438 c ^. \ ^^1^ NOAA Technical Report NMFS Circular 438 Marine Flora and Fauna of The Northeastern United States. Scleractinia Stephen D. Cairns July 1981 — — J Marine Biological Laboratory ': LIBRARY OCT 141992 ! I Woods Hole, Mc-ss. j 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 ihe 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 enforce- ment of domestic fisheries regulations, surveillance of foreign fishing off United States coastal waters, and the development and enforcement of interna- tional 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 investigations appear in the Circular series. NOAA Technical Report NMFS Circulars are available free in limited numbers to governmental agencies, both Federal and Stale. 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 D822, User Services Branch, Environmental Science Information Center, NOAA, Rockville, MD 20852. Recent Circulars are: 418. Annotated bibliography of four Atlantic scombrids: Scomberomorus brasiliensis, S. cavalla, S. maculalus, and S. regalis. By Charles S. Manooch III, Eugene L. Nakamura, and Ann Bowman Hall. December 1978, iii-i- 166 p. 419. Marine fiora and fauna of the northeastern United States. Protozoa: Sar- codina: Amoebae. By Eugene C. Bovee and Thomas K. Sawyer. January 1979, iii + 56 p., 77 figs. For sale by the Superintendent of Documents, U.S. Govern- ment Printing Office, Washington, DC. 20402, Stock No. 003-017-00433-3. 420. Preliminary keys to otoliths of some adult fishes of the Gulf of Alaska, Bering Sea, and Beaufort Sea. By James E. Morrow. February 1979, iii + 32 p., 9 pi. 421 . Larval development of shallow water barnacles of the Carolinas (Cirripe- dia: Thoracica) with keys to naupliar stages. By William H. Lang. February 1979, iv + 39 p., 36 figs.. 17 tables. 422. A revision of the catsharks, family Scyliohinidae. By Stewart Springer. April 1979, v+ 142 p., 97 figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, DC. 20402, Stock No. 003-020-00147-5. 423. Marine flora and fauna of the northeastern United Stales. Crustacea: Cumacea. By Les Watling. April 1979, iii + 23 p.. 35 figs. For sale by the Superintendent of Documents, Washington, D.C. 20402, Stock No. 003-017-00446-5. 424. Guide to the leptocephali (Elopiformes, Anguilliformes, and Notacanthi- formes). By David G. Smith. July 1979, iv + 39 p., 54 figs. 425. Marine fiora and fauna of the nonheastem United Slates. Anhropoda: Cirripedia. By Victor A. Zullo. April 1979, iii + 29 p., 40 figs. For sale by the Superintendent of Documents, Washington, D.C. 20402, Stock No. 003-017-00453-8. 426. Synopsis of biological data on the rock crab. Cancer irroralus Say. By Thomas E. Bigford. May 1979, v + 26 p., II figs., 21 tables. 427. Ocean variability in the U.S. Fishery Conservation Zone, 1976. By Julien R. Goulet, Jr., and Elizabeth D. Haynes. editors. July 1979, iii + 362 p. (427.) Introduction. By Julien R. Goulet, Jr. July 1979, p. 1-2. (427.) Summary. By Julien R. Goulet, Jr. July 1979, p. 3-10. (427.) Atmospheric circulation in 1976. By Elizabeth D. Haynes. July 1979, p. 11-18, 2 figs. (427.) Atmospheric climatology and its effect on sea surface temperature— 1976. By Robert R. Dickson and Jerome Namias. July 1979, p. 19-33, 6 figs. (427.) Eastern Pacific sea surface conditions in 1976. By Elizabeth D. Haynes. July 1979, p. 35-42. (427) Sea surface conditions in the western North Atlantic in 1976. By Julien R. Goulet, Jr., and Elizabeth D. Haynes. July 1976, p. 43-50. (427.) Anomalies of monthly mean sea level along the west coasts of North and South America. By Dale E. Bretschneider and Douglas R. McLam. July 1979, p. 51-64, 6 figs. (427.) Coastal upwelling off western North America, 1976. By Craig S. Nelson. July 1979, p. 65-75, 2 figs., 2 tables. (427.) Oceanic conditions during 1976 between San Francisco and Honolulu as observed from Ships of Opportunity. By J. F. T Saur and D R. McLain. July 1979, p. 77-92, 5 figs., 2 tables. (427.) The 1976 El Nino and recent progress in monitoring and prediction. By William H. Quinn. July 1979, p. 93-110, 7 figs., 4 tables. (427.) Sea surface temperature anomalies. By Douglas R. McLain. July 1979, p. 111-149, app. 1. (427.) Fluctuations of sea surface temperature and density at coastal stations during 1976. By Douglas R. McLain. July 1979, p. 151-166, 7 figs., app. I. (427.) Data on cold weather conditions along the Atlantic and Gulf coasts dur- ing the fall and winter of 1976-77. By J. Lockwood Chamberlin and Reed S. Armstrong. July 1979, p. 167-174, I fig., I table. (427.) Wind driven transport Atlantic coast and Gulf of Mexico. By Merton C. Ingraham. July 1979, p. 175-208, 4 figs., 1 table, app. I. (427.) Sea surface temperature distribution from Cape Cod, Massachusetts, to Miami, Florida - 1976. By Joseph W. Deaver III. July 1979, p. 209-229, 3 figs.. 2 tables, app. I . (427.) Water column thermal structure across the shelf and slope southeast of Sandy Hook, New Jersey, in 1976. By Steven K. Cook. July 1979, p. 231-257, 19 figs. (427.) Anticyclonic Gulf Stream eddies off the Northeastern United States dur- ing 1976. By David Mizenko and J. Lockwood Chamberlin. July 1979, p. 259-280, 16 figs., 2 tables. (427.) River runoff along the Middle Atlantic coast in 1976. By Elizabeth D. Haynes. July 1979, p. 281-287, I fig., 1 table, app. 1. (427.) Climatic conditions related to the fish kill and anoxia off New Jersey during the summer of 1976. By Reed S. Armstrong. July 1979, p. 289-300, 5 figs. (427.) Variations in Ihe position of the shelf water front off the Atlantic coast between Georges Bank and Cape Romain in 1976. By John T. Gunn. July 1979, p. 301-314, 8 figs., 1 table. (427.) Temperature structure on the continental shelf and slope south of New England during 1976. By R. Wylie Crist and J Lockwood Chamberlin. July 1979, p. 315-335,2 figs., app. 1. (427.) Continuous plankton records: Zooplankton and net phytoplankton in the Mid-Atlantic Bight, 1976. By Daniel E. Smith and Jack W. Jossi. July 1979, p. 337-348, 7 figs. (427.) Siphonophore ("lipo") swarming in New England coastal waters- update, 1976. By Carolyn A. Rogers. July 1979, p. 349-352, I fig. (427.) Bottom-water temperatures in the Gulf of Maine and on Georges Bank during spring and autumn, 1976. By Clarence W. Davis. July 1979, p. 353-362, 6 figs., 3 tables. fS^£JJg^,9, NOAA Technical Report NMFS Circular 438 Marine Flora and Fauna of the Northeastern United States. Scleractinia Stephen D. Cairns July 1981 Marine Biological Laboratory LIBRARY OCT 14 1992 Woods Hole, Mass. U.S. DEPARTMENT OF COMMERCE Malcolm Baldrige, Secretary National Oceanic and Atmospheric Administration National Marine Fisheries Service Terry L. Leitzell, Assistant Administrator for Fisheries FOREWORD This NMFS Circular is pari of the subseries "Marine Flora and Fauna of the Northeastern United States," which consists of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants and animals of the Northeastern United States. The manuals are published at irregular intervals on as many laxa of the region as there are specialists available to collaborate in their preparation. Geographic coverage of the "Marine Flora and Fauna of the Northeastern United States" is planned to include organisms from the headwaters of estuaries seaward lo 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 ihe manuals are deposited in the reference collections of major museums of the region. The "Marine Flora and Fauna of the Northeastern United Stales" is being prepared in collaboration with systematic specialists in the United Slates and abroad. Each manual is 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, includes an introduction, illustrated glossary, uniform originally illustrated keys, annotated checklist with information when available on distribution, habitat, life history, and related biology, references to the major literature of the group, and a systematic index. These manuals are intended for use by biology students, biologists, biological oceanographers, informed laymen, and others wishing to identify coastal organisms for this region. Often they can serve as guides to additional information about species or groups. The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods Hole Region;' edited by R. 1. Smith in 1964. and produced under the auspices of the Systematics Ecology Program, Marine Biological Laboratory, Woods Hole, Mass. After a .sufficient number of manuals of related taxonomic groups have been published, the manuals will be revised, grouped, and issued as special volumes, which will consist of compilations for phyla or groups of phyla. CONTENTS Iniroduction 1 Morphology 2 Keys to the Scleractinia of the northeastern coast of the United States 3 Dichotomous key 3 Tabular keys 9 Annotated systematic list 10 Selected bibliography 11 Systematic index 13 Acknowledgments 14 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. Marine Flora and Fauna of the Northeastern United States. Scleractinia STEPHEN D. CAIRNS' ABSTRACT This manual discussi-s Ihc 14 species inlot;> and ni nt Scleraclinia. an illiislraled dicholomons ke> and (ho (ahnlar ke>s are t£i%en t<»r ihese species. An annota(ed s\s(ema(ic lfs( includes complete i:eot>raphic and hath>me(rie ranges, relerences lo pi>rlinen( lileralurc, and. fur some species, ecological and (avonomic nt»les. /oo^eo^raphic urfinilies nl' the I'auna are brielU discussed. A selec(ed hihlio|^rapli> is provided. INTRODUCTION Fourteen species of stony corals (order Scleractinia) are known from off the northeastern coast from Chesapeake Bay to southern Nova Scotia of which only one, Astrangia astreiformis, occurs at depths shallow enough to be collected routinely by snorkeling or scuba diving. The remaining species are usually collected by benthic trawls or dredges and occur as deep as 3,200 m off the northeast coast. Much research has been done on the easily accessible A . asireiformis, but little more than physical descriptions and distributions are known for the deeper water species. All 14 species are included in this report. Of these 14 species, 13 are ahermatypic; i.e., they do not possess symbiotic zooxanthellae (a unicellular dinoflagellate) in their endodermal tissue. Individual colonies of A. asireiformis may or may not have zooxanthellae, depending perhaps on water temperature or light intensity; more frequently this species lacks the symbionts. The term ahermatypic is thus a physiological condition determined by ecological factors and therefore is not a character of great value in classification. Ahermatypic corals have been equated with deepwater corals, solitary corals, or nonreef building corals. This is an oversimplification; in fact, many species occur in shallow water, and some large, colonial deepwater (500-800 m) ahermatypes (e.g., Lophelia prolifera, Enallopsammia profunda) form the framework for reeflike structures. It is true, however, that all deepwater corals are ahermatypic because below the euphotic zone the zooxanthellae, being plants, cannot photosynthesize. Not restricted by the generally higher (often tropical) temperature and light requirements that zooxanthellae impose on hermatypic corals, the ahermatypes inhabit a more extensive geographic range. They are found in all oceans from the Norwegian Sea (lat.70°30 'N) to the Ross Sea, Antarctica (lat.78°29 S). from 0-6,328 m depth (Keller 1976), and in temperatures of - l.rc to over 29°C. Scleractinia are monoecious or dioecious; in either case their motile larva] form is a planula capable of remaining planktonic for weeks. In addition to sexual reproduction, some species propagate by asexual budding and others have remarkable powers of regeneration. For instance, the corallum of Dasmosmilia lymam usually splits longitudinally into several Ucpanmcni of Invcnibraii; Zoolog>, National Museum ol Naiutal Hisui Sniiihsoniaii Imiilulion. \Vashiiii;ioii. D.C. 20560. fragments, each wedge-shaped piece subsequently producing 1-30 small buds growing directly from the mesenterial tissue. It is rare to find a specimen of D. lymani that grew directly from a planula, i.e., not attached to an inside fragment of a parent specimen. All planulae need a hard substrate on which to settle. Following settlement and subsequent growth, corals may either remain attached to the substrate or become free, lying unattached on the substrate. The attached corallum usually reinforces its base of attachment by various means, whereas the subsequently unattached corallum becomes free by lacking reinforcement of an originally weak attachment or by completely overgrowing the substrate, as in the case of a sand particle. Some unattached species (e.g., Caryophyllia ambrosia) become top-heavy, fall to one side, but subsequently reorient their calices toward an upright position, producing a horn- shaped corallum. The scleractinian fauna of the northeastern coast of the United Stales is relatively low in diversity when compared to other areas in the western Atlantic, even when the reef corals (hermatypes) are excluded from consideration. For example, there are approximately 160 species of Scleractinia in the Caribbean: 76 deepwater (over 200 m) ahermatypes (Cairns 1979), about 30 exclusively shallow-water ahermatypes, and about 54 hermatypic species. In the region between north Florida and Cape Hatteras there are about 40 species known: 28 deepwater ahermatypes (Cairns 1979), about 8-10 exclusively shallow-water ahermatypes, and several hardy hermatypic species, i.e., Solenoasirea, Siderastrea, Oculina. In the western Atlantic north of the North Carolina-Virginia border there are 17 species known, 16 deepwater ahermatypes and one exclusively shallow-water species, A. asireiformis. Fourteen of these 17 species are treated in this work, the other three: Vaughanella margariiaia (Jourdan, 1895); Fungiacyathus durus Keller, 1976; and F. marenzelleri (Vaughan, 1906) are known from localities north of Maine. Other species occurring south of the area of consideration for this manual that may subsequently be found there include: Dendrophyllia gadilana (Duncan, 1873); Pourialosmilia conferla Cairns, 1978; Concentrolheca laevigala (Pourtales, 1871); and Polymyces fragilis (Pourtales, 1868). The 14 species of Scleractinia known from off the northeastern coast of the United States are, in general, widely distributed species. Six are cosmopolitan and seven are amphi- Atlantic. Of the latter, five are primarily cold temperate, one (/4 . astreiformis) is found both in temperate and tropical waters, and one (Deltocyathus italicus) is primarily a tropical species with northern limits in the cold temperate region. Only one species, Enaltopsammia profunda, is endemic to the western Atlantic and is most common in the warm temperate region. MORPHOLOGY The calcareous skeleton of a scleractinian coral, the coraUum, may be composed of numerous individual units producing a colonial corallum or it may be a solitar}' coral with only one unit produced by one polyp. The shape of the corallum of solitary corals is important at the generic and specific levels and is commonly described in geometric terms (e.g., conical, trochoid, cylindrical); shapes of colonial corals are described by branching pattern. Corals may be either attached to the substrate or unattached (free), this difference usually being consistent at the species level. If attached, the base of the coral is fu-mly cemented to a hard surface. A solitary attached coral usually has a stemlike pedicel directly above the base (Fig. 1), which supports the calice, the round to elliptical oral surface of the coraUum. The solitary attached coral often reinforces its attachment by ^PALUS COLUMELLA PEDICEL BASE thickening its pedicel, expanding its base, and, in some species, producing anchoring rootlets or adding successive rings of compartmentalized, concentric chambers around the base and pedicel. A solitary unattached coral usually has a flat or bowl- shaped base and lacks the pedicel. The sides or walls of solitary corals and corallites of colonies are termed the theca. This theca may be granular or porcelaneous in texture and often bears longitudinal ridges called costae corresponding to the larger septa. Skeletal deposits formed between individual coralhtes of a colony are called coenosteum. The calice of most species is regularly and hexamerally subdivided by radial partitions, called septa. The six largest septa, divide the calice into six equal areas or systems (Fig. 2), and comprise the first cycle of septa. The second cycle also con- sists of six septa which are generally smaller and occur halfway between the first cycle septa. The 12 third cycle septa are formed in each space created by the previous 12 septa. Calices with 6-7 cycles of septa (192-384 septa) are known. Septa with the upper edges extending above the theca are termed exsert septa. Sometimes one or several adjacent septa of a calice are extreme- ly exsert and bent over the calice, forming a hoodlike structure called a rostrum (e.g., see figure of Enaltopsammia rosiraia). Small accessory lobes are sometimes present on the inner edges of the septa of certain cycles. These are called pali (singular: palus), or paliform lobes, and are often used as a generic level character. Directly in the center of the calice there is often a structure called a columella, which may appear as a lamella (lamellar), a spongy mass (trabecular), a single rod (styliform), a field of simple or twisted rods (papillose, fascicular), or simply a fusion of the inner edges of the larger septa (rudimentary). The type of columella is often used to distinguish genera and subgenera. ' SYSTEM -Cutaway drawing of hypothetical : raorphological features. slitary coral illustrating Figure 2. — Diagram of relative sizes and position of septa in a calice containing three cycles (24 septa). Numben refer to the cycle to which the septum belongs; only the upper right system is completely numbered. KEYS TO THE SCLERACTINIA OF THE NORTHEASTERN COAST OF THE UNITED STATES The identification and classification of Scleractinia depend entirely on characteristics of the corallum; therefore, it is generally necessary to remove the tissue from the corallum before using the keys. This is easily accomplished by soaking the coral in full-strength commercial bleach for several hours followed by thorough rinsing. The specimens should then be stored dry. An illustrated, dichotomous key is presented first, followed by two tabular keys to the same species: one keying the colonial species and the other, the solitary ones. A blank space in the tabular key indicates that this character does not apply to this species. DICHOTOMOUS KEY 1 Corallum solitary (see tabular key 1 ) 2 1 Corallum colonial (see tabular key 2) 10 2 (/) Corallum firmly attached to substrate 3 2 (J) Corallum unattached or attached to small fragment of parent specimen 4 3 (2) Five or more cycles of septa (>96 septa); septa and theca thick, robust; theca granular and costate; inner septal edges straight (Fig. 3) Desmophyllum cristagatti Figure 3. — Desmophyllum crislagalli: side view. Scale: 3 cm. Figure 4. — Javania cailleti: A, side >iew of specimen with damaned upper septal edges; B. side view of specimen w illi intact septa, UlustrallnK liroad. encrusting attachment. Scales: 1 cm. 3 (2) Usually four cycles of septa (48 septa), occasionally 64 septa; septa and upper theca very delicate; theca smooth and porcelaneous; inner septal edges sinuous (Fig. 4A, B) Javania cailleti 4 (2) Pali or paliform lobes present 5 4 (2) Pali or paliform lobes absent 7 5 (4) Corallum attached to fragment of parent corallum or has broken base; corallum fragile, easily fragmented; paliform lobes often multilobate and indistinguishable from the columella (Fig. 5 A, B) Dasmosmilia lymani Figure 5. — Dasmosmilia lymani: A, fragment of larger parent corallum with three small coralla asexually budding from parent; B, side view of intact corallum. Scale: 2 cm. 5 {4) Corallum free, base intact; corallum strong, not easily fragmented; pali single-lobed and distinct from col- umella 6 6 (5) Pali (12-18) of uniform size occur on inner edges of third cycle septa; corallum a curved cone (horn- shaped); calicular diameter up to 40 mm (Fig. 6) Caryophyllia ambrosia ambrosia Figure 6. — Caryophyllia ambrosia ambrosia: side view showing crown of pali and several central columellar ribbons. Scale: 2 cm. 6 (5) Pali of different sizes occur on first, second (sometimes missing), and third cycle septa; corallum conical but not curved; calicular diameter rarely over 15 mm (Fig. 7 A, B) Deltocyathus italicus Figure l.—Dellocyathus italicus: A, calicular view showing septal and palar airangemenl and columella; larger pali before third cycle sepU, smaller before Hrst cycle sepU; pali missing before second cycle; B, side view. Scales: 5 mm. 7 {4) Corallum discoidal; base flat (Fig. 8) . . Fungiacyathus fragilis Figure S.—tungiacyathus fragilis: CalicuUir (w of slightly damaged specimen. Scale: I cm. 7 {4} Corallum conical, compressed-coni- cal, or fragmented; base not flat . . . . 8 (7) Rudimentary columella formed by fusion of lower inner edges of larger septa; corallum usually intact (7) No columella; corallum invariably fragmented (Fig. 9) Flabetlum macandrewi Figure 9. — FlabeBum macandrewi; side view. Scale: 1 cm. 9 {S) Calice compressed or constricted in center (Fig. lOA, B) Flabellum alabastrum Figure \0.— FlabeBum alabastrum: A, calicular view showing conslricled calice; B, side view. Scale: 2 cm. 9 (8) Calice a regular ellipse (Fig. 11) . . Flabellum angulare Figure 11. — flabellum angulare: side view. Scale: 2 cm. 10 (/) Corallum encrusting, rarely producing short, stubby branches; inner edges of septa dentate; corallites directly adjacent (Fig. 12) Astrangia aslreiformis Figure 12. — Astrangia astreiformis: an average-sized colony encrusting a rocli. Scale: 2 cm. 10 (/) Corallum branching, arborescent; inner edges of septa smooth; corallites not directly adjacent 11 1 1 (10) Colonies uniplanar, with calices arranged on only one side; each calice bordered proximally by a rostrum (hood) (Fig. 1 3 A, B) Enallopsaminia rosirata Figure M.—EnaUopsammia roslrala: A, a branch from pan of a larger colo- ny; note triangular swellings (rostra) beneath each calice. Scale: 2 cm. B, en- largement of rostrum of one calice. Scale: 3 mm. 1 1 (10) Colonies bushy, with calices randomly arranged on branches; no calicular rostra 7 12 (//) Columella present; septa not exsert; coenosteum finely porous, especially near calices; usually only 24 septa per calice(Fig. 14) Eiwllopsuiiuiiui profunda Figure 14. — EnaUopsammia profunda: a branch from part of a larger colony. Scale: 5 cm. 12 (//) Columella absent; septa exsert; coenosteum not porous; usually more than 24 (up to 60) septa per calice 13 13 (12) Calices in the process of intratentacular division common (various stages of splitting usually pre- sent); first three cycles of septa hexamerally- arranged, septa of fourth and fifth cycles added irregularly; nonsplitting calices rarely exceed 5 mm in calicular diameter; coenosteal costae often extend up to 10 mm from calicular edge (Fig. 15) Solenosmilia variabilis Figure i5.— Solenosmilia variabilis: a dislal branch of a larger colony. Scale: 3 cm. 13 (/2) Calices in process of intratentacular division rare (most calices, even terminal ones, appear discrete); septa not hexamerally arranged; calices up to 15 mm in calicular diameter; coenosteal costae rarely extend more than 5 mm from calicular edge (Fig. 16) Lophelia prolifera Figure 16. — Lophelia prolifera: a branch from part of a larger colony. Scale: 5 cm. Table I. — Tabular key for soGlary species. Table 2. — Tabular key for cokinial species. = E Fingiacyalhus fragilis Caryophyllia ambrosia Dasmosmitia lymani Deltocyalhus italicus Ftabellum alabastrum^ Ftabeltum macandrewi Ftabellum angulare^ Desmophyllum crisiagalli Javania cailteli ex "S T^ « 0. < o o 1 o 2 + 2 o O o 'Attachment of corallum: a — attached: f — unattached, flat base; c — unat- tached, conical base. ^Height of septa: e — exsert; n — not exsert. ^Texture of thcca: p — porcelaneous; g — granular. ^Presence of pali or paliform lobes: O — absent; 1 — on third cycle septa; 2 — on all but last cycle septa. ^Asexual budding: + — present; O — absent. 'Fragility of corallum: r— robust, usually intact; f— fragile, easily broken; vf — very fragile, usually found in fragments. ^Columella: O — absent; r — rudimentary; f — fascicular (rods, twisted rib- bons, papillose); s — trabecular (spongy). ^Ftabellum alabasirum and F. angulare differ mainly in the shape of their co- ralla, which is not a coded character in this key. ^ =a 3 = 2=2 3 o 8- O 8- ffl O crt U V) Aslrangia aslrei/ormis Lophelia prolifera Solenosmilia variabilis Enallopsammia profunda Enallopsammia roslrata 'Shape of corallum: a — arborescent; e — encrusting. ^Inner edges of septa: d — dentate; s — smooth. 'Orientation of calices on branch: I — calices occur on only one side of branch; colony uniplanar; 2 — calices occur on all sides of branch; colony bushy. •"Calicular rostrum: r— present; O — absent. 'Budding: e — extratenlacular; is — intratentacular. sympodial; id— intraten- tacular, dichotomous. ^Columella: s — spongy; p — papillose; O — absent. ^Height of septa: e— exsert; n— not exsert. 'Texture of coenosteum near calice: p — porous; s— solid. 'Arrangement of sepia: h— hexameral; i— irregular. ANNOTATED SYSTEMATIC LIST This list follows the classification proposed by Wells (1956). Genera are arranged alphabetically within families. Geographic ranges include all published records and data from additional specimens examined at the National Museum of Natural His- tory, Smithsonian; Yale Peabody Museum; and Museum of Comparative Zoology, Harvard. References to significant papers are cited in brackets at the end of each account. Suborder FUNGIINA Superfamily FUNGIICAE Family FUNGIIDAE Fiingiacyalluisfragilis Sars, 1872. Distribution probably world wide. In western Atlantic, known from continental slope south of Cape Cod, Mass. (412-460 m), 5.5"-6.1°C. Also known from eastern Atlantic, off Hawaii, and south of New Zealand (285-2,200 m). Because of its fragility it is usually damaged or in fragments when collected. It is distinguished from other closely related species by its possession of five cycles of septa and septa with sinuous upper edges. [Zibrowius 1980.] Suborder FAVDNA Superfamily FAVDCAE Fanuly RHIZANGITOAE Astrangia astreiformis Milne Edwards and Haime, 1849 (=As- trangia danae Agassiz, 1850; not Astrangia danae Milne Edwards and Haime, 1849). Star coral. Northern coral. Massachusetts to Texas from low tide to 35 m (- 1 °-22°C). This is the only scleractinian likely to be seen by snorkeling or with scuba off the northeastern coast of the United States. Also known from Puerto Rico and off tropical western Africa. Common on any hard substrate, i.e., stones, shells, pilings. May or may not have zooxanthellae. Often used as an experimental laboratory animal. Physiological studies include: Cummings (1976); Jacques et al. (1977); Szmant- Froelich and Pilson (1977); and Jacques (1978). Natural his- tory accounts include: Agassiz (1850); Fewkes (1889); and Bachand (1978). Milne Edwards and Haime (1849) described both Astrangia astreiformis and A. danae in the same paper; A. danae has page priority but, because the type is lost, the description is poor, and the type-locality is unknown, it is considered a nomen dubium. The type of A. astreiformis is also lost, but because Milne Edwards and Haime at least designated the type-locality of United States, it is chosen as the valid name for the common shallow-water Astrangia of the eastern and Gulf coasts. Louis Agassiz, in a paper read on 15 August 1849, independently described the same species and called it Astrangia danae, but because his account was not published until 1850, it is a junior synonym of A. astreiformis. Suborder CARVOPHYLLIINA Superlamily CARVOPHVLLIICAE Family CARYOPHYLLIIDAE Caryophyllia ambrosia ambrosia Alcock, 1898. Continental slope off Georges Bank (1,487-2,286 m), 3.3'=-4.2°C. Also known from eastern Atlantic and Indian Oceans (1,600-2,670 m). The other subspecies, C. ambrosia caribbeana Cairns, 1979, is found in more southern, shallower waters. It differs in having a broader, more open corallum and a rougher thecal texture. [Zibrowius 1980.] Dasmosmilia lymani (Pourtales, 1871). Common on outer edge of continental shelf from Alabama to south of Cape Cod, Mass. (48-366 m), 7°-21°C. Also known from off Venezuela, southeastern Brazil, and, in the eastern Atlantic, in area bordered by Portugal, the Azores, and Spanish Sahara (85-316 m). This species most frequently propagates by asex- ual budding from longitudinal fractures of the fragile coral- lum. Five to ten small buds originating from one wedge- shaped fragment are not uncommon. This is the most com- monly collected coral from the study area and is not likely to be confused with any other species. (Cairns 1979.) Deltocyathus iialicus (Michelotti, 1838). Common from Florida to southern Brazil, including Gulf of Mexico and Caribbean; Bermuda (403-2,634 m), 3°-7°C. One disjunct record on continental slope off New Jersey (Albatross station 2103), 4°C. Also known from eastern Atlantic and Azores ( 1 ,500-2,300 m). Some deepwaler trawls result in hundreds of specimens. Distinguished from other species in this genus by its distinctive conical base and frequent absence of pali on the second cycle septa. Coralla of some specimens have a pink pigmentation. [Cairns 1979.] Desmopliyl/iim cristagalli Milne Edwards and Haime, 1848. Cosmopolitan: widespread in Atlantic, Pacific, and Indian Oceans; Subantarctic; off Georges, Sable, and Grand Banks (off New England coast); Muir Seamouni and seamounts between San Pablo and Kelvin Seamounts (off New England coast). Worldwide depth range: 35-2,460 m. This species has no columella or pali. Polyp light orange. Found in great num- bers on undersides of ledges in Lydonia Canyon, off Massa- chusetts. [Cairns 1979.) Loplielia pro/ifera (Pallas, I766)( = ?Z-. pertusa (Linnaeus, 1758) nomen dubium). Common in western Atlantic from Nova Scotia to southeastern Brazil (95-1,000 m), 3''-12°C. Also known from eastern Atlantic, Indian, and Pacific Oceans (60-2,170 m). Abundant on Blake Plateau and in Straits of Florida as a major constituent of deepwaler coral banks. Growth rate 6-8 mm/yr. Systematics (Cairns 1979); ecology [Wilson 1979a, bj. Solenosmilia variabilis Duncan, 1873. Known from only two records off northeastern United States: Lydonia Canyon, off Cape Cod, Mass., and south of San Pablo Seamount. Also known from Muir Seamount; continental slopes from Georgia to southeastern Brazil (excluding Gulf of Mexico); eastern Atlantic; south of Greenland and Iceland; Indian Ocean; off southeastern Australia (280-2,165 m). Similar to L . proliferu but easily distinguished by its equal, inlratenlacu- lar budding, which always produces some terminal calices that are in the process of splitting in two. Polyps light orange. Superfamily FLABELLICAE Family FLABELLIDAE Flabellum alabusiriim Moseley, 1873 { = flabellum goodei Ver- rill, 1878. in part). Common on continental slope from Georgia to Davis Strait, including Gulf of Maine (357-1,977 m), 3.3°-7.0°C. Also known from eastern Atlantic from off Hebrides to Gulf of Guinea ( 1 ,200-2,000 m). Corallum some- times reddish brown. [Zibrowius 1980.] 10 Flahellum macandrewi Gray, 1849 ( = Flabellum goodei Verrill, 1878, in part). Fairly common on continental slope froin Virginia to Nova Scotia (180-667 m), 4.5°-8.3°C. Also known in eastern Atlantic from Norway to Senegal (128-1,170 m), 5°-7°C. Very similar to previous species but differs in that it 1) is invariably found in fragments having 3-24 septa, 2) lacks a columella, 3) has a more jagged calicular edge, and 4) has a shallower bathymetric range. (Zibrowius 1980.] Flabellum angulare Moseley, 1876. Known from southern Nova Scotia south to the continental slope of South Carolina (2,266-3,186 m), 2.5°-5.0°C. Also known in the eastern Atlantic from off Scotland south to Morocco and the Azores (1,647-2,800 m). Very similar to F. alabaslruin but differing in that 1) the outline of the calice is always elliptical, not con- stricted, and 2) the corallum is always white, never reddish brown. [Zibrowius 1980.) Javania cailleri (Duchassaing and Michelotti, 1864) ( = Desnwphyltum eburneuw Moseley, 1881; Desinophyllum nobile Verrill, 1885). Known from off Banquereau Bank, Nova Scotia, south to Oceanographer Canyon, off Cape Cod, Mass., and the continental slope of Georgia south to Burdwood Bank, Argentina; eastern Atlantic; Indian and Pacific Oceans (400-2,165 m), 6°-16°C. Distinguished from Desinophyllum crislagalli by a lesser number of septa and porcelaneous theca; however, probably indistinguishable in situ, i.e., from a submersible. (Cairns 1979.] Suborder DENDROPHYLLIINA Family DKNDROPHVLI.IIDAE Fnallopsammia profunda (Pourtales, 1867). Known from only three records off northeastern United States, all on con- tinental slope off Georges Bank (1,211-1 ,748 m), 3.5 °-3.7 °C. Also known from continental slope from South Carolina south through Straits of Florida and Lesser Antilles (403-1,337 m), 3°-12°C. Abundant on Blake Plateau and in Straits of Florida where, along with Lophelia prollfera, it is a primary constituent of deepwater coral banks. [Cairns 1979.] Fnallopsammia rosirala (Pourtales, \S1S)( = Fnallopsammia amphelioldes (Alcock, 1902)). Known from only three records off northeastern United States: continental slopes off San Pablo, New England, and Atlantis 11 Seamounts (1,174-1,646 m). Also known from continental slope from Georgia to Brazil (5°-13°C.); eastern Atlantic, western and central Pacific, Indian Oceans, and south of New Zealand (229-2,165 m). Requires hard substrate to support large cor- allum. Each calice bordered by a rostrum. [Cairns 1979.] SELECTED BIBLIOGRAPHY AGASSIZ. L. 1850. On the structure of coral animals. Proc. Am. Assoc. Adv. Sci. 2:68-77. ALCOCK, A. 1898. An account of the deep-sea Madreporaiia collected by the royal marine survey ship Investigator. Trustees Indian Museum. Calcutta, 29 p., 3 pU. 1902. Report on the Deep-sea Madreporaria of the Siboga-Expedi- ion. Siboga Exped. 16a, 31 p., 3 pU. BACHAND. R. G. 1978. Cold water coral. Sea Front. 24:283. CAIRNS, S. D. 1979. The deep-water Scleractinia of the Caribtwan Sea and adjacent waters. Stud. Fauna Curacao Other Caribb. Isl. 57(180), 341 p. including 40 pis. CUMMINGS, C. E. 1976. The effects of temperature and salinity on the survival and res- piration of Astrangia danae with and without zooxanthellae. M.S. Thesis, Univ. Rhode Island. DUCHASSAING, P., and J. MICHELOTTI. 1864. Supplement au memoire sur les coralliaires des Antilles. Mem. Acad. Sci. Turin, Ser. 2, 23:97-206. DUNCAN, P. M. 1873. A description of the Madreporaria dredged up during the expedi- tions of H.M.S. 'Porcupine' in 1869 and 1870. Part 1. Trans. Zool. Soc. Lond. 8:303-344, pis. 39^9. FEWKES, J. W. 1889. The anatomy of Astrangia danae. Smithsonian Inst., Wash., D.C., 22 p., 6 pis. GRAY, J. E. 1849. Description of some corals, including a new British coral discov- ered by W. MacAndrew, Esq. Proc. Zool. Soc. Lond. 17:74- 77. JACQUES, T. G. 1978. Metabolism and calcification of the temperate scleractinian coral Astrangia danae. Ph.D. Thesis, Univ. Rhode Island, Kingston, 187 p. JACQUES. T. G.. M. E. Q. PILSON, C. E. CUMMINGS, and N. MARSHALL. 1977. Laboratory observations on respiration, photosynthesis, and fac- tors affecting calcification in the temperate coral Astrangia danae. Proc. Third Int. Coral Reef Symp. 2:435-461. KELLER, N. B. 1976. Glubokovodnee madreporovee koralle roda Fungiacyathus iz kurilo-kamchatskogo i aleutskogo zhelobovoi nekolorekh drugikh ra- jonov mirovogo okeana. (The deep-sea madreporarian corals of the genus Fungiacyathus from the Kurile-Kamchatka Aleutian trenches and other regions of world ocean.) Tr. P. P. Shirshov Inst., Okeanol. Akad. Nauk SSSR 99:3 l^M, 3 pis. LINNAEUS, C. 1758. Systema naturae. lOth ed. Vol. I. 824 p. MICHELOTTI, G. 1838. Specimen zoophylologiae diluvianae. Turin, 222 p., 7 pis. MILNE EDWARDS, H., and J. HAIME. 1848. Recherches sur les polypiers. Memoire 2. Monographic des Tur- binolides. Ann. Sci. Nat., Ser. 3, 9:211-344, pis. 7-10. 1849. Recherches sur les polypiers. Memoire 4. Monographic des Astreides (1). Ann. Sci. Nat., Ser. 3, 12:95-197. MOSELEY, H. N. 1873. Figs. 2-3. In W. Thomson. Notes from the "Challenger" VII. Nature (Lond.) 8:400403. 6 figs. 1876. Preliminary report to Professor Wyville Thomson, F.R.S., Director of the Civilian Scientific Staff, on the true corals dredged by H.M.S. 'Challenger' in deep water between the dates Dec. 30th, 1870, and August 31st, 1875. Proc. R. Soc. Lond. 24:544-569, 1 fig. 1881. Part III. — On the deep-sea Madreporaria. Rep. Sci. Results Voyage H.M.S. Challenger, Zool. 2:127-208, 238-248, 16 pis.. 21 figs. PALLAS. P. S. 1766. Elenchus Zoophytorum. Hagae Comitum. 451 p. POURTALES, L. F. de 1867. Contributions to the fauna of the Gulf Stream at great depths. Bull. Mus. Comp. Zool. Harv. Coll. 1:103-120. 1871. Deep-sea corals. lUustr. Cat. Mus. Comp. Zool. Harv. CoU. 4, 93 p.. 8 pU. 1878. Reports on the results of dredging, under the supervision of Alexander Agassiz. in the Gulf of Mexico, by the United States Coast Survey steamer "Blake." Lieutenant-Commander C. D. Sigsbee. U.S.N.. commanding. Corals. Bull. Mus. Comp. Zool. Harv. CoU. 5:197-212. 1 pi. SARS. G. O. 1872. On some remarkable forms of animal life from the great deeps off the Norwegian coast. I. Bri^gger & Christie, Christiania, 82 p., 6 pis. SZMANT-FROELICH. A., and M. E. Q. PILSON. 1977. Nitrogen excretion by colonies of the temperate coral Astrangia danae with and without zooxanthellae. Proc. Third Int. Coral Reef Symp. 2:417424. VERRILL, A. E. 1878. Notice of recent additions to the marine fauna of the eastern coast of North America, no. 2. Am. J. Sci. Arts. Ser. 3. 16:371-378. im. Notice of the remarkable marine fauna occupying the outer banks (Pallas)) in the north-east Atlantic, J. Mar. Biol. Assoc. U.K. offthesoutherncoastof New England, no. 11. Am. J. Sci., Ser. 3, 59:149-164. 29:149-157. , , _^ ,. WELLS J W 1979b. 'Patch' development of the deep-water coral Lophelia perlusa 1956. Scleractinia. /n R. C. Moore (editor), Treatise on invertebrate (L.) on RockaU Bank. J. Mar. Biol. Assoc. U.K. 59:165-177. paleontology. Part F, Coelenterata, p. 328-444, figs. 222-339. Univ. Kansas Press, Lawrence. ZIBROWIUS, H. , „,,,c.«K, , D 1980 Les Scleractiniaires de la Medilcrranee el de I Atlantique nord- 1979a. The distribution of the coral Lophel.a perlusa (L.) [L. proUfera onental. Mem. Inst. Oceanogr., Monaco 1 1:284 p., 107 pis. i2 SYSTEMATIC INDEX A Strang ia astreiformis 1,2,7,9,10 danae 10 Caryophyllia ambrosia ambrosia 1 , 4, 9, 10 ambrosia caribbeana 10 Caryophylliidae 10 Caryophylliina 10 Concentrotheca laevigata 1 Dasmosmilia lymani 1 , 4, 9, 10 Deltocyathus italicus 2, 4, 5, 9, 10 Dendrophyllia gaditana 1 Dendrophylliidae 11 Dendrophylliina 11 Desmophyllum cristagalli 3,9,10 eburneum 11 nobile 11 Enallopsammia ampheliodes 11 profunda 1, 2, 8, 9, 1 1 rostrata 2, 7, 9, 1 1 Faviina 10 Flabellidae 10 F la be II urn alabaslrum 6, 9, 10, 1 1 angulare 6,9,11 goodei 10,11 macandrewi 6, 9, 1 1 Fungiacyathus durus 1 fragilis 5, 9, 10 marenzelleri 1 Fungiidae 10 Fungiina 10 Javania cailleti 3, 9, 1 1 Lophelia periusa 10 prolifera 1, 9, 10, 1 1 Polymyces fragilis 1 Pourtalosmilia conferta 1 Rhizangiidae 10 Solenosmilia variabilis 8,9,10 Vaughanella margariiata 1 ACKNOWLEDGMENTS Preparation of the "Marine Flora and Fauna of the Northeastern 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, 259 High Street, Coventry, CT 06238. Arthur G. Humes, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, MA 02543. Wesley N. Tiffney, Professor Emeritus, Boston University, 226 Edge Hill Road, Sharon, MA 02067. Ruth D. Turner, Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138. Roland L. Wigley. National Marine Fisheries Service, Northeast Fisheries Center, NOAA, Woods Hole, MA 02543. Robert T. Wilce, Department of Botany, University of Massachusetts, Amherst, MA 021 16. The Board, which 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. Thanks are expressed to Duane Hope, Department of Invertebrate Zoology, Smithsonian Institution, for permitting the author to work at the National Museum of Natural History during this study. Preparation of the illustrations by Charles G. Messing is also gratefully acknowledged. COORDINATING EDITOR'S COMMENTS Publication of the "Marine Flora and Fauna of the Northeastern United States" is most timely in view of the growing universal emphasis on environmental work and the urgent need for more precise and complete identiTication of coastal organisms than has been available. It is mandatory, where possible, that organisms be iden- tified accurately to species. Accurate scientiTic names unlock the great quantities of biological information stored in libraries, obviate duplication of research already done, and often make possible predic- tion of attributes of organisms that have been inadequately studied. Stephen Cairns started his research on Scleractinia at the Rosenstiel School of Marine and Atmospheric Science, University of Miami, where he studied the deepwater corals of the western Atlantic. He has continued his studies as a Research Associate at the Smithsonian Institution, with an emphasis on western Atlantic, Hawaiian, and Antarctic Scleractinia and Antarctic Stylasterina. 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 supported by the College of Marine Studies, University of Delaware. Manuals are available from the following: Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402, at a few cents a page. User Service Branch, Library and Information Services, Division D822, Washington Science Center, Building #4, Rockville, MD 20852, at no charge as long as the supply lasts. National Technical Information Service, U.S. Department of Commerce, 5285 Port Royal Road, Springfield, VA 22161, either as paper copy or microfiche, for a charge. Manuals are not copyrighted, and so may be photocopied from the NOAA Technical Report NMFS Circulars available in most major libraries. The manuals so far published in the NOAA Technical Report NMFS Circular Series are listed below by author, title, circular number, and NTIS accession number: Marine Flora and Fauna of the Northeastern United States: Circular No. NTIS No. COOK, DAVID C, and RALPH O. BRINKHURST. Annelida: Oligochaeta. BORROR, ARTHUR C. Protozoa: Ciliophora MOUL, EDWIN T. Higher Plants of the Marine Fringe. McCLOSKEY, LAWRENCE R. Pycnogonida. MANNING, RAYMOND B. Crustacea: Stomatopoda. WILLIAMS, AUSTON B. Crustacea: Decapoda. POLLOCK, LELAND W. Tardigrada. LARSON, RONALD J. Cnidaria: Scyphozoa. CAVALIERE, A. R. Higher Fungi: Ascomycetes, Deuteromycetes, and Basidiomycetes. COULL, BRUCE C. Copepoda: Harpacticoida. CUTLER, EDWARD B. Sipuncula. PAWSON, DAVID L. Echinodermata: Holothuroidea. HO, JU-SHEY. Copepoda: Lernaeopodidae and Sphyriidae. HO, JU-SHEY. Copepoda: Cyclopoids Parasitic on Fishes. CRESSEY, ROGER F. Crustacea: Branchiura. BOVEE, EUGENE C, and THOMAS K. SAWYER. Protozoa: Sarcodina; Amoebae. WATLING, LES. Crustacea: Cumacea. ZULLO, VICTOR A. Arthropoda: Cirripedia. TODD, RUTH, and DORIS LOW. Protozoa: Sarcodina: Benthic Foraminifera BUSH, LOUISE F. Turbellaria: Acoela and Nemertodermatida. CAIRNS, STEPHEN D. Cnidaria: Scleractinia. 374 378 384 386 387 389 394 397 398 399 403 405 406 409 413 419 423 425 COM 73 50670 COM 73 50888 COM 74 50019 COM 74 50014 COM 74 50487 COM 74 51194 PB 257 987 PB 261 839 PB 268 036 PB 268 714 PB 273 062 PB 274 999 PB 280 040 PB 281 969 PB 222 923 PB 285 538 PB 296 460 PB 297 676 • GPO — 1981— 798-545 NOAA TECHNICAL REPORTS NMFS Circular and Special Scientific Report— Fisheries Guidelines for Contributors CONTENTS OF MANUSCRIPT First page. Give the title (as concise as possible) of the paper and the author's name, and footnote the author's affiliation, mailing address, and ZIP code. Contents. Contains the text headings and abbreviated figure legends and table headings. Dots should follow each entry and page numbers should be omitted. Abstract. Not to exceed one double-spaced page. Foot- notes and literature citations do not belong in the abstract. Text. See also Form of the Manuscript below. 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