NOAA Technical Report NMFS Circular 397 Marine Flora and Fauna of the Northeastern United States Cnidaria: Scyphozoa Ronald J. Larson August 1976 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of (he National Marine Fisheries Service (NMFSI are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of these resources, and to establish levels for optimum use of the resources. NMFS is also charged with the development and implementation of policies for managing national fishing grounds, development and enforcement of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the development and enforcement of international fishery agreements and policies NMFS also assists the fishing industry through marketing service and economic analysis programs, and mortgage insurance and vessel construction • It collects, analyzes, and publishes statistics on various phases of the industry. The NOAA Technical Report NMFS CIRC series continues a series that has been in existence since 1941. The Circulars are technical publications of general interest intended to aid conservation and management. Publications that review in considerable detail and at a high technical level certain broad areas of research appear in this series. Technical papers originating in economics studies and from management investigations appear in the Circular series. NOAA Technical Reports NMFS CIRC are available free in limited numbers to governmental agencies, both Federal and State. They are also available in exchange for other scientific and technical publications in the marine sciences. Individual copies may be obtained (unless otherwise noted) from D83, Technical Information Division. Environmental Science Information Center, NOAA. Washington. D.C. 20235. Recent Circulars are: 315. Synopsis of biological data on the chum salmon. Oneorhynchus keta iWalbaum' 1792. By Richard G. Bakkala. March 1970. iu + 89 p., 15 figs.. 51 tables. 319. Bureau of Commercial Fisheries Great Lakes Fishery Laboratory. Ann Arbor. Michigan. By Bureau of Commercial Fisheries. March 1970. 8 p., 7 figs. 330. EASTROPAC Atlas: Vols. 1-7. Catalog No. I 49.4:330 (vol.) 11 vols. Available from the Superintendent of Documents. U.S. Government Printing Office. Washington. D.C. 20402. 331. Guidelines for the processing of hot-smoked chub. By H. L. Seagran. J. T. Graikoski. and J. A. Emerson. January 1970. iv + 23 p.. 8 figs.. 2 tables. 332. Pacific hake. il2 articles by 20 authors.) March 1970. iii + 152 p.. 72 figs.. 47 tables. 333. Recommended practices for vessel sanitation and fish handling. By- Edgar W. Bowman and Alfred Larsen. March 1970. iv + 27 p., 6 figs. 335. Progress report of the Bureau of Commercial Fisheries Center for Estuarine and Menhaden Research. Pesticide Field Station, Gulf Breeze, Fla., fiscal vear 1969. By the Laboratory staff. August 1970. iii + 33 p., 29 figs.. 12 tables. 336. The northern fur seal. Bv Ralph C. Baker. Ford Wilke, and C. Howard Baltzo. April 1970. iii + 19 p., 13 figs. 337. Program of Division of Economic Research. Bureau of Commercial Fisheries, fiscal year 1969. By Division of Economic Research. April 1970, iii + 29 p.. 12 figs!. 7 tables. 338. Bureau of Commercial Fisheries Biological Laboratory, Auke Alaska. By Bureau of Commercial Fisheries. June 1970. 8 p.. 6 figs. Bay, 339. Salmon research at Ice Harbor Dam. By Wesley J. Ebel. April 1970, 6 p.. 4 figs. 340. Bureau of Commercial Fisheries Technological Laboratory, Gloucester, Massachusetts. By Bureau of Commercial Fisheries. June 1970, 8 p., 8 figs. 341. Report of the Bureau of Commercial Fisheries Biological Laboratory, Beaufort. N.C.. for the fiscal year ending June 30. 1968. By the Laboratory staff. August 1970. iii + 24 p.". 11 figs.. 16 tables. 342. Report of the Bureau of Commercial Fisheries Biological Laboratory. St. Petersburg Beach. Florida, fiscal year 1969. By the Laboratory staff. August 1970. iii + 22 p.. 20 figs., 8 tables. 343. Report of the Bureau of Commercial Fisheries Biological Laboratory. Galveston. Texas, fiscal year 1969. Bv the Laboratory staff. August 1970. iii + 39 p.. 28 figs.. 9 tables. 344. Bureau of Commercial Fisheries Tropical Atlantic Biological Laboratory progress in research 1965 69. Miami. Florida. By Ann Weeks. October 1970. iv + 65 p.. 53 figs.. 346. Sportsman's guide to handling, smoking, and preserving Great Lakes coho salmon. By Shearon Dudley. J. T. Graikoski, H. L. Seagran, and Paul M. Earl. September 1970. iii + 28 p.. 15 figs. 347. Synopsis of biological data on Pacific ocean perch, Sebastodes alutus. By Richard L. Major and Herbert H. Shippen. December 1970. iii + 38 p., 31 figv. 11 tables. 349. Use of abstracts and summaries as communication devices in technical articles. By F. Bruce Sanford. February 1971. iii + 11 p., 1 fig. 350. Research in fiscal year 1969 at the Bureau of Commercial Fisheries Biological Laboratory, Beaufort, N.C. By the Laboratory staff. November 1970, ii + 49 p.. 21 figs., 17 tables. 351. Bureau of Commercial Fisheries Exploratory Fishing and Gear Research Base. Pascagoula, Mississippi, July 1, 1967 to June 30, 1969. By Harvey R. Bullis, Jr. and John R. Thompson. November 1970. iv + 29 p., 29 figs., 1 table. 352. Upstream passage of anadromous fish through navigation locks and use of the stream for spawning and nursery habitat. Cape Fear River, N.C, 1962 66. By Paul R. Nichols and Darrell E. Louder. October 1970. iv -I- 12 p., 9 figs.. 4 tables. 356. Floating laboratory for study of aquatic organisms and their environ- ment. By George R. Snyder. Theodore H. Blahm, and Robert J. McConnell. May 1971, iii + 16 p.. 11 figs.. 361. Regional and other related aspects of shellfish consumption — some preliminary findings from the 1969 Consumer Panel Survey. By Morton M. Miller and Darrel A. Nash. June 1971, iv + 18 p.. 19 figs., 3 tables, 10 apps. 362. Research vessels of the National Marine Fisheries Service. By Robert S. Wolf. August 1971, iii + 46 p., 25 figs., 3 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 364. History and development of surf clam harvesting gear. By Phillip S. Parker. October 1971, iv + 15 p., 16 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 365. Processing EASTROPAC STD data and the construction of vertical temperature and salinity sections by computer. By Forrest R. Miller and Kenneth A. Bliss. February 1972, iv + 17 p., 8 figs., 3 appendix figs. For sale by the Superintendent of Documents. U.S. Government Printing Office, Washington, D.C. 20402. 366. Key to field identification of andromous juvenile salmonids in the Pacific Northwest. By Robert J. McConnell and George R. Snyder. January 1972. iv + 6 p., 4 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 20402. 367. Engineering economic model for fish protein concentration processes. By K. K. Almenas, L. C. Durilla, R. C. Ernst, J. W. Gentry, M. B. Hale, and J. M. Marchello. October 1972, iii + 175 p.. 6 figs.. 6 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 368. Cooperative Gulf of Mexico estuarine inventory and study, Florida: Phase I. area description. By J. Kneeland McNulty. William N. Linda!!, Jr., and James E. Sykes. November 1972. vii + 126 p., 46 figs., 62 tables. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 369. Field guide to the anglefishes (Pomacanthidae) in the western Atlantic. By Henry A. Feddern. November 1972, iii + 10 p., 17 figs., For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. Continued on inside back cover. NOAA Technical Report NMFS Circular 397 ATMOSPi ''Went of Marine Flora and Fauna of the Northeastern United States. Cnidaria: Scyphozoa Ronald J. Larson August 1976 o u o «> Q vi U.S. DEPARTMENT OF COMMERCE Elliot L. Richardson, Secretary National Oceanic and Atmospheric Administration Robert M. White, Administrator National Marine Fisheries Service Robert W. Schoning, Director For Sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Stock No. 003-017-00387 FOREWORD This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna of the Northeastern United States." This subseries will consist of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants and animals of the Northeastern United States. Manuals will be published at irregular intervals on as many taxa oi the region as there are specialists available to collaborate in their preparation. The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region." edited by R. I. Smith, published in 1964, and produced under the auspices of the Sys- tematics-Ecology Program. Marine Biological Laboratory, Woods Hole, Mass. Instead of revising the "Woods Hole Keys." the staff of the Systematics-Ecology Program decided to expand the geographic coverage and bathymetric range and produce the keys in an entirely new set of expanded pub- lications. The "Marine Flora and Fauna of the Northeastern United States" is being prpared in collaboration with systematic specialists in the United States and abroad. Each manual will be based primarily on recent and ongoing revisionary systematic research and a fresh examination of the plants and animals. Each major taxon, treated in a separate manual, will include an introduction, illustrated glossary, uniform originally illustrated keys, annotated check list with information when available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a systematic index. These manuals are intended for use by biology students, biologists, biological oceanographers, in- formed laymen, and others wishing to identify coastal organisms for this region. In many instances the manuals will serve as a guide to additional information about the species or the group. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is plan- ned to include organisms from the headwaters of estuaries seaward to approximately the 200-m depth on the continental shelf from Maine to Virginia, but may vary somewhat with each major taxon and the interests of collaborators. Whenever possible representative specimens dealt with in the manuals will be deposited in the reference collections of major museums in the region. After a sufficient number of manuals of related taxonomic groups have been published, the manuals will be revised, grouped, and issued as special volumes. These volumes will thus consist of compilations of individual manuals within phyla such as the Cnidaria, Arthropoda, and Mollusca, or of groups of phyla. CONTENTS Introduction 1 Description of orders of Scyphozoa 1 Methods Collecting 4 Rearing 4 Preservation 4 Nematocysts 4 Natural history Stauromedusae 4 Scyphistomae 4 Ephyrae 4 Scyphomedusae 4 Stinging medusae 5 Glossary 5 Introduction to the keys to the Scyphozoa 6 Key to the Stauromedusae of the northeast coast of the United States 6 Synopsis of ephyrae of Chrysaora, Cyanea, and Rhopilema 9 Key to the pelagic scyphomedusae of the northeast coast of the United States 10 Annotated systematic list 14 Selected bibliography 16 Index to scientific names 17 Acknowledgments 18 Coordinating editor's comments 18 The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. Digitized by the Internet Archive in 2013 http://archive.org/details/marineflorafaunaOOIars Marine Flora and Fauna of the Northeastern United States. Cnidaria: Scyphozoa RONALD J. LARSON 1 ABSTRACT This manual is an introduction to the scyphomedusae found in coastal waters from Maine to the Chesapeake Bay. It includes a discussion of their identification, collection, rearing, preservation, and nematocysts. Also included is an introduction to the natural history of the scyphopolyps and medusae, a discussion of stinging scyphomedusae, a glossary of terms, an illustrated synopsis of ephyrae, an il- lustrating key to the scyphomedusae (including the Stauromedusae), an annotated systematic list, a bibliography of major references, and finally an index. INTRODUCTION The class Scyphozoa of the phylum Cnidaria or Coelenterata as it once was called, are commonly known as "jellyfish," "sea nettles," and "sea wasps." Scyphomedusae, as these jellyfish are termed, are the most conspicuous of the jellylike animals. Some species reach 1 m or more in diameter and most are very color- ful. All have stinging organs, hence the names "sea nettle" and "sea wasp." Scyphomedusae are seasonal; most species are seen during the warmer months. They represent the sexual stage of most scyphozoan species. The asexual stage is a small benthic (bottom-living) polyp (scyphopolyp) which is perennial. The scyphopolyp generally buds larval scyphomedusae (ephyrae) during the spring. The medusae are large by midsummer. Scyphomedusae have a muscular saucer-shaped or hemispherical bell or umbrella which propels the medusae through the water by contracting and expelling water behind. Long threadlike tentacles often hang from the umbrella and are covered by stinging organs (nematocysts) which are used to capture prey. Sur- rounding the mouth are ribbonlike, curtainlike, or gelatinous arms which transport prey to the mouth. Scyphopolyps are minute and rarely seen. They are saclike with a circle of tentacles around the oral end and are attached to the bottom by a stalklike peduncle. Scyphozoans usually have a scyphomedusa and a scyphopolyp stage in their life history but one group remains as a polyp only. The Stauromedusae or stalked medusae lack a medusa stage. Some scyphomedusae lack a polyp stage; Pelagia, an oceanic medusa, has eggs which, when fertilized, transform directly into a juvenile medusa and bypass the normal polyp stage. 'Department of Marine Sciences, University of Puerto Rico, Mayaguez, Puerto Rico 00708; present address: Division of Echinoderms, U.S. National Museum, Washington, D.C. 20560. Scyphomedusae are generally regarded as pests because of their irritating stings, but many are ecologically important. Chrysaora, well known as the sea nettle, consumes large numbers of ctenophores which might otherwise be very detrimental to oyster and clam populations by feeding on their larvae. Other scyphomedusae prey on jellyfish which have few predators and feed on commercially important inver- tebrate and fish larvae. Five orders of Scyphozoa are recognized. The Stauromedusae, or "stalked jellyfish," are not well known even though they may be abundant. They are in- active and cryptically pigmented and are difficult to see when attached to algae. The Cubomedusae, or sea wasps, are also infrequently seen but are infamous because of their potent sting. The Coronatae, or "crown medusae," are mostly deepwater (bathypelagic) . The Semaeostomeae, commonly known as sea nettles, are the most familiar order, and most shallow-water scyphozoans belong to this group. The Rhizostomeae, which lack tentacles, are mostly tropical and are un- common in New England waters. DESCRIPTION OF ORDERS OF SCYPHOZOA Order Stauromedusae — Small benthic scyphozoans which lack a free-swimming medusa stage. Stauromedu- sae are urn- or funnel-shaped and are attached to the substrate by a stalklike peduncle. In some forms, the margin is divided into eight arms, the knobbed tentacles project from the arm tips; in others the tentacles project from the margin. Modified tentacles, which may act as anchors or sense organs, occur on the margin between the groups of tentacles in some species. The gonads are paired, leaflike, or folded structures extending along the length of the calyx. A wormlike planula larva develops from the zygote and creeps over the substrate, even- tually attaches itself and becomes a juvenile stauromedusa. Stauromedusae occur mainly in cold, shallow marine waters. Adult Stauromedusae usually oc- cur only during the summer, and the wintering juveniles may be difficult to find because of their small size. Tropical and deepwater forms are rare. Order Cubomedusae (Fig. 1) — Small- to medium-sized medusae with a somewhat cuboidal or rectangular- shaped transparent umbrella. A velarium, or shelflike projection, encircles the inner margin of the umbrella. Tentacles vary from four to many and are on bladelike tions of the bell, termed pedalia. Four rhopalia are located about midway between the corners of the um- brella near the margin. The mouth is at the end of a flask-shaped stomach which hangs within the umbrella. The life cycle of the Cubomedusae is unlike that of other scyphozoans. The cubopolyp has several features in com- mon with hydrozoan polyps, e.g., solid tentacles, steno- tele nematocysts. Cubopolyps metamorphose directly and totally into a single medusa, other scyphopolyps undergo transverse fission producing one to many ephy- rae. Werner (1973) has placed the Cubomedusae in their own class, Cubozoa, because of this group's unique char- acteristics. Cubomedusae are tropical or subtropical me- Figure 1. — Cubomedusa. Figure 2. — Coronate medusa. Figure 3.— Semaeostome medusa. Figure 4.— Rhizostome medusa. dusae but are occasionally transported into temperate regions by warm currents. Cubomedusae are known as sea wasps because of their virulent sting. Order Coronatae (Fig. 2) — Small- or medium-sized medusae with flattened or dome-shaped umbrella which is divided into two regions by a circular (coronal) furrow. The tentacles occur between marginal lappets. Radial thickenings of the margin, termed pedalia, give rise to the tentacles, lappets, and rhopalia. The gonads are on four septa which project into the coelenteron. The Coronatae are mostly bathypelagic and darkly pig- mented. The coronate polyp, all known species being placed in the genus Stephanoscyphus, is covered by a chitinous cuticle which has numerous circular an- nulations (Kramp 1959). Order Semaeostomeae (Fig. 3) — Large medusae, usually saucer-shaped. The mouth has four long oral arms. These may be folded once as in Pelagia, or folded a number of times as in Cyanea. Tentacles occur between marginal lappets as in the Coronatae, or on the subum- brella. Gonads occur on folds of the gastrodermis. The polyp stage is small and naked or has a thin cuticle around its base and can encyst under certain conditions (Fig. 7). A typical ephyra (Fig. 5) is produced, usually by polydisc strobilation (Fig. 9). Semaeostomes are the most common scyphomedusae in temperate coastal waters. Order Rhizostomeae (Fig. 4) — Large medusae, usually with a hemispherical umbrella. Marginal tentacles are absent, but eight oral arms serve both to capture and transport food to the gastric cavity by way of a canal system. A central mouth is usually absent; instead the mouth is formed by a large number of bifurcating grooves which lie on the ventral and lateral sides of the oral arms. The oral arms may have terminal appendages which have a high concentration of nematocysts at their tip. The gonads are folds of the gastrodermis as in the semaeostomes. The polyp stage has a long stalklike peduncle which is partially covered with a thin cuticle. Ephyrae are usually produced by monodisc strobilation (Fig. 10). Rhizostomes are mostly tropical, with only a few species extending into temperate waters. lappet rhopalium radial canal gastric cirri mouth rhopaliar cleft .tentacles gastric cirri- \^ /i"n^ anchor peduncle Figure 5. — Ephyra, newly released. Figure 6. — Stephanoscyphus. Figure 7. — Semaeostome polyp. Figure 8.— Stauromedusa. 10 Figure 9. — Semaeostome strobila. Figure 10. — Rhizos tome strobila. METHODS Collecting To obtain medusae in the best possible condition they should be dipped from the water surface using a bucket or other large container. Deepwater species can be caught in plankton nets equipped with large sample buckets. Scyphopolyps can be obtained by dredging, collecting at low tide, or diving. Polyps commonly attach to shells, oysters being a favorite substrate. Polyps will also settle on glass slides placed in areas where medusae spawn. Rearing Scyphozoans can be raised from embryos to maturity in the laboratory (Spangenberg 1965). Polyps can be reared in small dishes and fed newly hatched Artemia. Planktonic stages are more difficult to raise, requiring a large volume of circulating water and suspended food. Medusae should be fed daily, preferably more often. Artemia can be fed to such plankton-feeding forms as Aurclia and rhizostome medusae. Chrysaora can be fed ctenophores, Artemia, or other medusae. (See Russell 1970.) Preservation Scyphozoans should be fixed in a 20°? formaldehyde solution and stored in a 5°} neutralized formaldehyde solution. Alcohol dehydrates specimens, making them distorted. Ordinary paper disintegrates in Formalin, thus labels should be made from high-rag content and water-resistant paper. NEMATOCYSTS Nematocysts occur in all cnidarians. They are produced by specialized cells, cnidoblasts. A great amount of work has been done on the cnidoblast and how the nematocyst functions (Picken and Skaer 1966). Weill U934) showed that the cnidome is taxonomically impor- tant; and recently Calder (1971) used nematocysts as a means of identifying polyps of Chrysaora, Cyanea, and Aurelia. (Also see Calder 1974.) with crustaceans being the major part of the food. Berrill (1962) noted that Lucernaria fed on the amphipod Amphithoe and the small gastropod Lacuna. At Woods Hole, Mass., I found mostly harpacticoid copepods and gammarid amphipods, and a few chironomid fly larvae in gastric cavities of Craterolophus. Prey, caught by the tentacles, cause one or all arms to fold over the mouth which then expands and engulfs the food. Stauromedusae, like scyphistomae, occur year-round. Berrill (1962) found that Stauromedusae, along the coast of Maine, spawn during the summer and then die. Juveniles appear during the fall and become sexually mature by summer. Apparently Stauromedusae are very sensitive to en- vironmental conditions and have been difficult to raise in aquaria. Berrill (1962) remarked that they are becoming rare in Massachusetts Bay due to pollution. Scyphistomae Scyphistomae are the polyp stages of scyphomedusae; in the semaeostome (Fig. 7) and rhizostome medusae, they are small, about 2-4 mm high, and naked, or with a chitinous theca around the peduncle. They are flask- shaped with a single whorl of tentacles and a cruciform mouth. They are often found in shallow water attached to shells or other substrates. Those of the coronate medusae are completely covered by a chitinous theca and are found in deeper water. Scyphistomae feed on small crustaceans and can be reared in the laboratory on a diet of Artemia nauplii. They undergo asexual reproduction to form more polyps. Some produce resistant stages known as pedal cysts which can resist temperature extremes at which medusae are not found (Cargo and Schultz 1967). Scyphistomae also undergo strobilation, a process which produces from one to many ephyrae by transverse constrictions of the polyp's oral region (Figs. 9, 10). Scyphistomae are difficult to identify. Calder (1971) used nematocysts to aid in identifying the polyps of Aurelia, Chrysaora, and Cyanea. They can also be reared in the laboratory until they strobilate and then the ephyrae or postephyrae can be identified. Increased feeding and/or change in water temperature or addition of thyroxin may induce strobilation. NATURAL HISTORY Stauromedusae Stauromedusae, contrary to what their name implies, are really polyplike. Some are permanently attached. Others can move in a somersaulting motion by adhering to the substrate with the oral end and releasing the pedal disc, then reattaching the disc at a new location; but none have been observed to swim. They attach to algae, sea grass (Zostera), and other substrates in shallow areas which have adequate water circulation. They may be abundant, but because of their inconspicuous coloration and inactivity, they are rarely seen. Most of the diet consists of other benthic animals, Ephyrae Ephyrae (Fig. 5) are small, 1-4 mm in diameter, flat- tened larval medusae which usually have eight arms. At the tip of each arm there is a pair of blunt or pointed lappets, and between the lappets, a rhopalium or sense organ. The mouth is cruciform and a few gastric cirri may occur in the coelenteron. Ephyrae feed on plankton and grow rapidly. They soon appear like small scypho- medusae, at which time they can usually be identified with certainty. (See synopsis of Ephyrae.) Scyphomedusae By being planktonic, scyphomedusae may find food, which is unavailable to the polyp, and can also dis- tribute gametes over an extensive area thereby in- creasing the distribution of the species and also insuring that uninhabited substrates are utilized if conditions are favorable for the polyp. Scyphomedusae are usually short-lived. They grow rapidly and later die due to changes in water tem- perature or other factors, which for the most part are not understood. Chrysaora ephyrae from the Chesapeake Bay are released in early June. By July these medusae are mature, and most have died by mid-September. Aurelia and Cyanea may live somewhat longer. The occurrence of some scyphomedusae is related to water temperature. Cyanea in the Gulf of Maine are first seen in April or May and are mature by early summer (Bigelow 1926). Cyanea is found in the Chesapeake Bay early in the winter and matures by February or March. Chrysaora occurs only at higher temperatures (about 20° C) in the Chesapeake Bay in early May, and slightly later at Woods Hole, Mass. Because of this dependence on high temperatures, it apparently cannot strobilate in colder water and is not found north of Cape Cod. Aurelia, whose range overlaps the two previous species, occurs during the warmer months in both the Gulf of Maine and in the Chesapeake Bay, but appears earlier further south. In the Gulf of Mexico and off Florida, both Aurelia and Chrysaora can occur at almost any time. Rhopilema and Stomolophus also may occur nearly year-round. Medusae feed on a variety of prey. These include ctenophores, planktonic microcrustacea (i.e., copepods, amphipods, and larvae), larval and small fishes, pelagic polychaetes, siphonophores, and even other medusae. Fraser (1969) calculated that a Cyanea less than one-half meter in diameter would eat over 1.5 million copepods of the genus Calanus in its lifetime. One can only guess at the amount of food eaten by a 2-m Cyanea. Chrysaora feeds mostly on ctenophores and other medusae. Studies which I have made in Puerto Rico (unpublished) indicate that a Chrysaora medusa would have to consume about 15 kg of ctenophore Mnemiopsis in order to attain a diameter of 150 mm. In Aurelia, food is trapped on the umbrella and by the short tentacles, and is transferred, not directly to the lips, as in most medusae, but to a food groove formed by the small velarium. Cilia move the food which is mixed with mucus to the adradial canals where it is then transferred to the lips (Southward 1955). Rhizostomes are filter feeders and many are active swimmers. In forms such as Rhopilema, water is forced around the oral arms by the contracting umbrella during swimming. In Stomolophus, water is forced through the keellike appendages. Minute tentacles, which line these structures, filter the suspended food from the water. The food is then carried to the stomach by the cilia-lined canals. Stomolophus collected at Beaufort, N.C., fed mostly on bivalve veligers and harpacticoid copepods (author, unpublished). The gonads of scyphozoans are always near the site of digestion. In semaeostomes and rhizostomes the gonads are infoldings of the subumbrella which also bear the gastric cirri. Some scyphomedusae spawn while still very small. Chrysaora reared by the author began spawning at the size of 55 mm and spawned almost nightly thereafter. Fertilization usually takes place within the gastric cavity or on the surface of the ovary. In Cyanea and Aurelia the embryos are retained on the oral arms. Aurelia has brood pockets on the oral arms of the female where embryos are retained until the planula is released. The embryology is quite variable even within the same species. Segmentation is usually total but gastrulation depends upon egg size, with ingression in small eggs and invagination in large ones and by both methods in medium-sized ova. Usually a planula is produced, which settles and forms a polyp, but very large eggs may bypass both the planula and polyp and form a modified ephyra as they do in Pelagia (Berrill 1949). Symbiosis is widespread in the Cnidaria and occurs frequently in the Scyphozoa. There are numerous reports of juvenile fishes associated with scyphomedusae; Men- sueti (1963) discussed these associations. Associations between juvenile fish and medusae range from the fish seeking refuge under the umbrella, or between the ten- tacles, to predation with the larger fish eating bits of the tentacle or gonad. It is not fully understood how certain species of fishes, e.g., Peprilus and Poronotus, are able to feed on medusae without ill effects, but it is generally thought that antibodies may be involved and that the secretion of mucus by the fish may prevent the nematocysts from discharging. Crustacea are also known to parasitize scyphomedusae; hyperiid amphipods may both steal food from medusae and feed on medusal tissue. Phillips et al. (1969) discussed the parasitic habits of the two species of brachyuran crabs on medusae in the Gulf of Mexico. STINGING MEDUSAE Chrysaora is popularly known as the sea nettle because its sting can be very painful. Cyanea also is a stinger; Russell (1970) noted that Cyanea was the cause of a mysterious death in Conan Doyle's story, "Adventure of the Lion's Mane." The majority of the Cubomedusae are stingers, as are coronate medusae, some producing a very severe sting. Aurelia, Rhopilema, Stomolophus, Phacellophora, and the Stauromedusae are not known to be severe stingers. Halstead (1965) gave an account of the nature of the wounds and medical aspects of stings. Cargo and Schultz (1967) found that when meat ten- derizers containing proteolytic enzymes were mixed with water and applied to the affected area, the pain was quickly relieved. GLOSSARY anchors (Fig. 8) adhesive organs between arms of some Stauromedusae. calyx funnel-shaped umbrella of Stauromedusae. cnidoblast cell in which the nematocyst forms. cnidome complement of nematocyst types present. coelenteron gastric cavity or stomach. coronal furroiv (Fig. '2) circular groove which divides the exumbrella of the coronate medusae. cruciform cross-shaped. • : (Fig. 5) planktonic larva of Scyphozoa except Cubomedusae and Stauromedusae. Usually with 8 pairs of arms and 8 rhopalia. exumbrella aboral or upper surface of umbrella. gastric cirri or phacellae (Fig. 1) hollow fingerlike pro- jections in the coelenteron which aid in digestion. flaplike structure making up the scalloped mar- gin oi the umbrella. disc strobilation (Fig. 10) a single ephyra produced per strobila at one time. oral arm armlike extensions of the corners of the mouth which hang from the subumbrella of the Scyphozoa, 4 in number in the semaeostomes (Fig. 3) and 8 in the rhizostomes (Fig. 4). pedalia radial thickenings between lappets and coronal furrow of the coronate medusae (Fig. 2): bladelike pro- jections of Cubomedusae to which the tentacles are attached (Fig. 1). peduncle (Fig. 8) stalk of polyps, or Stauromedusae. polydisc strobilation (Fig. 9) several ephyrae produced from a polyp at the same time. rhopalium (Figs. 1, 2) hollow, club-shaped sense organ generally located near the umbrella margin, usually with a statocyst and, rarely, an ocellus or ocelli. scyphistoma polyp of scyphozoan medusae excluding the Stauromedusae and Cubomedusae. sensory niche open structure partially enclosing rho- palium in Cubomedusae. strobila scyphistoma which is forming ephyrae. st?-obilation process of producing ephyrae. subumbrella oral or underside of umbrella. velarium (Fig. 1) shelflike structure at margin of um- brella in Cubomedusae and Aurelia. INTRODUCTION TO THE KEYS TO THE SCYPHOZOA An attempt has been made in constructing the keys to include those features which are most obvious or which occur in both juvenile and adult specimens. As medusae grow, structures such as tentacles usually increase in number and oral arms become increasingly folded or complex, while other structures, e.g., rhopalia, usually remain unchanged. Because of the delicate nature of medusae they are often injured and parts may be missing or abnormal. Therefore, in using the key some discretion must be used for each specimen to determine which characters are usable. Key to the Stauromedusae of the Northeast Coast of the United States 1 With anchors 2 1 Without obvious anchors 4 2 I 1 1 Anchors large; no shieldlike covering; gonads extend into arms 3 2 (/) Anchors small with shieldlike covering at base; arms short; gonads do not extend into arms Thaumatoscyphus atlanticus Figure 11.— Thaumatoscyphus atlanticus. a. side view b. oral view c. anchor, side view d. anchor, front view 3 (2) Arms short; anchors bean-shaped; calyx as high as broad Haliclystus auricula Figure 12. — Haliclystus auricula. a. side view c. anchor, side view b. oral view d. anchor, front view 3 (2) Arms long; anchors trumpet-shaped with short projection at center; calyx broader than high Haliclystus salpinx Figure 13.— Haliclystus salpinx. a. side view c. anchor, side view b. oral view d. anchor, front view 4 (1) Peduncle and arms long; arms paired; calyx broader than high; largest New England Stauro- medusa Lucernaria quadricornis Figure 14. — Lucernaria quadricornis. a. side view b. oral view / (1) Peduncle and arms short; arms not paired; calyx higher than broad Craterolophus convovulus Figure 15.— Craterolophus convovulus. a. side view b. oral view Synopsis of Ephyrae of Chrysaora, Cyanea, Aurelia, and Rhopilema. (See Russell 1970.) Newly released ephyrae ca. 2.5 mm in diameter (Fig. 5) Shape of lappets; position of rhopalia Morphology of ephyral arms Gastric cirri Postephyrae 5 mm in diameter Morphology Chrysaora (Fig. 16a, b) lappets slender pointed tips; rhopaliar cleft deep ca. 1/2 arm length a pair of nematocyst clusters on each arm; radial canals in arms short, consists of 1 pair lack oral arms split; long ribbonlike; marginal tentacles; no ring canal Cyanea (Fig. 16c, d) lappets slender pointed tips; rhopaliar cleft deep ca. 1/2 arm length radial canals in arms long, consists of 2 pairs usually 4 oral arms not split, wide and flaring; subumbrellar tentacles; no ring canal Aurelia (Fig. 16e, f) lappets broad, tips blunt, rounded; rhopaliar cleft short 1/2 arm length radial canal to rhopalium only with 4 or more oral arms split; short; marginal tentacles; ring canal present Rhopilema (Fig. 16 g, h) lappets slender, tips pointed; rhopaliar cleft deep ca. 1/2 arm length radial canals in arms long, consists of 1 pair with 4 or more oral arm split several times; no marginal or subumbrellar tentacles Figure 16. — Ephyrae (a, c, e, g); postephyrae (b, d, f, h). a, b Chrysaora quinquecirrha c, d Cyanea capillata e, f Aurelia aurita (from D. Calder 1972, unpublished; and Calder 1973) g, h Rhopilema verrilli Key to the Pelagic Scyphomedusae of the Northeast Coast of the United States 1 Umbrella, saucer-shaped, dome-shaped, or hemispherical; with 8 or more tentacles, or no tentacles; with 8 01 more rhopalia 2 1 Umbrella tall rectangular; 4 tentacles; 4 rhopalia; sensory niche with covering scale above and below; stomach supported by 4 mesenteries Order Cubomedusae Tamoya haplonema Figure 17. — Tamoya haplonema. a. side view b. sensory niche, front view 2 (7) Umbrella divided into two regions by a deep circular groove; stomach darkly pigmented; bathy- pelagic. rarely at surface or coastal Order Coronatae 3 2 (1) Umbrella not divided by circular groove, not darkly pigmented; mostly coastal or at surface 4 (2) Umbrella flat, disc-shaped; tentacles and rhopalia equal in number, usually 22; gonads button- like Atolla wyvilld Figure 18. — Atolla wyvillei, side view. 10 3 (2) Umbrella conical or dome-shaped; usually 3 tentacles between rhopalia; 4 rhopalia; 12 ten- tacles; gonads horseshoe-shaped Periphylla periphylla Figure 19. — Periphylla periphylla, side view. 4 (2) With marginal or subumbrella tentacles; 4 oral arms; distinct mouth; saucer-shaped umbrella Order Semaeostomeae . . 4 (2) Without tentacles on umbrella; 8 tough gelatinous oral arms; mouth small or absent; umbrella usually hemispherical Order Rhizostomeae . . 5 (4) With numerous narrow radial canals extending from stomach to margin; ring canal near mar- gin; tentacles either on margin, short and numerous or in 16 linear groups on subumbrella Family Ulmaridae . . Figure 20. — Family Ulmaridae, radial and ring canals in black. Figure 21. — Cyanea and family Pelagidae, stomach pouches in black. 5 (4) With broad stomach pouches extending to margin; without ring canal; tentacles long or 8 U- shaped subumbrellar groups 11 6 (5) Oral arms ribbonlike; tentacles on margin Family Pelagidae 7 6 (5) Oral arms wide, curtainlike; tentacles in 8 U-shaped groups on subumbrella Cyanea capillata Figure 22.— Cyanea capillata. a. side view b. oral view, oral arms removed and four groups of tentacles removed {6) With 8 tentacles; exumbrella with prominent wartlike projections; umbrella nearly hemis- pherical in shape, usually less than 100 mm in diameter; oceanic Pelagia noctiluea Figure 23. — Pelagia noctiluea. a. side view b. oral view, oral arms removed (6) With more than 8 tentacles; exumbrella smooth; umbrella saucer-shaped, up to 200 mm in diameter; estuarine or coastal Chrysaora quinquecirrha Figure 24. — Chrysaora quinquecirrha. a. side view b. oral view, oral arms removed 12 8 (7) Marginal tentacles short and very numerous; oral arms folded, ribbonlike; gonads in 4 horse- shoe-shaped groups; 8 rhopalia Aurelia aurita Figure 25. — Aurelia aurita. a. side view b. oral view, oral arms removed (7) Tentacles long, in 16 subumbrellar linear groups; oral arms broad, curtainlike; 16 rhopalia Phacellophora camtschatica Figure 26.— Phacellophora camtshatica. a. side view b. oral view, oral arms and one- half of tentacles removed 13 9 (4) Oral arms tree along most of their length; spindlelike appendages hang from oral arms; um- brella hemispherical or flattened Rhopilema verrilH Figure 27 '.— Rhopilema verrilli, side view. 9 (-/) Oral arms fused: no spindlelike appendages; umbrella higher than a hemisphere (globular) Stomolophus meleagris Figure 28. — Stomolophus meleagris, side view. ANNOTATED SYSTEMATIC LIST The following list is arranged according to the classification of Kramp (1961). Distribution notes are from published records and from museum specimens. Most of the species listed are represented by specimens in the U.S. National Museum, Washington, D.C., and the Gray Museum, Marine Biological Laboratory, Woods Hole, Mass. Notes on seasonal occurrence, habitat and life history, and coloration are included. References to important papers are cited for both the families and for each species. Class SCYPHOZOA Order STAUROMEDUSAE Family ELEUTHEROCARPIDAE References: Berrill (1963); Kramp (1961). Haliclystus auricula (Rathke 1806) Massachusetts northward, Northeast, Atlantic, North Pacific, lower intertidal and subtidal on algae and Zostera, at Woods Hole most common on brown algae Scytosiphon. Juveniles easily confused with H. salpinx, but arms are very short. Coloration: highly variable, brown, green, red. Berrill (1962); Mayer (1910). Haliclystus salpinx Clark 1863 Massachusetts northward, lower intertidal and sub- tidal on algae and Zostera. May be confused with juvenile H. auricula but with long arms. Coloration: variable, green, brown. Berrill (1962, 1963). Lucernaria quadricornis O. F. Muller 1776 Massachusetts northward, Northeast Atlantic, Arc- tic, lower intertidal and subtidal mostly on Laminaria. Largest New England stauromedusa M reaching over 50 mm in length. Coloration: highly variable, brown, green, red. Berrill (1962, 1963). Family CLEISTOCARPIDAE References: Berrill (1963); Kramp (1961). Craterolophus convolvulus (Johnston 1835) Massachusetts, Northeast Atlantic, intertidal on Fucus where it attaches near the bladders, incon- spicuous because of its shape and color. Colora- tion: olive green when attached to Fucus. Kramp (1961). Thaumatoscypus atlanticus Berrill 1962 Maine, subtidal on Laminaria. This species is un- usual in that ocelli are located on the anchors. Coloration: not known. Berrill (1962, 1963). Order CUBOMEDUSAE Family CARYBDEIDAE References: Bigelow (1938); Kramp (1961). Tamoya haplonema F. Muller 1859 Tropical Atlantic, Gulf of Mexico to Massachusetts. Probably carried north in the Gulf Steam. Mayer's (1910) report of this species in Long Island Sound is the only report of T. haplonema in the New England coastal waters. Coloration: colorless except ten- tacles, pink. Bigelow (1938); Phillips and Burke (1970). Three other Cubomedusae also occur in the northwestern Atlantic. Carybdea alata Reynaud, a specimen in the U.S. National Museum (USNM) was collected off Cape Hatteras at lat. 35°N, long. 75°W. Carybdea marsupialis (Linneaus) occurs in the Gulf Stream near Bermuda but has not been reported on our coast. Chiropsalmus quadrumanus (Muller) occurs along U.S. coast south of Virginia. Bigelow (1938); Phillips and Burke (1970). Order CORONATAE Family ATOLLIDAE References: Kramp (1961); Russell (1970). Atolla wyvillei Haeckel 1880 Cosmopolitan in deep oceans. A deep-sea species, bathypelagic, seldom occuring at the surface or on the continental shelf. Commonly taken in midwater samples in deep water usually below 500 m. Coloration: stomach dark red-brown. Russell (1970). Family PERIPHYLLIDAE References: Kramp (1961); Russell (1970). Periphylla periphylla (Peron and Lesueur 1809) Cosmopolitan bathypelagic in deep oceans. Com- mon in midwater trawl samples from deep water. It has been taken near the surface off New England. Coloration: stomach dark red- brown. Russell (1970). At least four other coronate medusae may also occur in the northwestern Atlantic: Linuche unguiculata (Schwartz), a specimen at the U.S. National Museum, collected off Cape Hatteras, lat. 35°N, long. 75°W. Nausithoe punctata Kolliker, a specimen at the USNM collected off Georges Bank, lat. 40°N, long. 68° W. Nauphantopsis diomedeae Fewkes, not reported since the original description, off New England. (Kramp 1961.) Atolla vanhoeffeni Russell, a specimen at the USNM, collected off Virginia lat. 36°39'N. long. 74°39'W. (Russell 1970.) Order SEMAEOSTOMEAE Family PELAGIDAE References: Kramp (1961); Russell (1970). Chrysaora quinquecirrha (Desor 1848) Possibly cosmopolitan, western Atlantic distribu- tion from Cape Cod to the Gulf of Mexico. Abun- dant during the summer months in estuaries, not common in the open ocean. Scyphistoma occurs on oyster shells in upper estuary. Popularly called "sea nettle" because of its stinging ability. Coloration: highly variable, pink, yellow, often with 16 yellow- ocher or red radiating stripes. Cargo and Schultz (1966, 1967). Pelagia noctiluca (Forskal 1775) Cosmopolitan open ocean species in warm and tem- perate oceans. Carried north in the Gulf Stream, sometimes seen in coastal New England waters. Lacks scyphistoma stage. Coloration: highly variable, blue, yellow, pink, tentacles red. Russell (1970). Family CYANEIDAE References: Kramp (1961); Russell (1970). Cyanea capillata (Linnaeus 1758) Cosmopolitan in colder oceans. Occurs in the Gulf of Mexico and Florida northward. Most abundant during winter through spring in the south, summer to fall in the north (New England waters). Popularly called "lion's mane" because of the long red-yellow tentacles. Largest New England medusa reaching a meter or more in diameter. Coloration: highly variable, pink, red, yellow, brown. Russell (1970). Family ULMARIDAE References: Kramp (1961); Russell (1970). Aurelia aurita (Linnaeus 1758) Cosmopolitan species in nearly all oceans except the Arctic. Popularly known as "moon jelly" because of the whitish umbrella. Coloration: bell clear; gonads, tentacles, pink, purple. Russell (1970). Phacellophora camtschatica Brandt 1838 Occurs north of Cape Cod in the western Atlantic, probably the entire Pacific coast of the Americas and in the Mediterranean and off western Africa. Rarely taken in New England waters. Coloration: yellowish. Mayer (1910). 15 Order RHIZOSTOMEAE Family RHIZOSTOMATIDAE References: Kramp (,1961); Mayer (1910). Rhopilema verrilli (Fewkes 1887) Occurs in the northern Gulf of Mexico, North Carolina to Connecticut. An uncommon species, found along coast and in mouths of es- tuaries. Coloration: bell milky; oral arms brown. Mayer (1910); Calder (1973). Family STOMOLOPHIDAE References: Kramp (1961). Stomolophus meleagris L. Agassiz 1862 Occurs in the western Atlantic from Brazil to Cape Hatteras and from Panama to San Diego in the eastern Pacific. Specimens occasionally carried north to New England waters by the Gulf Stream. Popularly known as the "cabbage head jellyfish" because of its globular shape. Coloration: brown band at margin. Mayer (1910). SELECTED BIBLIOGRAPHY BERRILL. M 1962. The biology of three New England Stauromedusae, with a description of a new species. Can. J. Zool. 40:1249-1262. 1963. Comparative functional morphology of the Stauromedusae. Can. J. Zool. 41:741-752. BERRILL. N. J. 1949. Developmental analysis of Scyphomedusae. Biol. Rev. iCamb.) 24:393-410. BIGELOW. H. B. 1926. Plankton of the offshore waters of the Gulf of Maine. Bull. U.S. Bur. Fish. 40:1-509 (see p. 340-380). 1938. Plankton of the Bermuda Oceanographic Expeditions, VIII. Medusae taken during the years 1929 and 1930. Zoologica (N.Y.) 23:99-189. CALDER. D. R. 1971. Nematocysts of polyps of Aurelia, Chrysaora, and Cyanea, and their utility in identification. Trans. Am. Microsc. Soc. 90:269-274. 1973. Laboratory observations on the life history of Rhopilema uerrilli (Scyphozoa: Rhizostomeae). Mar. Biol. (Berl.) 21:109- 114. ;974 Nematocysts of the coronate scyphomedusa, Linuche un- guiculata. with a brief reexamination of scyphozoan nematocyst classification. Chesapeake Sci. 15:170-173. CARGO, D. G., and L. P. SHULTZ. 1966. Notes on the biology of the sea nettle, Chrysaora quinque- cirrha. in Chesapeake Bay. Chesapeake Sci. 7:95-100. 1967. Further observations on the biology of the sea nettle and jelly- fishes in Chespeake Bay. Chesapeake Sci. 8:209-220. FRASER, J. H. 1969. Experimental feeding of some medusae and Chaetognatha. J. Fish. Res. Board Can. 26:1743-1762. HALSTEAD, B. W. 1965. Poisonous and venomous marine animals of the world. I. In- vertebrates. U.S. Gov. Print. Off., Wash., D.C., 994 p. (see Phylum Coelenterata, p. 297-371). KRAMP, P. L. 1959. Stephanoscyphus (Scyphozoa). Galathea Rep. 1:173-185. 1961. Synopsis of the medusae of the world. J. Mar. Biol. Assoc. U.K. 40:1-469. MANSUETI, R. 1963. Symbiotic behavior between small fishes and jellyfishes, with new data on that between the stromateid, Peprilus alepido- tus, and the scyphomedusa, Chrysaora quinquecirrha. Copeia 1963:40-80. MAYER, A. G. 1910. Medusae of the world. III. The Scyphomedusae. Carnegie Inst. Wash. Publ. 109:499-735. PHILLIPS, P. J., and W. D. BURKE. 1970. The occurrence of sea wasps (Cubomedusae) in Mississippi Sound and the northern Gulf of Mexico. Bull. Mar. Sci. 20:853- 859. PHILLIPS, P. J., W. D. BURKE, and E. J. KEENER. 1969. Observations on the trophic significance of jellyfishes in Mis- sissippi Sound with quantitative data on the associative behavior of small fishes with medusae. Trans. Am. Fish. Soc. 98:703-712. PICKEN, L. E. R., and R. J. SKAER. 1966. A review of researches on nematocysts. In W. J. Rees (edi- tor), The Cnidaria and their evolution, p. 19-50. Zool. Soc. Lond. RUSSELL, F. S. 1970. The medusae of the British Isles. II. Pelagic Scyphozoa. Cambridge LIniv. Press, 284 p. SOUTHWARD, A. J. 1955. Observations on the ciliary currents of the jelly-fish Aurelia aurita L. J. Mar. Biol. Assoc. U.K. 34:201-216. SPANGENBERG, D. B. 1965. Cultivation of the life stages of Aurelia aurita under con- trolled conditions. J. Exp. Zool. 159:303-318. WEILL, R. 1934. Contribution a l'etude des cnidaires et de leurs nematocystes. Trav. Stn. Zoo. Wimereux 11:351-701. WERNER, B. 1973. New investigations on systematics and evolution of the class Scyphozoa and the phylum Cnidaria. Publ. Seto Mar. Biol. Lab. 20:35-60. 16 SYSTEMATIC INDEX Atolla vanhoeffeni 15 wyvillei 10, 15 Atollidae 15 Aurelia aurita 9, 13, 15 Carybdea alata 15 marsupialis 15 Carybdeidae 15 Chiropsalmus quadrumanus 15 Chrysaora quinquecirrha 9, 12, 15 Cleistocarpidae 15 Coronatae 10, 15 Craterolophus convovulus 8, 15 Cubomedusae 10, 15 Cyanea capillata 9, 12, 15 Cyaneidae 15 Eleutherocarpidae 14 Haliclystus auricula 7, 14 salpinx 7, 14 Linuche unguiculata 15 Lucernaria quadricornis 8, 14 Nausithoe punctata 15 Nauphantopsis diomedeae 15 Pelagia noctiluca 12 Pelagidae 11, 12, 15 Periphylla periphylla 11, 15 Periphyllidae 15 Phacellophora camtschatica 13, 15 Rhizostomatidae 16 Rhizostomeae 11, 16 Rhopilema verrilli 9, 14, 16 Semaeostomeae 11, 15 Stauromedusae 6, 14 Stomolophidae 16 Stomolophus meleagris 14, 16 Tamoya haplonema 10, 15 Thaumatoscyphus atlanticus 6, 15 Ulmaridae 11, 15 17 370. Collecting and processing data on fish eggs and larvae in the" California Current region. By David Kramer, Mary J. Kalin, Elizabeth G. Stevens, James R. Thrailkill, and James R. Zweifel. November 1972, iv + 38 p., 38 figs., 2 tables. For sale by the Superintendent of Documents, U.S. Govern- ment Printing Office, Washington, D.C. 20402. 371. Ocean fishery management: Discussions and research. By Adam A. Sokoloski (editor). (17 papers, 24 authors.) April 1973. vi + 173 p., 38 figs., 32 tables, 7 appendix tables. 372. Fishery publications, calendar year 1971: Lists and indexes. By Thomas A. Manar. October 1972, iv + 24 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 381. Fishery publications, calendar year 1967: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. July 1973, iv + 22 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 382. Fishery publications, calendar year 1966: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. July 1973, iv + 19 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washing ton, D.C. 20402. 383. Fishery publications, calendar year 1965: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. July 1973, iv + 12 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washing ton, D.C. 20402. 374. Marine flora and fauna of the northeastern United States. Annelida: Oligochaeta. By David G. Cook and Ralph O. Brinkhurst. May 1973, iii + 23 p., 82 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 384. Marine flora and fauna of the northeastern United States. Higher plants of the marine fringe. By Edwin T. Moul. September 1973, iii + 60 p., 109 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 375. New Polychaeta from Beaufort, with a key to all species recorded from North Carolina. By John H. Day. July 1973, xiii + 140 p., 18 figs., 1 table. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 385. Fishery publications, calendar year 1972: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. November 1973. iv + 23 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 376. Bottom-water temperatures on the continental shelf, Nova Scotia to New Jersey. By John B. Colton, Jr. and Ruth R. Stoddard. June 1973, iii + 55 p., 15 figs., 12 appendix tables. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. 386. Marine Flora and fauna of the northeastern United States. Pycnogo- nida. By Lawrence R. McCloskey. September 1973, iii + 12 p., 1 fig. For sale by the Superintendent of Documents. U.S. Government Printing Office. Washington, D.C. 20402. 377. Fishery publications, calendar year 1970: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. December 1972, iv + 34 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 387. Marine flora and fauna of the northeastern United States. Crustacea: Stomatopoda. By Raymond B. Manning. February 1974, iii + 6 p., 10 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 378. Marine flora and fauna of the northeastern United States. Protozoa: Ciliophora. By Arthur C. Borror. September 1973, iii + 62 p., 5 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 388. Proceedings of the first U.S. -Japan meeting on aquaculture at Tokyo, Japan, October 18-19, 1971. William N. Shaw (editor). (18 papers, 14 authors.) Februarv 1974, iii + 133 p. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 379. Fishery publications, calendar year 1969: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. April 1973, iv + 31 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washing- ton, D.C. 20402. 389. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda. By Austin B. Williams. April 1974. iii + 50 p., Ill figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 380. Fishery publications, calendar year 1968: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. May 1973. iv + 24 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washing- ton. D.C. 20402. 390. Fishery publications, calendar year 1973: Lists and indexes. By Mary Ellen Engett and Lee C. Thorson. September 1974. iv + 14 p., 1 fig. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 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