403 NOAA Technical Report NMFS Circular 403 mm Marine Flora and Fauna of the Northeastern United States Sipuncula Edward B. Cutler July 1977 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 the National Marine Fisheries Service (NMFS) 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 subsidies. It collects, analyzes, and publishes statistics on various phases of the industry. The NOAA Technical Report NMFS Circular 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 in- vestigations appear in the Circular series. NOAA Technical Report NMFS Circulars 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 D825, Technical Information Division, Environmental Science Information Center, NOAA, Washington, D.C. 20235. Re- cent Circulars are: 365. Processing EASTROPAC STD data and the construction of ver- tical temperature and salinity sections by computer. By Forrest R. Miller and Kenneth A. Bliss. February 1972, iv + 17 p., 8 figs., 3 app. figs. For sale bv the Superintendent of Documents, U.S. Government Printing Of- fice, Washington, D.C. 20402. 366. Key to field identification of anadromous juvenile salmonids in the Pacific Northwest. By Robert J. MacConnell 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 bv 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. Lindall. 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 Of- fice. Washington, D.C. 20402. 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. Government Printing Office, Washington, D.C. 20402. 371. Ocean fishery management: Discussion and research. By Adam A. Sokoloski (editor). (17 papers, 24 authors.) April 1973, vi + 17.3 p., 38 figs.. 32 tables, 7 app. tables. 372. Fishery publications, calendar year 1971: Lists and indexes. By Thomas A. Manar. October 1972. iv + 24 p., 1 fcg. For sale by the Superintendent of Documents, U.F. 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 bv 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. 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. Washington, D.C. 20402. 180. 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 Of- fice. 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 bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 382. Fisherv 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 Of- fice. Washington, D.C. 20402. 383. Fisherv publications, calendar year 1965: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. July 1973, iv + 12 p., 1 fig. For sale bv 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. Govern- ment Printing Office, Washington, 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. 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 bv the Superintendent of Documents, U.S. Government Printing Of- fice, Washington, D.C. 20402. 175. New Polvchaeta 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. 386. Marine flora and fauna of the northeastern United States. Pyc- nogonida. By Lawrence R. McCloskey. September 1973. iii + 12 p., 1 fig. For sale bv 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 app. tables. 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. Govern- ment Printing Office, Washington, D.C. 20402. Continued nn inside back cover NOAA Technical Report NMFS Circular 403 ,^ WMOSgft Marine Flora and Fauna of the Northeastern United States. Sipuncula Edward B. Cutler July 1977 I z ■J c D U.S. DEPARTMENT OF COMMERCE Juanita M. Kreps, 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. 003-012-00404-0 FOREWARD 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 ir- regular intervals on as many taxa of the region as there are specialists available to collaborate in their preparation. The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region," edited by R. I. Smith, published in 1964, and produced under the auspices of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. In- stead 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 publications. The "Marine Flora and Fauna of the Northeastern United States" is being prepared in collaboration with systematic specialists in the United States and abroad. Each manual will be based primarily on recent and ongoing revisionary systematic research and a fresh examination of the plants and animals. Each major taxon, treated in a separate manual, will include an in- troduction, illustrated glossary, uniform originally illustrated keys, annotated check list with in- formation when available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a systematic index. These manuals are intended for use by biology students, biologists, biological oceanographers, informed laymen, and others wishing to identify coastal organisms for this region. In many instances the manuals will serve as a guide to additional information about the species or the group. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is planned to include organisms from the headwaters of estuaries seaward to approximately the 200-m depth on the continental shelf from Maine to Virginia, but may vary somewhat with each major taxon and the interests of collaborators. Whenever possible representative specimens dealt with in the manuals will be deposited in reference collections of the major museums in the region. After a sufficient number of manuals of related taxonomic groups have been published, the manuals will be revised, grouped, and issued as special volumes. These volumes will thus con- sist of compilations of individual manuals within phyla such as the Coelenterata, Arthropoda, and Mollusca, or of groups of phyla. CONTENTS Page Introduction 1 Key to the Sipuncula, Maine to Virginia 3 Annotated systematic list of species 5 Selected bibliography - 5 Index to scientific names 6 Acknowledgments 7 Coordinating editor's comments 7 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. Marine Flora and Fauna of the Northeastern United States. Sipuncula EDWARD B. CUTLER 1 ABSTRACT This report includes an account of the five species of Sipuncula living in shallow waters (down to 200 m) from Maine to Virginia. Four of these are widespread elsewhere in the world— Go Ifingia eremita, G. margaritacea, G. minuta, and Phascolion atrombi. Phxwcolopsis gouldi is endemic to the east coast of North America. An introduction to their biology, an annotated systematic list, selected bibliography, and an illustrated key are presented. INTRODUCTION The Sipuncula (peanut worms) are marine, seden- tary, vermiform coelomates which lack any trace of seg- mentation, although related to the annelids. They are found in all oceans and live within the sediment or in- side a protective shelter such as a discarded mollusc shell (pteropod, gastropod, or scaphopod), foraminiferan test, polychaete tube, or crevice in rock or coral. Sipunculans have two body regions: the trunk (or body) and a retractable introvert (Fig. 1A). The mouth is located at the tip of the introvert and is usually sur- rounded by tentacles of varying degrees of development and complexity. Behind the tentacular region is a zone which may bear chitinous, posteriorly directed hooks which are scattered or arranged in regular rings. The in- trovert may be less than half the trunk length in some species to many times its length in others. The demarca- tion between these two regions is usually defined by the abrupt narrowing of the diameter where the introvert begins. This area generally coincides with the position of the anus or nephridiopores, or both. Sipunculans range in trunk length from less than 2 mm to over 300 mm, but in the northeastern Atlantic they are more commonly between 3 and 30 mm. The shape of the body may vary from almost spherical to cylindrical. In those forms inhabiting discarded gas- tropod shells, the living worm retains the spiral shape of its shell even after it is removed. Sipunculans have a variety of epidermal structures such as papillae, hold- fasts, and hooks. The papillae are glandular structures which produce material which is thought may function in constructing or maintaining burrows or tubes. Generally, pigmentation is scant but when present, it is in shades of yellow or brown and is not ordinarily useful for iden- tification. The internal anatomy of these worms is relatively sim- ple (Fig. IB). The digestive tract has a straight esoph- agus and a double-coiled intestine extending towards 'Utica College of Syracuse University, Utica, NY 13502. the posterior end of the body. The tract terminates in a rectum, which may bear a small caecum. The dorsal anus is located at the anterior end of the trunk except in a few species where it is in the midregion and in one genus (Onchnesoma) where it is on the introvert. The distal part of the rectum is anchored to the body wall by a sheet of muscle tissue referred to as the wing muscles. Near the anus a long strand of muscle tissue originates from the body wall and extends down the center of the gut coil, increasing its stability; this is the spindle mus- cle. It may terminate within the coil or extend beyond to the posterior end of the trunk. In addition, the worm may have fixing muscles, which are fine muscle strands anchoring the gut coils or esophagus to the body wall. One or usually two simple, saclike nephridia (sometimes called segmental organs) open on the ven- tral side at the anterior end of the trunk. The nephridia are always attached to the body wall at the neph- ridiopores and sometimes are attached by more exten- sive mesentery along some portion of their length. The nephridiopores themselves, in most species, are not dis- cernible externally. The longitudinal and circular muscles of the body wall are frequently arranged in smooth, uniform layers but also may be gathered into bundles. One, two, or four retractor muscles control the introvert. A ventral nerve cord with lateral nerves and a pair of cerebral ganglia are present. Two or four pig- mented eyespots may be present on the cerebral ganglia. Our knowledge of the reproductive biology of east coast sipunculans is scanty. Rice (1967, 1973, 1975) has given the most current statement of our knowledge on develop- mental patterns in Sipuncula. Gerould (1907) gave an ac- count of the development of Phascolopsis gouldi. Good information on breeding cycles is unavailable for most species. Gametes are produced from a strip of gonadal tissue at the base of the ventral retractor muscles. They are released into the coelom at an early stage where they undergo the remainder of their growth and differentia- tion as freely floating cells. Gibbs (1975), working with Golfingia minuta pop- ulations, found monoecious animals in the eastern At- lantic but not in the western Atlantic. With the possible Mouth v- Tentacles Hooks ntrovert svn Trunk •M '°^¥- Anus Esophagus Ventral Nerve Cord Nephridium Gut Coil Right Dorsal Retractor — Rectum RightVentral Retractor Figure 1. — General morphology of Sipuncula. A. External form of Phascolion strombi. B. Internal form of Phascolopsis gouldi. 5 v.i; Holdfasts exception of G. minuta, sipunculans are dioecious and lack any sexual dimorphism. The gametes are retained in the coelom until mature then released through the neph- ridiopores, fertilization occurring externally. Free-swim- ming trochophore larvae generally occur but some brooding of early stages within the maternal "home" (crevice or hole in substrate) has been observed in G. minuta and may occur in other cold- or deepwater forms. After metamorphosis, the juveniles settle onto a suitable substratum and probably remain in that immediate vicinity throughout their lives. Our knowledge of growth rate or longevity is nonexistent. Sipunculans are either filter or deposit feeders, depend- ing on the species. The filter feeders have a more elabor- ate tentacular apparatus than the deposit feeders. As such, these worms play their part in the recycling of detri- tus and probably consume smaller invertebrates in the process. They are in turn preyed on by fish and probably other predators. It has been reported in the literature that sipunculans are used as food for man in Indonesia and the Indo-Pacific (Hyman 1959) and are also used as fish bait in these regions. The methods for collecting Sipuncula vary according to the area. Phascolopsis gouldi can be collected with a shovel at low tide; in warmer waters intertidal forms often inhabit coral or rock which must be carefully broken with a hammer. To collect the deeper dwelling species a dredge or trawl is required. These should be equipped with a fine mesh bag (less than 2 mm) to retain the smaller individuals. The sediment is then gently washed through a sieve (1 mm or less) and the remaining material inspected with a dissecting microscope. Initial fixation should be with 4% formaldehyde but, if pos- sible, should be preceded by a 30-60 min period of nar- cotization with a compound such as menthol crystals or a dilute solution (5-10%) of ethanol and seawater, to allow the introverts to protrude. This greatly facilitates iden- tification. For long-term storage 70% ethanol should be used. In order to identify most animals to species, dissection is required. The recommended procedure is to find the dorsal anus and make an incision down the dorsal side passing just to the left of the anus, thereby avoiding the internal organs. Hyman (1959) gave a general account of the biology of the sipunculans. Gerould (1913) and Cutler (1973) have discussed the western North Atlantic fauna in more detail. Stephen (1960) has brief descriptions of all these species except Phascolopsis gouldi. Stephen and Ed- monds (1972) presented the most comprehensive treat- ment of the phylum, but it is not without error. Theel (1905) contains good illustrations and descriptions of all the species discussed here except for P. gouldi. Stephen (1964) first proposed that the proper spelling of this phy- lum is Sipuncula, familiarly known as sipunculans. This usage has been accepted by specialists but has been slow to spread throughout the biological community. The five species found in the area between Maine and Virginia to a depth of 200 m are all common forms, four of them being found in all major oceans. One, P. gouldi, is endemic to the Atlantic coast of the United States and is rare south of Long Island. Onchnesoma steenstrupi and Golfingia catharinae are common on the continen- tal slope but occasionally occur on the outer shelf southeast of Long Island. Because they are restricted to the outer shelf and occur there only infrequently, they have not been included here (see Cutler 1973). KEY TO THE SIPUNCULA, MAINE TO VIRGINIA This is an artificial key designed only for the identification of the five inshore species found in the area covered by this manual. Accurate identification requires a well-preserved, entire organism with the internal parts intact. If the in- trovert is not fully extended, the nature of the tentacles can usually be determined by dissection of the retracted in- trovert. Measurements given are trunk lengths of average adults. 1 Trunk surface with chitinized epidermal structures (holdfasts) as in Figures 1A and 2A; one nephridium; irregular gut coils (Fig. 2B IGC); usually living in mollusc shell; 3-15 mm . . Phascolion strombi IGC 1 Trunk surface without chitinized epidermal structures; two nephridia; regular double helix of gut coils (Fig. IB); usually living in sediment 2 2 (7) Longitudinal body musculature separated into distinct bundles (Fig. 3A); common in shallow subtidal waters; up to 130 mm (Fig. 3B) Phascolopsis gouldi 3A 2 (1) Longitudinal body musculature forms continuous sheet; rare in water less than 10 m deep 3 3 (2) Introvert with two pair of retractor muscles (dorsal and ventral, Fig. IB); 5-25 mm (Fig. 4) ... Golfingia margaritacea 3 (2) Introvert with one pair of retractor muscles (ventral only) 4 4 (3) Trunk smooth; well-developed tentacles as in G. margaritacea; mature forms are stout with thick body walls; free-living; 5-25 mm (Fig. 5) Golfingia eremita 4 (3) Posterior of trunk may have papillae; tentacles reduced to lobes (Fig. 6A); small (2-10 mm) and slender with thin body walls (Fig. 6B); sometimes found in hyalinaceous polychaete or arena- ceous foraminiferan tubes Golfingia minuta 6A ANNOTATED SYSTEMATIC LIST OF SPECIES Phylum Sipuncula This taxon has been considered to be at a variety of systematic levels and has been spelled many ways at various times in the past. Since its elevation to the rank of phylum there has been a void of intermediate taxa. The description of families occurred recently (Stephen and Edmonds 1972), but there are still no orders or classes. Golfingia (Phascoloides) minuta (Keferstein 1865) From Maine to North Carolina, also in the eastern North Atlantic to lat. 70°N, Mediterranean, off South Africa and Chile; 1-3,750 m, uncommon on the shelf; medium sand-silt sediment; temperature range 0-10°C. This is a poorly understood species and opinions vary as to its nature. Gerould's (1913) G. cinctum is definitely a synonym and G. improvisa (Theel) may be a synonym but this question is unresolved (see Cutler 1973). A dis- cussion of the history of Golfingia minuta and G. im- provisa can be found in Murina (1958). Family SIPUNCULIDAE Phascolopsis gouldi (Pourtales 1851) Endemic to the western North Atlantic, common in southern New England, scattered reports south of lat. 30°N; no sexually mature forms known south of Cape Hatteras; depth 0-165 m, most common in shallow water; sediment gravelly sand to silty sand; temperature range 0-25°C. During the past 100 yr this species has been assigned to the genera Sipunculus, Phascolosoma, and Golfingia. The most recent shift (Stephen 1964) elevated Phas- colopsis from subgeneric to generic rank. This is now a monotypic genus. I believe this is a mistake and that it is more closely related to Siphonosoma; until we better un- derstand the development of Siphonosoma I have not suggested this change (see Cutler 1973). It is unfor- tunate that a species so commonly used by experimental biologists has undergone so many generic changes. An- drews (1890) presents a detailed account of the anatomy of this species. Family GOLFINGIIDAE Golfingia {Golfingia) margaritacea (Sars 1851) A widely distributed species found in all oceans but most common in the North Atlantic and absent from the tropics; along the coast from Maine to Cape Hatteras, on the outer shelf and upper slope; sandy sediments; tem- perature range 2-12°C. Several dubious subspecies, forms, and varieties have been described in the literature for this widespread form. Golfingia (Phascoloides) eremita (Sars 1851) Common on both sides of the North Atlantic up into the Arctic Ocean, one record from 500 m off California; on the east coast from Maine to Cape Hatteras, rare south of Long Island; outer shelf and slope; sandy-silt sediment; temperature range 0-10°C. Phascolion strombi (Montague 1804) Common on the shelf from Maine to Long Island (rare on Georges Bank), ranging in deeper water to North Carolina (lat. 32°N); common throughout the Atlantic from the Arctic to the Antarctic, one record from the Pacific off southern Chile; depth 14-3,800 m; tem- perature range 2-26°C. This is the most common, eury- thermal, eurybathyl, and morphologically plastic species in the area (Gerould 1913, Cutler 1973). SELECTED BIBLIOGRAPHY ANDREWS, E. A. 1890. Notes on the anatomy of Sipunculus gouldii Pourtales. Stud. Biol. Lab. Johns Hopkins Univ. 4:384-430. CUTLER, E. B. 1973. Sipuncula of the western North Atlantic. Bull. Am. Mus. Nat. Hist. 152:105-204. GEROULD, J, H. 1907. The development of Phascolosoma. Zool. Jahrb., Abt. Anat. Ontogen. Tiere 23:77-162. 1913. The sipunculids of the eastern coast of North America. Proc. U.S. Natl. Mus. 44:373-437. GIBBS, P. E. 1975. Gametogenesis and spawning in a hermaphroditic population of Golfingia minuta (Sipuncula). J. Mar. Biol. Assoc. U. K. 55:69-82. HYMAN. L. H. 1959. The protostomatous coelomates — phylum Sipunculida. In The invertebrates, Vol. 5, p. 610-696. McGraw-Hill Book Co., N.Y. MURINA, V. V. 1958. On the systematics of two closely related species of the deep- water sipunculids of the genus Golfingia according to the materials of the "Vitiaz" expeditions in 1949-1955. [In Russ.] Zool. Zh. 37:1624-1633. RICE, M. E. 1967. A comparative study of the development of Phascolosoma agassizii, Golfingia pugettensis, and Themiste pyroides with a dis- cussion of developmental patterns in the Sipuncula. Ophelia 4: 143-171. 1973. Morphology, behavior, and histogenesis of the pelagosphera larva of Phascolosoma agassizii (Sipuncula). Smithson. Contrib. Zool. 132, 51 p. 1975. Observations on the development of six species of Caribbean Sipuncula with a review of development in the phylum. In M. E. Rice and M. Todorovic (editors), Proc. Int. Symp. Biol. Sipuncula and Echiura. Vol. 1. p. 141-160. Naunco Delo Press, Belgrade. STEPHEN, A. C. 1960. British echiurids (Echiuroidea), sipunculids (Sipunculoidea), and priapulids (Priapuloidea). Synopses Br. Fauna 12:1-27. 1964. A revision of the classification of the phylum Sipuncula. Ann. Mag. Nat. Hist., Ser. 13, 7:457-462. STEPHEN. A. C, and S. J. EDMONDS. 1972. The phyla Sipuncula and Echiura. Trustees Br. Mus. (Nat. Hist.). Lond., 528 p. THEEL. H. 1905. Northern and Arctic invertebrates in the collection of the Swedish State Museum I. Sipunculids. K. Sven. Vetenskapakad. Handl. 39:1-130. INDEX TO SCIENTIFIC NAMES Golfingia eremita 4,5 margaritacea 4, 5 minuta 1, 2, 4, 5 Golfingiidae 5 Phascolion strombi 3, 5 Phascolopsis gouldi 1, 2, 3, 5 Sipunculidae 5 ACKNOWLEDGMENTS Preparation of the "Marine Flora and Fauna of the Northeastern United States" is being coordinated by the fol- lowing board: Coordinating Editor: Melbourne R. Carriker, College of Ma- rine Studies, University of Delaware, Lewes, DE 19958. Roland L. Wigley, National Marine Fish- eries Service, Biological Laboratory, Woods Hole, Mass. Robert T. Wilce, Department of Botany, University of Massachusetts, Amherst, Mass. Editorial Advisers: Marie B. Abbott, Marine Biological Lab- oratory, Woods Hole, Mass. Arthur G. Humes, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Mass. Wesley N. Tiffney, Department of Biol- ogy, Boston University, Boston, Mass. Ruth D. Turner, Museum of Compara- tive Zoology, Harvard University, Cambridge, Mass. The Board established the format for the "Marine Flora and Fauna of the Northeastern United States," invites systematists to collaborate in the preparation of manuals, review manuscripts, and advises the Scientific Editor of the National Marine Fisheries Service. The completion of this paper was made possible by the assis- tance of Norma J. Cutler. The artwork was done by Mary Ann Pritchard and Henry Iwanicki. COORDINATING EDITOR'S COMMENTS Publication of the "Marine Flora and Fauna of the Northeastern United States" is most timely in view of the grow- ing universal emphasis on environmental work and the urgent need for more precise and complete identification of coastal or- ganisms than has been available. It is mandatory, wherever pos- sible, that organisms be identified accurately to species. Ac- curate scientific names 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. Edward B. Cutler began his studies of Sipuncula in 1964 as a participant in the International Indian Ocean Expedition. As a graduate student at the University of Rhode Island he worked with several large collections from eastern North American and Bermudan waters. That work was published in 1973. His on- going research is related to the systematics and zoogeography of the Indian Ocean and the sub-littoral Atlantic Ocean Sipun- cula. Particular attention has been paid to the slope fauna off the Carolinas. He has been at Utica College of Syracuse Univer- sity since 1967. Preparation of this manual was supported in part by a grant from the Environmental Protection Agency to the Editorial Board of the "Marine Flora and Fauna of the Northeastern United States." Work on the "Marine Flora and Fauna of the Northeastern United States" by the Coordinating Editor is sup- ported by the College of Marine Studies, University of Delaware. Manuals are available for purchase from the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. The manuals so far published in the series are listed below. COOK, DAVID G., and RALPH 0. BRINKHURST. Marine flora and fauna of the Northeastern United States. Annelida: Oligochaeta. BORROR, ARTHUR C. Marine flora and fauna of the Northeastern United States. Protozoa: Ciliophora. MOUL, EDWIN T. Marine flora and fauna of the Northeastern United States. Higher plants of the marine fringe. McCLOSKEY, LAWRENCE R. Marine flora and fauna of the Northeastern United States. Pycnogonida. MANNING, RAYMOND B. Marine flora and fauna of the Northeastern United States. Crustacea: Stomatopoda. WILLIAMS, AUSTIN B. Marine flora and fauna of the Northeastern United States. Crustacea: Decapoda. POLLOCK, LELAND W. Marine flora and fauna of the Northeastern United States. Tardigrada. LARSON, RONALD J. Marine flora and fauna of the Northeastern United States. Cnidaria: Scyphozoa. CAVALIERE, A. R. Marine flora and fauna of the Northeastern United States. Higher fungi: Ascomycetes, Deuteromycetes, and Basidiomycetes. COULL, BRUCE C. Marine flora and fauna of the Northeastern United States. Copepoda: Harpacticoida. CUTLER, EDWARD B. Marine flora and fauna of the Northeastern United States. Sipuncula. 6 US. GOVERNMENT PRINTING OFFICE: IQ77-797-854' 22 REGION 10 388. Proceedings of the first U.S. -Japan meeting on aquaculture at Tokyo, Japan, October 18-19, 1971. William N. Shaw (editor). (18 papers, 14 authors.) February 1974, iii + 133 p. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 389. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda. By Austin B. Williams. April 1974, iii + 50 p., Ill figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, 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. 391. Calanoid copepods of the genera Spinocalanus and Mimoealanus from the central Arctic Ocean, with a review of the Spinocalanidae. By David M. Damkaer. June 1975, x + 88 p., 225 figs., 4 tables. For sale bv the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 392. Fishery publications, calendar year 1974: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. June 1975, iv + 27 p., 1 fig. 393. Cooperative Gulf of Mexico estuarine inventory and study — Texas: Area description. By Richard A. Diener. September 1975, vi + 129 p., 55 figs., 26 tables. 394. Marine Flora and Fauna of the Northeastern United States. Tar- digrada. Bv Leland W. Pollock. May 1976, iii + 25 p., figs. For sale b\ i he Superintendent of Documents, U.S. Government Printing Office. Washington. D.C. 2(1402. 395. Report of a colloquium on larval fish mortality studies and their relation to fishery research. January 1975. By John R. Hunter. May 1976, iii + 5 p. For sale by the Superintendent of Documents, U.S. Government Printing Office. Washington, D.C. 20402. UNITED STATES DEPARTMENT OF COMMERCE NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION NATIONAL MARINE FISHERIES SERVICE SCIENTIFIC PUBLICATIONS STAFF ROOM 450 1107 N E 45TH ST SEATTLE. 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