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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 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 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 (Cirripedia: 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 Notacanthiformes). 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 during 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 during 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 watersupdate, 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 <it silcrucliniuii corals known IVom (he nortlu-aslcrn I niied Males coasi from \ ir^inia (o No\a SeoCia. tollowin^ a hrJet inlroduelion to (he ceiUTal l>inlot;> and ni<irphnlo};;> 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 consists 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 extremely 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) .

.

Fungiacyathusfragilis

Figure S.—tungiacyathus fragilis: CalicuUir (w of slightly damaged specimen. Scale: I cm.

7 {4} Corallum conical, compressed-conical, 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 colony; 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 present); first three cycles of septa hexamerallyarranged, 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
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 History, 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 (=Astrangia 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); SzmantFroelich 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 asexual budding from longitudinal fractures of the fragile coral- lum. Five to ten small buds originating from one wedgeshaped fragment are not uncommon. This is the most commonly 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 numbers on undersides of ledges in Lydonia Canyon, off Massachusetts. [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, inlratenlacular 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 Verrill, 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 sometimes 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 constricted, 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 continental 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-Expediion. 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 expeditions 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 Turbinolides. 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 nord1979a. 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

AStrangia 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 Fla beIIurn 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 identified 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 prediction 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.

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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

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