gill cavities of silver hake, Merluccius bilinearis, from Massachusetts to Florida; cape hake, M. capensis, off CRESSEY, R. F. South Africa; European hake, M. merluccius, in 1975. A new family of parasitic copepods (Cyc\opoida. Shiinoid.e). Crustace:.na 28:211·219. eastern North Atlantic (Ho 1971b). DELAMARE.DEBOUTIEVILLE. C. Chondracanthus nodosus (Muller 1776). In gill cavity 1962. Prodrome d'une faune d'Europe des copepodes parasites de of redfish, Sebastes marinus, in North Atlantic; deep­ poissons. Les copepodes Philichthyid.e. Bull. Inst. Oceanogr. water rosefish, S. mentella, in eastern North Atlantic; Monaco 57:1-44. HO..J.•8. Norway haddock, S. viviparus, in North Sea (Ho 1967a. Cyc\opoid copepods of the genus Tucea (Tuocidae), parasi­ 1971b). tic on diodontid and tetraodontid fishes. U.S. Fish Wild!. SeN.. Pseudochondracanthus diceraus Wilson 1908. On gills Fioh. Bull. 66:285-298. of smooth puffer, Lagocephalus laevigatus, from Gulf 1967b. Cyc\opoid copepods of the genus Telson par.sitic on ur'no· of Mexico; northern puffer, Sphoeroides maculatus, scopid fishes in the Gulf of Mexico. J. Parasitol. 53:852·858. 1970. Revision of the genera of the Chondracantbidae. a copepod from Massachusetts to North Carolina; southern family parasitic on marine fishes_ Beaufortia 17:105-218. puffer, S. neDhelus, from Gulf of Mexico; bandtail 1971a. Pharodes Wilson, 1935, genus ofcyclopoid copepods (pharo. puffer, &. sp~ngleri, from North Carolina to Florida didae) parasitic on marine fishes. J. Nat. Hist. 5:349-359_ (Ho 1970). 1971h. Parasitic copepods of the family Chondracanthidne from fishes of eastern North America. Smithonian Contrib. Zoo1., no. 87. 39 p. Family ERGASILIDAE HUMES. A. G.• and D. C. ROSENFIELD. Aflehi.~,troto,'i 1960. ocr:identalis C. B. Wilson l 1924 (Crustacea. Copepoda). a parasite of the orange fileflsh. CruBtaceana 1:179­ Ergasilus labracis Krglyer 1863. On gill filament of lS7. striped bass, J"forone saxatilis, from Massachusetts to IZAWA. K. Virginia (Roberts 1970). 1973. On the development of parasitic copepoda. L Sarcotacp... 10 paci/icu. Komlli (Cydopoidll: Philichthyidae). Publ. Seto Mar. of the life·histories nna ml1les of two Chinese species from food BioI. Lab. 21:77-86. fishes. re\'e81in~ their relationship with Lf.'rnut'u and of both to the PILLAI. N. K. CyciopoideA, Sinensill. New Ser. 1:51·84. 196.1. Redese!iption of seven species of homolochids from the col· VERVOORT. W. lections of the United States National Museum. J. Mar. BioI. 1969. CarihbeAn Bomolochidae (Copepoda: Cyclopoida). Stud. A"oc. India. 7:243·269. Fauna Cura,an lind nther Cacihb. Is. 28:1·125. WILSON. C. B. ROBERTS. L. S. 1911. North Amerielln parasitic copepads belongir,g to the family (Cnpepoda: Cyclopoidal: Revision and key to 1970. Erga.itu., Ergllsilidae. Proc, U.S. Nlltl. Mus, 39:263-400. species in North Amerira. Truns. Am. Microse. Soc. 89:134·161. YAMAGUTI. S. SPROSTON, N. G.• W. Y. YIN. and Y. T. HU. 196.1. Pllrasitic Copepoda lind BranchinCll of fishes. Inlerscience 1950. The genus I.amprog/eno (Copepoda PArasitical. the discovery Pub! .. New York. London. Sydney. 1104 p.

SYSTEMATIC INDEX

Acanthachandria manica/us . .... 10 carnuta 2,7,9 Grandiunguidae . .. .. 1 exilipes 7, )0 Grandiungus pramicraps ....1 galerita .. 7,10 Halabomalaehus albidus .. 3,9 phycidis 6,7, 10 Lernaea . .2 Acanthachandrites annulatus 6, 10 Lernaeidae . .1 Anchistrotas accidentalis · .. 4, 10 Nathobomolachus Blias prianati ..... · 5,6, 10 saetiger . ..9 Bomolochidae ...... 1,3,4,9 teres . .9 Chondracanthidae . ·. 1,3,9 Philichthyidae . 1, 3, 10 Chandracanthades deflexus . 8, 10 Philichthys xiphiae " .. 5, 10 Chandracanthus Pseudachandracanthus diceraus 5,6, 10 cottunculi . 9, 10 Shiinoidae ...... 1 merluceii .. 9, 10 Taeniacanthidae .1,3,4,10 nadasus ... 8, 10 Telsidae ...... 1 Ergasilidae .. ..1, 3, 4,10 Tucca impressus 4,5,10 Ergasilus . . . 2 Tuccidae ..... 1,3, 10 labracis ...... 4,10

11 ACKNOWLEDGMENTS

Preparation of the "Marine Flora and Fauna of the Ncrt~c' Roland L. Wigley, National Fisheries Services Northeast eastern United States" is bein~ coordinated by the following Fisheries Center, NOAA, Woods Hole, Mass. Board: Robert T. Wilce, Department of Botany, University of Massachusetts, Amherst, Mass. Coordinating Editor: Melbourne R Carriker, College of Mar­ ine Studies, University of Delaware, Lewes, DE 19958. The Board, which established the format for the "Marine Editorial Advisers: Flora and Fauna of the Northeastern United Stat.s," invites Marie R Abbott, Marine Biological Laboratory, Woods systematists to collaborate in the preparation of manuals, re­ Hole, Mass. views manuscripts, and

COORDINATING EDITOR'S COMMENTS

Pubtic;,tion of the "Marine Flora and Fauna of the North­ Copepoda. His studies have resulted in more than 50 papers on eastern United States" is most timely in view of the growing un­ the systematics of marine parasitic copepods, including a iversal emphasis on environmental work and the urgent need for monographic revision of Chondracanthidh" at the generic level. more precise and complete identification of coastal organisms Currently he is working On a series of copepod parasites from the than has been available. It is mandatory, wherever possible, fishes of the Great Barrier Reef, Australia. that orgnnisms be identified accurately to species. Accurate Preparation of this manual was supported in part by'a grant scientific names unlock the great quantities of biological infor­ from the Environmental Protection Agency to the Editorial mation stored in libraries, obviate duplication of research al­ Board of the "Marine Flora anr! Fauna of the Northeastern ready done, and often make possible prediction of attributes of United States." Work on the "Marine Flora and Fauna of the organisms that have been inadequately studied. Northeastern United States" by the Coordinating Editor is sup­ .Ju-shey Ho began his study of the systematics of the parasitic ported by the College of Marine Studies, University of Dela­ Copepoda in 1960 when he was a teachmg assistant at the ware. Department of Zoology, National Taiwan University, Taipei, Manuals are available for purehl!.se from the Superintendent Taiwan. In 1962 he went to Boston University to pursue of Documents, U.S. Government Printing Office, Washington, graduate studies on the copepod parasites of marine animals, DC 20402. The manuals so far published in the series are listed Ho joined the faculty of California State University, Long below: Beach. in 19',0 where he bas continued his research on para~itic

Marine Flora and Fauna ofthe Northeastern United States Circular COOK, DAVID G., and RALPH O. BRINKHURST. Annelida: Oligoehaeta. 374 BORROR, ARTHUR C. Protozoa: Ciliophora. 378 MOUL. EDWIN T. Higher Plants of the Marine Fringe. 384 McCLOSKEY, LAWRENCE R. Pycnogunida. 386 MANNING, RAYMOND B. Crustace,,: Stomatopoda. 387 WILLIAMS, AUSTIN B. Crustacea: Decapoda. 389 POLLOCK. LELAND W. Tardigrada. 394 LARSON, RONALD.J. Cnidaria: Scyphozoa. 397 CAVALIERE. A. R. Higher Fungi Ascomycetes, Deuteromycetes, and Basidiomycetes. 398 COULL, BRUCE C. Copepoda: Harpacticoida. 399 CUTLER. EDWARD B. Sipuncula. 403 PAWSON, DAVID L. Edhinodermnta: Holothuroidea. 405 HO, .JU-SHEY. Copepoda: Lernaeopodidae and Sphyriidae 406 HO, ./U·SHEY. Copepodn: Cyelopoids Parueitic on Fishes 409

12 '" GPO: 1978-799-756/9-10