C53./3--A//'/S< 405 / ^ 0Fc°<, Mm c <5> & ^rEs o* * NOAA Technical Report NMFS Circular 405 Marine Flora and Fauna of the Northeastern United States. Echinodermata: Holothuroidea David L. Pawson September 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. 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. 37M. 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. 367. Engineering economic model for fish protein concentration processes. Bv 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 bv 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 bv 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 + 173 p., 38 figs., 32 tables, 7 app. tables. 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 bv 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 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 bv 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. 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. 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 bv 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. 375. 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 bv 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. Bv 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 on inside back cover ^D WMOS^ ^"^To^P- NOAA Technical Report Circular 405 Marine Flora and Fauna of the Northeastern United States. Echinodermata: Holothuroidea David L. Pawson September 1977 Q. O c c. o Q U.S. DEPARTMENT OF COMMERCE Juanita M. Kreps, Secretary National Oceanic and Atmospheric Administration Richard A. Frank, 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. 003-017-00+0-4 FOREWORD This issue of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna of the Northeastern United States." This subseries will consist of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants and animals of the northeastern United States. Manuals will be published at irregular intervals on as many taxa of the region as there are specialists available to collaborate in their preparation. The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region," edited by R. I. Smith, published in 1964, and produced under the auspices of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. Instead of revising the "Woods Hole Keys," the staff of the Systematics-Ecology Program decided to ex- pand the geographic coverage and bathymetric range and produce the keys in an entirely new set of expanded publications. The "Marine Flora and Fauna of the Northeastern United States" is being prepared in collaboration with systematic specialists in the United States and abroad. Each manual will be based primarily on recent and ongoing revisionary systematic research and a fresh examination of the plants and animals. Each major taxon. treated in a separate manual, will include an in- troduction, illustrated glossary, uniform originally illustrated keys, annotated check list with in- formation when available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a systematic index. These manuals are intended for use by biology students, biologists, biological 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 the reference collections of major museums in the region. After a sufficient number of manuals of related taxonomic groups have been published, the manuals will be revised, grouped, and issued as special volumes. These volumes will thus con- sist of compilations of individual manuals within phyla such as the Cnidaria, Arthropoda, and Mollusca, or of groups of phyla. CONTENTS Page Introduction 1 Morphology 1 Systematic characters 2 Examination procedures 2 Glossary 2 Key to the holothurians of the northeastern United States inshore and shelf waters 4 Annotated systematic list 12 Selected bibliography 13 Systematic index 14 Acknowledgments 15 Coordinating editor's comments 15 The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. in Marine Flora and Fauna of the Northeastern United States. Echinodermata: Holothuroidea DAVID L. PAWSON1 ABSTRACT The holothurian fauna of the east coast of the United States from southern New Jersey to Nova Scotia, to depths of 200 m, comprises 21 species in 17 genera and 8 families. An introduction to the general biology and morphology of the holothurians, systematic characters and examination procedures, and a selected bibliography are included, together with an illustrated key to the species, an annotated systematic list, and an index. Affinities of this fauna are chiefly boreal; some species also occur south of Cape Hatteras, while others are essentially Arctic in distribution. INTRODUCTION Holothurians or sea cucumbers are usually incon- spicuous, soft-bodied animals, often unattractive in ap- pearance, which at first glance seem to belie their close relationship with the more aesthetically pleasing sea stars and sea urchins. But closer examination reveals the five-part (pentamerous) symmetry that is typical of liv- ing echinoderms, although in the holothurians the pen- tamery is often obscured by a more or less conspicuous bilateral symmetry. A skeleton of calcium carbonate in the form of calcite is present, though the elements are usually reduced to microscopic ossicles (see Hyman 1955 and Boolootian 1966). In the tropics, large aspidochirotid species 30 cm or more in length are very common, and the holothurians are a conspicuous element in shallow-water faunas, but in the temperate waters of the northeastern United States most are secretive, living buried in sand or mud, or concealed under rocks. A single relatively large species, Cucumaria frondosa (Gunnerus), is common on rocks along the coast of Maine, just below lowtide level (Jordan 1972). Some species are, however, important members of offshore benthic communities, and can reach population densities of many hundreds of individuals per square meter. They are efficient reworkers of seafloor sediments (for example, see Rhoads and Young 1971), and in many areas of the deep sea can comprise more than 90"? of the total biomass present (Zenkevitch 1963). With few exceptions, holothurians are sedentary; some, such as the psolids, may remain fixed to the same spot for years at a time. Dispersal is effected by trans- port of larval stages in currents to other localities. Some have planktotrophic larvae; others have yolky ciliated vitellaria larvae. Commonly, populations of a species may die off in an area and remain absent for a con- siderable number of years, until the area is repopulated by a new settlement of larvae. 'Department of Invertebrate Zoology, Smithsonian Institution, Wash- ington, D.C. 20560. Affinities of the northeastern United States holothurians are chiefly boreal; several of the species also occur around Europe and in the Arctic. Some species ex- tend southward below Cape Hatteras, even into the Caribbean. Others are known only from the northeastern United States and Canadian coasts (Clark 1901; Coe 1912). The fauna lacks representatives of the Order Aspidochirotida. The large Parastichopus californicus (Stimpson), so typical of the northwestern United States' shallow waters, has no counterpart on the east coast. The Order Dendrochirotida is well represented and at least one species, Sclerodactyla briareus (Lesueur), is a commonly used experimental animal (Farmanfarmaian 1969, Ellington 1976). The 21 species dealt with here occur in depths of 200 m or less off the northeastern United States; a small number of forms of doubtful status were omitted. Only those species recorded from the area bordered to the south by southern New Jersey and to the north by Nova Scotia, Canada, were included. Literature on holothurians of the northwest Atlantic is scanty. Very little information is available on ecology, physiology, and other aspects of the biology of most species. The best systematic study of the fauna is that of Deichmann (1930), which remains the standard work, al- though some of the nomenclature is now outdated. The key provided here includes illustrations of impor- tant taxonomic characters. Most species are illustrated, and where desirable in the key, illustrations of ossicles or other anatomical features are included. MORPHOLOGY Figures 1 and 2 show basic features of the external and internal morphology of a holothurian. Holothurians generally tend to be cylindrical, with mouth and anus at opposite ends of the body. The mouth is surrounded by 10 or more tentacles. Typically there are five rows of suc- torial tube feet, which form radii, the radii running from the anterior end of the body to the posterior end. Three ALOTMO BAD Figure 1. -External features of a holothurian, viewed from the ven- tral side. radii are ventral and two are dorsal; spaces between the radii are termed interradii. The dermal layer of the body wall contains numerous microscopic calcareous ossicles (often termed calcareous deposits or spicules), which assume a great variety of shapes and which are very useful in classification. The extensive coelomic cavity is lined with circular muscles, and five conspicuous longitudinal muscles define the radii. The mouth opens into an esophagus, which leads to a stomach (often poorly differentiated) and intestine. The intestine describes a large loop, then runs to the terminal anus via the rectum. Two branching respiratory trees may arise from the rectum and extend anteriorly in the coelom; these are used in respiration and excretion of waste materials. A ring of calcareous plates, the calcareous ring, surrounds the esophagus, and this serves as the point of attachment for the retractor muscles, which serve to withdraw the anterior end of the body (the introvert) into the coelomic cavity when the animal contracts. A part of the coelom forms the water vascular system, which is a system of fluid-filled vessels used in feeding, locomotion, and respiration. A ring vessel immediately posterior to the calcareous ring carries a stone canal in the middorsal interradius, the stone canal terminating in an internal madreporite. A polian vesicle arises from the ventral side of the ring vessel and hangs into the coelom. From the ring vessel five radial water vessels run an- teriorly, giving rise to vessels serving the tentacles, and then posteriorly in the radii, giving rise to the tube feet, which penetrate the body wall to the exterior. The sexes are separate. The gonad comprises one or two bunches of caeca hanging free in the body cavity. A genital duct arises from the confluence of the caeca and runs anteriorly in the dorsal mesentery to open to the ex- terior in the middorsal interradius near the tentacles. TENTACLE CALCAREOUS RING POLIAN VESICLE RETRACTOR MUSCLE INTESTINE RESPIRATORY TREE RAOIAL LONGITUDINAL MUSCLE MADREPORITE MADREPORIC DUCT RING VESSEL GENITAL DUCT GENITAL TUBULE Figure 2.— Internal features of a cucumariid holothurian, dissected from the left side. 2 SYSTEMATIC CHARACTERS External Characters Shape is highly variable. Some species are U-shaped, others cylindrical or even wormlike. A conspicuous tail may be present. The tube feet may be restricted to the radii in well defined rows, or they may be scattered all over the radii and interradii in various characteristic pat- terns. The Orders Apodida and Molpadiida lack tube feet. Tentacles may be dendritic, digitate, or pinnate. There are usually 10 tentacles, but some groups have 15- 30, and many apodous species have 12 or 13. Internal Characters The calcareous ring is of particular importance (Paw- son and Fell 1965). The ring may have well developed posterior projections, moderately developed projections, or none at all. The ring itself is typically composed of 10 pieces, but many apodous species have 12, and in some forms the ring is composed of a mosaic of numerous pieces. Other features of the water vascular system and musculature are useful in classification, but are not rele- vant to the species being considered here. Ossicles These are found in the body wall, tentacles, and tube feet, and they display a bewildering variety of shapes and sizes. They are of paramount importance in classifica- tion, particularly at the level of the genus and species. Certain types of ossicles have convenient names, such as buttons, plates, tables, wheels, anchors (see Fig. 3), and the various major groups of holothurians have charac- teristic types of ossicles. In the Order Dendrochirotida smooth or knobbed plates are common; small plates are often termed buttons. Baskets or cups occur frequently, and some taxa have tables, which consist of a flat per- forated disk surmounted by a spire composed of two or more columns joined at their distal extremity. Rods oc- cur in the tube feet and tentacles. Members of the Order Figure 3. — Some typical holothurian ossicles: A. knobbed button; B. shallow cup; C. plate; D. rod; E. smooth button; F. wheel; G. anchor plate; H. anchor. A-E. Dendrochirotida; F. Apodida, Chiridotidae; G, H. Apodida, Synaptidae. Apodida may have either anchors and anchor plates (Family Synaptidae) or wheels (Families Chiridotidae and Myriotrochidae) (see Clark 1924). Rods of various types may also be present. In the Order Molpadiida the ossicles can take the form of tables, and sometimes characteristic anchors and plates are present. In many molpadiids the ossicles are transformed into wine-red "phosphatic deposits" as the animal grows; these give the body wall a distinctive light to dark red color. EXAMINATION PROCEDURES Holothurians generally contract when preserved, with- drawing their tentacles, and changing their shape dramatically. They can be effectively narcotized with Epsom salts (MgS04) in seawater before fixation. Specimens should be preserved in 70% ethyl alcohol; acid conditions in formaldehyde tend to dissolve the os- sicles with passage of time. Dissection is best effected by making a longitudinal in- cision along the left or right side of the body. Most organs are easily seen when the body cavity is opened in this way. Ossicles are usually smaller than 1 mm in diameter (many are 50 /i m or less) and must be examined with a compound microscope. They can be extracted by im- mersion of a small piece of the body wall in household bleach solution, or in sodium hypochlorite. The soft tis- sues are dissolved away, leaving the ossicles. Usually, body wall samples from the middorsal area will reveal typical ossicles. Permanent microscope slides can be made by washing the ossicles several times in distilled water, and mounting them in Canada balsam or some other permanent medium. GLOSSARY Aspidochirotida An Order of Holothuroidea which in- cludes forms with shield-shaped tentacles and a thick body wall. Comprises mostly tropical and deep-sea species. calcareous ring A ring of solid plates of calcite surround- ing and supporting the esophagus and related struc- tures. dendritic Branching profusely. Dendrochirotida An Order of Holothuroidea which in- cludes forms with richly branching tentacles. digitate Possessing fingerlike projections. genital duct A tube which links the gonad to the exter- ior and carries eggs or sperm into the surrounding sea- water. introvert In Dendrochirotida, the anterior part of the body, comprising tentacles, calcareous ring and related structures, which can be retracted by contraction of retractor muscles. madreporite A sievelike plate through which water may be drawn into the water-vascular system. Internal in most shallow-water holothurians, external in many deep-sea forms. ossicles Usually small, distinctively shaped, skeletal elements composed or calcium carbonate in form of calcite. pentamerous Having five-part symmetry. pinnate Featherlike. Used for tentacles with numerous digits. planktotrophic Able to feed on planktonic organisms. polian vesicle In water-vascular system, a contractile sac in which excess fluid may be stored. psolids Dendrochirotid holothurians in which the upper surface of the body is covered with large overlapping ossicles, and the lower surface is in the form of a soft sole. retractor muscles In dendrochirotids, set of five mus- cles used to retract the introvert. ring vessel In water-vascular system, a vessel surround- ing esophagus, which gives rise to polian vesicle, stone canal, and radial vessels. sole In psolids, the soft ventral surface of the body. stone canal In water-vascular system, a tube, often cal- cified, which links the madreporite to the ring vessel. tentacles Dendritic, digitate, or shield-shaped anterior extensions of the water-vascular system which sur- round the mouth and are used in feeding. tube feet Hollow extensions of water-vascular system which project through body wall. Locomotory organs; display great variety in number and arrangement. vitellaria A type of larval stage richly supplied with yolk, upon which the larva feeds. water-vascular system A system of fluid-filled vessels used in respiration, locomotion, nutrition, and sensory perception. On external surface of body manifested as tube feet and tentacles. KEY TO HOLOTHURIANS OF THE NORTHEASTERN UNITED STATES INSHORE AND SHELF WATERS Body with typical tube feet, which may be scattered or restricted to well-defined areas (radii). Ten or 20 tentacles, richly branched (dendritic). Body wall generally firm, leathery Body totally lacking tube feet. Twelve or 15 tentacles, with fingerlike branches (digitate or pin- nate). Body wall leathery and opaque or fragile and translucent .15 2(1) Body generally cylindrical, often tapering anteriorly and posteriorly, never partly enclosed in a test of conspicuously overlapping scales 2(1) Body with flattened ventral surface; dor- sal surface arched, covered by a test of overlapping scales. Ventral surface (sole) soft, with tube feet around edge and some- times along midventral radius 14 3 (2) Calcareous ring with no posterior projections; posterior margin of ring undulating. Tentacles 10 or 20. Tube feet scattered over body wall or tending to be confined to radii 3 (2) Calcareous ring with long (a) or short (b) poster- ior projections. Tentacles 10. Tube feet confined to radii or scattered over body wall 11 4 (3) Tentacles 20, 10 large and 10 small, arranged in 2 rings. Ossicles tables, which tend to disappear with age 5 4(3) Tentacles 10, of which 2 ventral ones are smaller than others. Notables 6 5 (4) Tube feet tend to be scattered uniformly over body. Ossicles robust tables with low spire ending in numerous teeth. Body wall thin, translucent. Maximum size 15 cm, color whitish or brownish Thyonidium pellucidum ^nfyniVirt^ ^S 5 (4) Tube feet tend to be aggregated in radii and scattered in interradii. Ossicles delicate tables with high spire ending in few teeth. Body wall thick, not trans- lucent. Maximum size 10-15 cm, color whitish or yellowish Duasmodactyla commune 6 (4) Tube feet uniformly scattered over radii and interradii. Calcareous ring with undulating posterior edge or with rudimentary projections. Ossicles but- tons (a) and cups (b). Maximum size 7 cm, color whitish with brown spots, or uniform pale brown Thyonella pervicax 1 6 {4) Tube feet tend to be aggregated along radii, especially ventrally, but may also be present, scat- tered, in interradii (6) Ossicles include numerous buttons and numerous cups, the latter rudimentary or with perfora- tions 7(6) Ossicles may include plates or buttons. No cups 10 8 (7) Feet absent from interradii. Ossicles knobbed buttons (a) and rudimen- tary cups (b) lacking perforations. Maximum size 2 cm, color light orange to white Ocnus diomedeae 8 (7) Feet present in interradii, at least dorsally. Ossicles smooth or knobbed buttons and perforated flat cups 9(8) 9(8) Feet numerous, scattered in all in- terradii. Ossicles knobbed buttons (a) and cups (b). Maximum size 15 cm, color mottled brown Thyonella gemmata Feet very few, scattered in dorsal interradii only. Ossicles smooth buttons (a) and cups (b). Maximum size 1.5 cm, color white Trachvthvone nina 10 (7) Ossicles plates, which disappear as the animal grows. Tube feet generally restricted to radii, but also scattered dorsally. Maxi- mum size 50 cm, color brown . . . Cucumaria frondosa 10 (7) Ossicles four-holed buttons with smooth to knobbed surface. Tube feet in double row in midventral radius, scattered else- where on body wall. Maximum size 5 cm, color white or pale tan to brown Stereoderma unisemita 11 (3) Posterior projections of calcareous ring short. Tube feet hairlike, scattered all over body wall. Ossicles tables with spire of four rods; tables few or often absent. Maximum size 10- 12 cm. color brownish or greenish to nearly black Sclerodactvla briareus 11 (3) Posterior projections of calcareous ring very long. Tube feet scattered over body wall or restricted to radii. Ossicles include tables with spires usually composed of two rods (exceptionally 3-4 rods mPentamera calcigera) 12 12 (11) Tube feet hairlike, scattered all over body wall. Ossicles very numerous tables (a). Maximum size 5-10 cm, color whitish to brownish Havelockia scabra 12 (//) Tube feet restricted to radii, plates in five distinct crowded bands. Ossicles tables and few to many .13 13 (12) Tables (a) variable in shape, over- lying densely crowded layer of plates (b). Maximum size 9 cm, color white Pentamera calcigera 13 (12) Tables (a) usually oval, with four holes; elongate plates occasionally found. Maximum size 5 cm, color white or dirty brown Pentamera pulcherrima 14 (2) Taillike projection present on posterior end of dorsal surface of body. On sole, midventral ra- dius with complete double row of tube feet. Maximum size about 15 cm, largest specimens black, smaller specimens often yellowish-brown Psoitis phantapus 14 (2) No taillike projection. On sole, midventral ra- dius without complete double row of tube feet. Maximum size about 19 cm, color bright scarlet red or reddish-orange Psoitis fabricii 15 (1 ) Plump, tapering posteriorly to form a well- defined tail. Body wall opaque, leathery. Fifteen digitate tentacles (a) 16 15 (2) Cylindrical and worm-like, with 12 pin- nate tentacles (a) and translucent body wall 17 16 (15 ) Color whitish. Ossicles include tables (a) with about 14 holes and a central spire. Maximum size 25 cm Caudina arenata 16 (15) Color light red to very dark red or blackish. Ossicles in body replaced by wine-red phosphatic deposits as animal grows. Deposits in tail (a) unaffected. Maximum size 20 cm Molpadia oolitica 10 17(7) Ossicles as wheels, or as anchors with anchor plates 18 17 (7) Ossicles rods only; no wheels, anchors or anchor plates. Maximum size 4 cm, color red-brown Toxodora ferruginea 18(77) Ossicles wheels, gathered into papillae 19 18 (77) Ossicles anchors (a) associated with anchor plates (b). 20 19 (78) In addition to wheel papillae, body wall con- tains numerous curved rods with bifurcated ends. Tentacles with 5-7 pairs of digits. Max- imum size 12 cm, color dark reddish-brown . Chiridota wigleyi 19 (78) Curved rods absent. Tentacles with 2-3 pairs of digits. Maximum size 20 cm, color usually pinkish Chiridota laevis 11 20 (18) Tentacles with 4-6 pairs of digits. Radial pieces of calcareous ring (a) perforated for passage of radial nerve. Maximum size 15 cm, color generally whitish . . .Leptosynapta tenuis K7 20 (18) Tentacles with 2-4 pairs of digits. Radial pieces of calcareous ring (a) not perforated for radial nerve, but notched anteriorly. Maximum size 10 cm, color generally pinkish Epitomapta roseola ANNOTATED SYSTEMATIC LIST The species are arranged here according to currently accepted classifications (see Deichman 1930, Pawson and Fell 1965). Notes on distribution and habitat are in- cluded. Class HOLOTHUROIDEA Order Dendrochirotida FAMILY PSOLIDAE Psolus phantapus (Strussenfeldt 1765). Off coast of New England to Labrador; Greenland, Iceland, Great Brit- ain, Scandinavia; 20-250 m. Lives attached to hard substrates on gravelly bottoms, or on surface of silty sand. Psolus fabricii Duben and Koren 1846. Off Massachusetts and northwards to Ungava Bay, Greenland, Norway; intertidal (in north) — 1800+ m. FAMILY PHYLLOPHORIDAE Pentamera pulcherrima Ayres 1854. Coast of New England to North Carolina; also Gulf coast of Texas and Colombia; shallow water. In North Carolina buried in mud. Occasionally washed ashore after storms in New England. Pentamera calcigera (Stimpson 1851). New England coast to Labrador and northward, western Greenland, Alaska and Bering Sea; 10-100 m. Often found in sand, and frequently washed ashore after storms. Eaten by fish. Havelockia scabra (Verrill 1873). Halifax, Nova Scotia, to Long Island; 10-1,170 m. Most common on sandy-silt bottoms. FAMILY SCLERODACTYLIDAE Sclerodactyla briareus (Lesueur 1824). Nova Scotia south along Atlantic coast to Gulf coast of Texas; 0-20 m. Often found in muddy localities, buried or associated with eelgrass. A widely used experimental animal, re- ferred to formerly as Thy one briareus. FAMILY CUCUMARIIDAE Cucumaria frondosa (Gunnerus 1770). Nantucket, New England northward; Iceland, Greenland, northern Europe; 0-260 m. Commonly found clinging to rocks below low water mark; also occurs on sand. Trachythyone nina (Deichmann 1930). East of Cape Cod; 190-250 m. On gravel and shell-sand bottoms. Stereoderma unisemita (Stimpson 1851). Newfoundland to Long Island Sound; 10-200 m. Sand or gravel bot- tom. Ocnus diomedeae Pawson 1976. East of Cape Cod; 190- 250 m. Gravel bottoms. Thyonidium pellueidum Duben and Koren 1844. Casco Bay, Maine, to Labrador, Newfoundland, Iceland, Greenland, Great Britain, Scandinavia; 40-120 m. Gravelly or shelly bottom. Duasmodactyla commune (Forbes 1841). Coast of New England and Newfoundland; northern Europe; 0-190 m. Commonly found under stones or buried in gravel near low water mark. L2 Thyonella gemmata (Pourtales 1851). Woods Hole, Mass., to Yucatan; shallow water. Common in Gulf of Mexico. Buried in mud or sand; frequently washed ashore in large numbers. Thyonella pervicax (Theel 1886). Vineyard Sound, Mass., to Gulf of Mexico and Brazil; shallow water. Order Molpadiida FAMILY MOLPADIIDAE Molpadia oolitica (Pourtales 1851). From Newfoundland to New England; possibly extending to Florida; Bering Sea; 10-720 m. Partially buried in muddy substrates. FAMILY CAUDINIDAE Caudina arenata (Gould 1841). New Brunswick and coast of New England; 0-2,500 m. Buried in sand or mud, often washed ashore in great numbers after storms. Anatomy of this species studied by Gerould (1896). Order Apodida FAMILY SYNAPTIDAE Leptosynapta tenuis (Ayres 1851). Coast of New England; Bermuda (?); shallow water. Usually buried near low water mark in sand or mud. Epitomapta roseola (Verrill 1874). Massachusetts and Connecticut coasts; Bermuda; shallow water. Buried in sand or mud near low water mark, or under stones. FAMILY CHIRIDOTIDAE Chiridota laevis (Fabricius 1780). Massachusetts north- ward to Labrador and Greenland (exact range not known); 0-100 m. In sandy to muddy areas, often also among stones and seaweed. Chiridota wigleyi Pawson 1976. East of Long Island, N.Y.; 90 m. Silty sand. Toxodora ferruginea Verrill 1882. South of Nantucket and Martha's Vineyard; 140-280 m. SELECTED BIBLIOGRAPHY BOOLOOTIAN, R. A. (editor). 1966. Physiology of Echinodermata. John Wiley, N.Y.. 822 p. CLARK, H. L. 1901. Synopses of North-American invertebrates. XV. The Holo- thurioidea. Am. Nat. 35:479-496. 1924. The holothurians of the Museum of Comparative Zoology. The Synaptinae. Bull. Mus. Comp. Zool. Harv. Univ. 65(13): 457-501. COE, W. R. 1912. Echinoderms of Connecticut. Conn. Geol. Nat. Hist. Surv., Bull. 19, 152 p. DEICHMANN, E. 1930. The holothurians of the western part of the Atlantic Ocean. Bull. Mus. Comp. Zool. Harv. Univ. 71(3):43-226. ELLINGTON, W. R. 1976. L-lactate dehydrogenase in the longitudinal muscle of the sea cucumber Sclerodactyla briareus (Echinodermata: Holothu- oidea). Mar. Biol. (Berl.) 36:31-36. FARMANFARMAIAN, A. 1969. Intestinal absorption and transport in Thyone. II. Observa- tions on sugar transport. Biol. Bull. (Woods Hole) 137:132-145. GEROULD, J. H. 1896. The anatomy and histology of Caudina arenata Gould. Bull. Mus. Comp. Zool. Harv. Univ. 29:123-190. HYMAN, L. H. 1955. The invertebrates: Echinodermata. Vol. IV. McGraw-Hill, N.Y., 763 p. JORDAN, A. J. 1972. On the ecology and behavior of Cucumaria frondosa (Echino- dermata: Holothuroidea) at Lamoine Beach, Maine. Ph.D. The- sis, Univ. Maine, Orono, 77 p. PAWSON, D. L., and H. B. FELL. 1965. A revised classification of the dendrochirote holothurians. Breviora 214, 7 p. RHOADS, D. C, and D. K. YOUNG. 1971. Animal-sediment relations in Cape Cod Bay, Massachusetts. II. Reworking by Molpadia oolitica (Holothuroidea). Mar. Biol. (Berl.) 11:255-261. ZENKEVITCH, L. 1963. Biology of the seas of the U.S. S.R. Allen and Unwin, Lond., 955 p. 13 SYSTEMATIC INDEX Apodida 3, 13 Aspidochirotida 1, 3 Caudina arenata 10, 13 Caudinidae 13 Chiridota laevis 11, 13 wigleyi 11, 13 Chiridotidae 3, 13 Cucumaria frondosa 1, 7, 12 Cucumariidae 12 Dendrochirotida 1, 3, 12 Duasmodactyla commune 5, 12 Epitomapta roseola 12, 13 Havelockia scabra 8, 12 Leptosynapta tenuis 12, 13 Molpadia oolitica 10, 13 Molpadiida 3, 13 Molpadiidae 13 Mvriotrochidae 3 Oncus diomedeae 6, 12 Parastichopus californicus 1 Pentamera calcigera 8, 12 pulcherrima 9, 12 Phyllophoridae 12 Psolidae 12 Psolus fabricii 9, 12 phantapus 9, 12 Sclerodactyla briareus 1, 8, 12 Sclerodactylidae 12 Stereoderma unisemita 7, 12 Synaptidae 3, 13 Thyonella gemmata 7, 13 pervicax 6, 13 Thyonidium pellueidum 5, 12 Toxodora ferruginea 11, 13 Trachythyone nina 7, 12 14 ACKNOWLEDGMENTS Preparation of the "Marine Flora and Fauna of the Northeast- ern United States" is being coordinated by the following Board: Coordinating Editor: Melbourne R. Carriker, College of Marine Studies, University of Delaware, Lewes, DE 19958. Editorial Advisers: Marie B. Abbott, Marine Biological Labora- tory, Woods Hole, Mass. Arthur G. Humes, Boston University Ma- rine Biological Laboratory, Woods Hole, Mass. Wesley N. Tiffney, Department of Biology, Boston University, Boston, Mass. Ruth D. Turner, Museum of Comparative Zoology, Harvard University, Cambridge, Mass. Roland L. Wiglery, National Marine Fish- eries Service, Biological Laboratory, Woods Hole. Mass. Robert T. Wilce, Department of Botany, University of Masschusetts, Amherst, 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, reviews manuscripts, and advises the Scientific Editor of the National Marine Fisheries Service. Collections and records of the National Museum of Natural History, Smithsonian Institution, Washington, D.C.; the Museum of Comparative Zoology, Harvard University; the Northeast Fisheries Center, National Marine Fisheries Service, Woods Hole; and the Gray Museum, Marine Biological Laboratory, Woods Hole, provided reference materials for this paper. Thanks go to Roland L. Wigley of the National Marine Fisheries Service for his help with collections, John H. Dear- born, University of Maine, for critically reading the manuscript, and Irene Jewett, for preparing most of the illustrations. 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. David L. Pawson commenced his study of the systematics and zoogeography of echinoderms in 1958, working mainly on echinoids and holothuroids of the southern Pacific Ocean. In 1964 he joined the staff of the National Museum of Natural His- tory, Smithsonian Institution, where he continues studies on echinoids and holothuroids of the Atlantic and Indo-Pacific regions. 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 LTnited States. Cnidaria: Scyphozoa. CAVALIERE, A. R. Marine flora and fauna of the Northeastern United States. Higher fungi: Ascomycetes, Deuteromycetes, and Basi- diomycetes. 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. PAWSON, DAVID L. Marine flora and fauna of the Northeastern United States. Echinodermata: Holothuroidea. 15 -CtGPO 798-587 388. 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 by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 392. Fishery publications, calendar year 1974: Lists and indexes. By Lee C. Thorson and Mary Ellen Engett. June 1975, iv + 27 p., 1 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. By Leland W. Pollock. May 1976, iii + 25 p., figs. For sale by the 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 1 107 N E 45TH ST SEATTLE WA 98105 PETlPiT|A.!E UNIVERSITY LIBRARIES A0D0Q7E01flfi?3 OFFICIAL BUSINESS NOAA SCIENTIFIC AND TECHNICAL PUBLICATIONS NOAA, the National Oceanic and Atmospheric Administration, was established as part of the Department of Commerce on October 3, 1970. 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