NOAA Technical Report NMFS 141

Marine Flora and Fauna

of the Eastern United States

Anthozoa: Acitniaria, Corallimorpharia,

Ceriantharia, and Zoanthidea

Kenneth P. Sebens

December 1998

U.S. Department of Commerce

Seattle, Washington
Foreward _

This NOAA Technical Report NMFS is part of the subseries "Marine Flora and

Fauna of the Eastern United States" (formerly "Marine Flora and Fauna of the

Northeastern United States"), which consists oforiginal, illlustrated, modern manuals

on the identification, classification, and general biology of the estuarine and

coastal marine plants and animals of the eastern United States. The manuals are

published at irregular intervals on as many taxa ofthe region as there are specialists

available to collaborate in their preparation. These manuals are intended for use by

students, biologists, biological oceanographers, informed laymen, and others wishing to identify coastal organisms for this region. They can often 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. I. Smith, and produced in 1964 under the

auspices of the Systematics Ecology Program, Marine Biological Laboratory, Woods

Hole, Massachusetts. Geographic coverage of the "Marine Flora and Fauna of the

Eastern United States" is planned to include organisms from the headwaters of

estuaries seaward to approximately the 200-m depth on the continental shelf from

Maine to Florida, but can vary somewhat with each major taxon and the interests of

callaborators. Whenever possible, representative specimens dealt with in the manuals are deposited in the reference collections of major museums.

The "Marine Flora and Fauna of the Eastern United States" is being prepared in

collaboration with systematic specialists in the United States and abroad. Each

manual is based primarily on recent and ongoing revisionary systematic research

and a fresh examination of the plants and animals. Each manual, treating a

separate major taxon, includes an introduction, glossary, uniform originally illustrated key, 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.

Manuals are available from the National Technical Information Service, United

States Department of Commerce, 5285 Port Royal Road, Springfield, VA 22161,

either as a paper copy or microfiche, for a charge. Manuals are not copyrighted,

and so can be photocopied from the NOAA Technical Report MFS Circulars and

Reports available in most libraries and listed at the end of this manual.

The 1 ational Marine Fisheries Service (NMFS) does not approve,

recommend 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 promotion which would indicate or

imply that NMFS approves, recommends or endorses any

proprietary product or proprit>tary material mentioned herein,

or which has as its purpose an intt>nt to cause directly or

indirectly the advertised product to be used or purchased

because of this NMFS publication.

ii
Note to recipients ofNOAA Technical Report NMFS 141, "Manne Flora

and Fauna of the Eastern United States. Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea, " by Kenneth P. Sebens:

Please insert this Errata sheet in all copies ofthe report.

Errata

NOAA Technical Report NMFS 141

Replace Acknowledgntents, p. 66-67, with:

Aclmowledgxnents _

Preparation ofthis manual was supported in part by a grant

from the National Science Foundation to the Editorial

Board of the "Marine Flora and Fauna of the Eastern

United States." The author thanks Frederick M. Bayer and

Steven Cairns for providing access to the U.S.N.M.

collections, Ardis Johnston for help with specimens and

curating at the Museum of Comparative Zoology, Harvard

University, Kathy Durante for nematocyst determinations,

Mel Carriker for encouragement and editing, Ronald W.

Hardy and Sharyn Matriotti for careful editing, J. C. den

Hartog and D. G. Fautin for helpful reviews of the

manuscript, and Melody Chen, Elaine Cole, Susan

Thornton, Julia Miles, Cecile Villars, Trish Dahill, Ellie

Valminuto, and Karen Vandersall for preparation of the

manuscript. Barbara Hecker (Lamont-Doherty Geological

Observatory) supplied an excellent recent collection of

anemones and photographs procured under the auspices of

the U.S. Bureau of Land Management. Ruth D. Turner

made valuable suggestions on the manuscript. Most

drawings were done by Tom Siebert from specimens at the

M.C.Z. and at the U.S.N.M. and from photographs

supplied by Barbara Hecker and by the author. Figures 1-6

were done by Sarah Jordan. Figures 10, 11, 12, 15,24,44,

and 54 are by the author.

Corrections to tables and figures:

Table 2. p. 12. Adamsia sociabilis. Shift both "SHH" and

"NJR" one column left.

Table 2. p. 13. Ceriantheopsis americanus. Move "OS" from

sphincter type column to Bottom no. column.

Table 2, p. 13. Corynactis delawarei. Add "R" below "BR" in

column 6.

Table 3, p. 14. Haloclava producta. Remove "MB 50-112"

from third column.

Table 2, p. 16. Corynactis delawarei. Move "H 69-85" left one

column.

Figure 33, p 32. Change Aiptasiogeton to Aiptasia.

Index, p. 64. Actinauge verrilli. Add p. 11, 15, 31.

Index, p. 65. Daontesia praelonga. Change p. 56 to 51.

Index, p. 65. Diadumene leucolena. Remove p. 55.

Index, p. 65. Drillactis pallida. Remove p. 56.

Index, p. 66. Hexacorallia, not italics.

Index, p. 66. Thenaria. Add p. 37.

Marine Flora and Fauna of the Eastern United States

Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea

KENNETH P. SEBENS

Department of Biology

Univmity of Maryland

College Park, Maryland 20742

and

Centerfor Environmental Science, Hom Pt. Laboratory

Cambridge, Maryland 21613

ABSTRACT

This key incl udes 60 species ofsea anemones and their relatives in the orders Actiniaria,

Corallimorpharia, Ceriantharia, and Zoanthidea. Species from the intertidal zone, continental slope, and deep sea are included over a geographic range from Atlantic Canada to

approximately South Carolina. In addition to the illustrated key itself, characteristics of

each species are summarized in tabular form, including morphology, distribution, and

types and sizes of cnidae. Ecological and taxonomic information on each species are also

included in an annotated species list.

Introduction

Anthozoans of the orders Actiniaria, Ceriantharia, and

Corallimorpharia are all commonly referred to as "sea

anemones." However, only the actiniarians are considered true sea anemones. The others are more appropriately termed "cerianthids," collectively, or "tube anemones" and "false corals", respectively. Zoanthids, sometimes called "colonial anemones" or "encrusting anemones", resemble the other three groups at the level ofthe

individual polyp but typically form connected colonies.

Corallimorpharians and, to a lesser extent actiniarians,

share their basic anatomy with scleractiniarian coral

polyps, from which they may have evolved; they are

essentially corals without skeletons. The cerianthids and

zoanthids have diverged much farther from the

actinarians, corallimorpharians, and scleractinians in

both external and internal anatomy.

Anthozoans are primarily carnivorous and are important constituents of many benthic marine communities

owing to their biomass and predatory activities. The

tropical reef corals (Scleractinia) are a good example

of an abundant and ecologically important anthozoan

group, butsea anemones and cerian thids can be equally

important; they are often the most conspicuous large

organisms in deep-sea photographs or community

samples. As a rule, anemones with short knoblike or

long thick tentacles prey on relatively large benthic

invertebrates such as sea urchins, gastropods, bivalves,

or crustaceans that crawl or swim into their grasp

(Sebens, ] 987; Shick, 1991). A strong sphincter muscle

around the margin, just below the tentacles, allows

some anemones (e.g. most Actiniidae) to cover the

retracted tentacles that hold their prey and to close the

upper column over the prey, preventing its escape.

Initial prey capture is effected by the cnidae, nematocysts and spirocysts, microscopic capsules that eject

threads and darts that either stick to or penetrate the

prey. Short thin tentacles (e.g. in Metridiidae) and the

long thin streaming tentacles (of cerianthids) are specialized for capturing zooplankton. The prey either

swim into contact with the tentacles or are carried into

the tentacle crown by water currents. Anemones with

tentacles of intermediate length and thickness (e.g.

many Actiniidae) may prey on both benthic invertebrates and zooplankton.

Sea anemones inhabit sandy, muddy, or rocky habitats and sometimes attach themselves to the hard parts

or products of other organisms, such as worm tubes,

hermit crab shells, crab claws, gastropod and bivalve

shells, gorgonian corals, and sea pens. A broad flat base

on an anemone generally indicates a hard substrate
2 NOAA Technical Report NMFS 141

marginal sphincter

acontium

marginal stoma

mesenteric filament

siphonoglyph ---'l't---II'---++++

oral disc

mesentery --------------4Jactinopharynx -------------\\----\\-,----!H

coelenteron

tentacle~

mouth --------------~§~~;;:;;;;i.f

Figure I

Anatomy of a generalized sea anemone (Aetiniaria).

origin. This type of base can also wrap around cylindrical structures such as polychaete tubes or gorgon ian

branches, and can even meet and adhere to itself on

the other side (amplexicaulous base). The sand or mud

burrowing anemones and the cerianthids have rounded

or almost pointed bases that can be used to burrow and

to anchor themselves into the substrate. These anemones have elongate, sometimes almost wormlike, bodies

capable of peristaltic burrowing movements. Cerianthids

represent the only group that consistently builds a permanent tube; only a few species of burrowing anemone

(Actiniaria) have this ability. The cerianthid tube is a

long soft black or brown construction, composed of

ptychocyst nematocysts, that collapses when a specimen

is collected. Its shiny, smooth inner surface allows a

large cerianthid to withdraw rapidly into the sediment

to depths of a meter or more, providing some protection from predators.

Morphology

An anthozoan polyp (Fig. 1) is basically a cylinder (the

column) surrounding a second cylinder (the actinopharynx), which is supported in a central position by

radial mesen teries (the mesenteries are paired in

actiniarians, zoan thids, and corallimorpharians, unpaired in cerianthids). The distal (oral) end of the

actinopharynx forms the mouth and lips. One or more

thickened ciliated grooves (siphonoglyphs) can run the

length of the actinopharynx. The siphonoglyphs pump

water into the body cavity (coelenteron) which is compartmentalized by the mesenteries and which includes

the hollow center (lumen) of each tentacle. Corallimorpharians lack siphonoglyphs, as do scleractinian corals.

Most actiniarians have two siphonoglyphs although three

or four have been observed in certain species. The

aboral end of the column is closed and forms the base,

which has an opening (pore) only in the cerianthids.

The oral end of the column gives rise to the tentacles

and the oral disc that surrounds the mouth. Tentacles

of all species are contractile, i.e. able to bend or shorten

by muscular contraction. Many species can also retract

the tentacles, covering them completely or almost so by

contracting the marginal sphincter, a circular muscle

that surrounds the margin or upper edge of the column of these anemones (Fig. 1, 2).

The column may end simply at the tentacles with no

demarcation or, in some Actiniaria, it may be divided

into recognizable regions: the scapus (lower or entire

column); the scapulus (upper column, thinner-walled

and often smooth); and the capitulum (column just

below the tentacles). The column of actiniarians can

bear distinctive sculpturing including tubercles, vesicles,
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 3

B

G

c

Figure 2

Types of marginal sphincters, redrawn from Carlgren, 1949. Endodermal: (A)

pinnate circumscribed; (B) restricted diffuse; (C) circumscribed; (D) restricted

diffuse (or palmate); and (E) diffuse. Mesogleal: (F) reticulate; (G) alveolar; (H)

transverse stratified; and (I) longitudinally stratified.

adhesive verrucae, or raised tenaculi (see definitions).

Corallimorpharians and cerianthids have smooth columns, as do many zoanthids, except for a few species

with tubercles. Zoanthid columns also incorporate sand,

sponge spicules, and other foreign material, which give

them a textured appearance.

Internally, the mesenteries are important structural

elements that also help to identify species. They are

either perfect (connected to column wall and

actinopharynx) or imperfect (not connected to

actinopharynx). Some or all mesenteries bear gonads.

The free edge of a large mesen tery bears a filamen t that

is trilobed in cross section along its upper (distal) portion and single-lobed toward the base (proximal) in the

Actiniaria and Zoantharia. In certain Actiniaria the basal

region of the mesentery forms an acontium, a threadlike structure that is free of the mesentery except at one

end. Acontia move by means of ciliary beating, muscular contraction, and hydrostatic pressure within the

coelenteron, and can be ejected through the mouth,

through pores in the body wall (cinclides), or through

temporary lesions. Their only known function is to
4 NOAA Technical Report NMFS 141

A B

E

c

F

Figure 3

Schematic mesenteric arrangements. (A) Actiniaria-most common arrangement; (B)

j;,'dwardsia, an abasilarian anemone; (C) Ceriantharia; (D) Zoanthidea; (E) Actiniaria,

Actinostola, illustrating "Actinostola rule" that third and higher cycles of mesenteries have

members of pair unequal; (F) Actiniaria, Endocoelantheae: Halcurias; new cycle of

mesenteries arises in endocoels of older pairs. a = aetinopharynx; d = directive pair; s =

siphonoglyph; m = retractor muscle.

deter predation by adhering to a predator's body a71d

stinging it with nematocysts. The mesenteric filaments

secrete digestive enzymes and digest (by phagocytosis)

pieces of prey that have been partially digested extracellularly. Cerianthids and zoanthids have mesenteric

arrangements very different from those of actinians

and corallimorpharians (Fig. 3),

The column wall, tentacles, and actinopharynx of

an thozoans consist of three tissue layers: the epidermis

(or ectoderm), the mesoglea, and the gastrodermis (or

endoderm). Mesenteries have only the latter two layers:

a central mesogleal lamella covered on both sides by

gastrodermis. The mesoglea consists of layers of collagenous fibers running in several directions. By acting

antagonistically to the hydrostatic pressure within the

coelenteron, the mesoglea enables an anthozoan to

maintain its shape. Shick (1991) has provided a thorough review ofsea anemone morphology and biology.

Cnidae: Nematocysts, Spirocysts, and Ptychocysts

The microscopic capsules (organelles) produced by

specialized cells (cnidocytes) in anthozoans are termed

cnidae. All anthozoan tissues, in all body regions, (except the mesoglea) can contain cnidae. These structures either penetrate or adhere to prey organisms.

Nematocysts are double-walled structures used for feeding, as defense against potential predators, and in competition among anthozoans. Spirocysts, the second category of cnidae found in the Hexacorallia, are generally confined to the tentacles and the oral disc but

sometimes occur in the column and base. They have a

single wall, and their threads do not penetrate prey;

instead they adhere to the surface of prey. They are

important for the capture of prey with hard shells or

exoskeletons, such as certain crustaceans, bivalves, and

gastropods. Ptychocysts, the third category, are unique

to ceriantharians; they are used to build the tube that

lines the burrow,

Nematocysts of the Actiniaria are often reliable specific characters. Their types, sizes, and distribution

among the tissues are consistent, although there is considerable intraspecific variability in sizes and in relative

numbers of types among individuals. Although these

structures are a great aid to systematists, they are not

necessary for identification of most specimens. Nematocysts can, however, be examined as a final check for

species identification. Table 1 gives the distribution for

types of cnidae among the anthozoan orders, and Table
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 5

Table I

Cnidae of several anthozoan orders, following Schmidt (1972a, 1972b) and other sources. Terminology follows Weill (1934) and

Carlgren (1949), with additions and alternative designations of Schmidt (1972a; 1972b) and Mariscal et al. (1977). Abbreviations: S =

spirocyst; A = atrich; H = holotrich; MB = microbasic b-mastigophore and basitrichs (b-rhabdoids); MP = microbasic p-mastigophore

(p-rhabdoids) types A [MPA], B [MPB], C [MPC] and D [MPD], PT = ptychocysl. ote: microbasic amastigophores are equivalent to

p-rhabdoids Band C.

Column Mesenteric

Tentacles (scapus) Actinopharynx filaments

Ceriantharia

(Cerianthidae) S,MB,A S,MB,PT,H S,MB,A MB,A

(Aracnanthidae) S,MB,MPB,A S,MB,MP,PT S,MB,MPB,A MB,MPB

Zoanthidea

(Epizoanthus and most other Zoantharia) S,MB,H S,MB,H MB,MPA,H MPA,H

(Palythoa) S,MB,H S,MB,MPA,H MB,MPA,H MB,MPA,H

Corallimorpharia (and Scleractinia) S,MB,MPD,H S,MB,MPD,H S,MB,MPD MPD,H

Actiniaria (suborder Endomyaria) S,MB S,MB S,MB,MPA S,MB,MPB,MPA

Actiniaria (other suborders) S,MB,MPB, S,MB,MPB,A S,MB,MPB, S,MB,MPA

H,MPC MPA

Antipatharia S,MB S,MB MB MB,MPA

Octocorallia MB MB MB MB

3 provides the most recen t data on spirocysts, nematocyst, and ptychocyst types and size ranges for the species in this key. Cnidae are much easier to examine and

identify in live or newly dead specimens than in preserved ones. They are small (10-150 11m long) and are

thus best observed with a high-powered compound microscope (I,OOOx, oil immersion). Phase contrast or

Nomarski interference optics are great aids in nematocyst identification because they make the small spines

and barbs on the thread much more visible.

Cnidae can be observed by making smears from each

tissue type. A scalpel or razor blade is scraped across the

surface of the column, tentacle, or actinopharynx and

is then wiped on a microscope slide. Fragments of

acontia, mesenteries, and mesenteric filaments can be

removed with fine forceps and then transferred to and

smeared on a slide. A cover slip is placed on the slide

and then pressed with a fingertip or blunt object, such

as the wooden end ofa dissecting pin. The tissue should

be smeared thin enough so that there is only a single

layer of cnidae when observed under the microscope.

In unpreserved specimens, the pressure on the cover

slip will cause many ofthe capsules to eject their threads.

Osmotic shock from a drop offreshwater introduced at

one edge of the cover slip also causes thread ejection,

as does a very dilute solution of methylene blue dye.

This dye becomes concentrated in the capsules and

may, if it becomes too dense, actually obscure the internal structure; therefore caution is needed if it is to be

used. Preserved tissues are usually stiffer and must be

macerated before they are smeared onto a slide. Investigators familiar with cnidae may wish to check these

important species-specific characters and should refer

to references by Weill (1934), Carlgren (1940; 1949),

Cutress (1955), Schmidt (1969; 1972a; 1972b), Westfall

(1965; 1966), Mariscal (1974; 1977a; 1977b; 1984) and

Conklin et al. (1977) to gain familiarity with nemotocyst

identification. Shick (1991) has reviewed nematocyst

function and toxins in sea anemones.

In addition to determining the various categories of

cnidae correctly, which can be difficult with the average

light microscope, there may be further complications.

First, there is often great inter-individual variability in

the density of nematocysts in a given tissue type. An

individual may be missing a characteristic type of cnidae

from a given tissue. Some of this variability may be due

to the location where the sample was taken, population

differences, or the presence of specific prey, competitors or predators that may induce the formation of

certain categories of cnidae. Similarly, sizes of cnidae

can vary between individuals and populations; larger

nematocysts of a given type tend to occur in larger

individuals within a species. With cnidae, small differences on the order of 10-20% of maximum or minimum capsule length should not be considered definitive for separation among species. Larger differences,

and the presence or absence of a specific category of

cnidae in several specimens, are more reliable. Several
6 NOAA Technical Report NMFS 141

Categories of Cnidae

N ematocysts

Haplonemes: Atrichs and Holotrichs-This group of

nematocysts is characterized by a tube (thread) that

lacks a basal "shaft" portion and is either loosely or very

regularly coiled within the symmetrical capsule (Fig. 4).

Tubes with a noticeable change in diameter in the section

proximal to the capsule are termed "anisorhizas," although both forms change tube diameter from the

most proximal to most distal portions. Holotrichs have

small barbs along the entire tube which atrichs appear

to lack under the light microscope. Electron microscopy has determined that atrichs actually have very

small spines along the tube .and must therefore be

considered a special form of holotrich.

Heteronemes: Basitrichs and b-MastigophoresHeteroneme nematocysts have a distinct demarcation between a thicker basal "shaft" and a thinner distal "thread"

or "tube" (Fig. 5, 6). The arrangement ofbarbs and spines

along the two portions often differs as well. In this first

group of heteronemes, the shaft has a comparatively

gradual transition to the tube, resulting in a smooth appearance of this zone in the unexploded state (Fig. 2).

Thus, no "funnel" shape is produced as in p-mastigophores

and amastigophores. Basitrichs have a thin straight shaft

and a tightly coiled tube along most of the capsule length.

b-mastigophores (Fig. 6) have a wider shaft, sometimes

with several types of barbs along it, and a loosely coiled

tube within the capsule, probably shorter than that ofthe

basitrich tube. Small basitrichs (<12 ~m), found in some

tentacles and columns of actiniarians, have often been

neglected in published descriptions. Similarly, very large

basitrichs are sometimes scattered in the tips of tentacles

Ptychocysts-Previously thought to be atrichous nematocysts, this group was recognized as a distinct category

of cnidae only recently (Mariscal et. aI., 1977). These

large structures (to over lOa ~m) are by far the most

abundant cnidae in the column of cerianthids and are

used to construct the smooth lining of their permanent

burrow. They are roughly symmetrical, about twice to

three times as long as wide, and contain an irregularly

coiled tube (Fig. 4). The tube may evert from a position

that is distinctly off center on one end of the capsule. This

tube (or thread) is not helically folded; instead it has a

variable number of pleats arranged circumferentially, but

not lengthwise. The discharged tube appears very wide,

about one fourth the capsule diameter, and varies in

diameter along its length. The tube has no spines, barbs,

or rod-shaped bodies and it shows fine ridges running

lengthwise. The tip is closed, unlike that of nematocysts.

"glutinants" in the older literature. Certain atrichous

nematocysts resemble spirocysts but have much thinner

threads, are usually symmetrical, double-walled, and

lack pronounced curvature.

a b c

Figure 4

Categories of cnidae (a) spirocyst; (b)

nematocyst, atrich (form ofhololrich); (c)

nematocyst, holotrich. (Figs. 2-4 redrawn

from Schmidt, 1972a). See also Figures 5

and 6.

Spirocysts-Spirocysts are single-walled capsules with a

tightly packed spiral tube that fills the interior (Fig. 4).

The capsule is relatively uniform in diameter, sometimes slightly wider at the proximal end and more

pointed at the distal end. There is often a pronounced

curvature to the whole capsule, especially in large

spirocysts. The tube has no barbs or spines and is thick

in relation to the thread of nematocysts. Upon firing,

the tube everts, and rod-shaped bodies (tubules) packed

inside the tube hydrate to form a long sticky filament

that wraps loosely around the tube. This filament then

breaks up to become an anastomosing network of

strands. Such cnidae correspond to the category of

specimens should be examined whenever possible given

the known variability among individuals. Nematocysts

identified as "rare" in the literature (or in Table 3 of

this key) are not likely to be appropriate for positive

identification. They may be rare for several reasons,

including contamination of one tissue surface by another during collection, dissection, or handling. Fortunately, only a small piece of tissue is necessary for study

of the cnidom, thus it is usually possible to examine

another sample from the same specimen, even if a

permanent slide has not been prepared.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 7

a b c a b c d

Figure 5

Categories of cnidae (nematocysts): (a-c)

bastrichs, three forms in unexploded and

exploded condition.

and thus may be missed in tentacle scrapings (e.g.

Urticina1). Schmidt (1972a, 1972b) hasillustrated the broad

diversity in this group and the gradation from basitrichs to

b-mastigophores; he terms both "b-rhabdoids."

Heteronemes: p-Mastigophores and Amastigophores-This group differs from the group ofbasitrichs

and b-mastigophores by having a sharp transition from

shaft to tube, which produces a distinctive "funnel"

shape in the unexploded state (Fig. 6). The tube is

rarely tightly coiled and is sometimes reduced to a

fragment or is absent. The reduced condition defines

the amastigophores; the tube in this group is never

"armed" with barbs. The p-mastigophores have a tube

at least as long as the shaft, and the tube is usually

armed. Both groups can have very long shafts, up to

several times the capsule length, that coil loosely within

the capsule. When the shaft is over three times the

capsule length, the condition is termed "macrobasic."

The more common situation, where the tube is less

than this length, is termed "microbasic." Note that both

I Hand, C. 1976. Bodega Bay Marine Laboratory, Univ. Calif.,

Bodega Bay, CA 94923. Personal commun.

Figure 6

Categories of cnidae (nematocysts): (a and b) microbasic

b-mastigophores; (c and d) microbasic p-mastigophores,

each in unexploded and exploded condition (c includes

two shapes of unexploded capsule).

types of nematocysts in this group can form terminal

"darts" composed of fused tube spines; these darts can

penetrate prey surfaces, such as crustacean exoskeletons. There is some confusion in the literature between amastigophores and b-mastigophores because

some of the former have small "funnels" that can be

hidden if the shaft is long. Schmidt (1972a; 1972b) has

suggested that the above terms be replaced by new

terms for four categories offunnel-bearing heteronemes:

p-rhabdoids (types A, B, C, D). His argument is that

neither the shaft length nor the tube length are characters that show an abrupt transition.

Specimen Preservation and Examination:

Most sea anemones contract tightly when collected. If

they are preserved in such a state, their tentacle number, column divisions, column texture, and mesenteric

arrangements are difficult or impossible to determine.

A solution of 7.5% by weight MgCI 2 or MgS04 dissolved

in distilled or tap water, then mixed with an equal
8 NOAA Technical Report NMFS 141

volume of sea water, usually relaxes them quite well.

The mixture should be kept refrigerated and the anemones left in it 12-18 hours at I-SoC (i.e. in a refrigerator

or cold room). The coelenterons of large specimens

should be flushed with the solution by means of either a

large syringe or a plastic poultry baster. Small anemones will relax in a few hours; larger ones may take

overnight to do so. A relaxed anemone does not look

like an expanded one because of the lack of hydrostatic

pressure. The marginal sphincter, if present, relaxes

and the tentacles are thus not hidden, as in contracted

specimens, and the column will also be free of strong

contraction furrows. If the animal is kept at a cold

temperature, the chance that it will begin to deteriorate, if it dies, is lessened. Tropical and subtropical

anemones often relax completely when kept in a refrigerator (at ::;SOC) even without an anaesthetic solution.

Once the anemone has been relaxed, it should be

transferred to 10% chilled formalin in seawater (i.e. 1

part 37% formaldehyde solution to 9 parts seawater;

the coelenteron should be flushed if needed) and left

to fix overnight or longer, at I_SOC, if possible. At this

point, the specimens can be stored at room temperature. After a week or more, they should be transferred

to 70% ethanol for permanent storage. If specimens

are to be used for histological study, Bouin's solution,

rather than formalin in seawater, is often recommended.

The tentacles of contracted preserved specimens can

be examined by bisecting the anemone longitudinally

(oral to aboral) through the actinopharynx. The specimen should be pinned to a wax layer in the bottom 01'1

glass dish, then immersed in water to allow internal

structures to separate. Mesenteries, acontia, tentacles,

siphonoglyphs, and sometimes the marginal sphincter

are easily observed by this method. The number of

mesenteries that reach below the actinopharynx can

also be checked, although the mesenteries in the oral

(distal) half of the specimen will not be fully visible. To

see the mesenteric arrangement, a transverse section

should be cu t just below the oral disc. The oral end of

the specimen can then be flipped over to examine ar,d

count the mesenteries. Cuts can be made cleanly, even

in small specimens, by using two pieces of a doubleedged razor blade that has been snapped in half (the

two halves are slid against each other as the cut is

made). Large anemones can be dissected with a stout

single-edged razor blade or scalpel.

Using the Key _

Given the lack of hard structures and other read' I)'

visible characters, anemones and their relatives are notoriously difficult to identify. One must spend the time

necessary to get to know their anatomy well before

proceeding with identification. This key can be used

three ways. For each method, begin by listing the obvious characteristics of the specimen, including the size,

shape, number of tentacles, type of base, and column

texture. The first method is to work from the beginning

of the key, making choices based on the description,

until further information is needed (e.g. hand sections). Once a final species determination has been

made, make certain the specimen looks like the drawing, taking into account the effects of preservation and

degree of contraction. If the identification is questionable, use Table 2 to confirm that the specimen's characteristics are within the listed ranges. Dichotomous keys

are problematic because they rely heavily on few characters and ignore some useful information. If the specimen does not match some of the characteristics noted

in Table '2, check other species close to the first selection in Table 2 and refer back to the illustrations of

these other species. For example, a wrong turn can be

made in the key because the specimen was judged to

have "prominent tubercles" when, compared with other

related species, the tubercles are really rather small.

The second method for using the guide is to dismiss

the branching key entirely and scan the pictures, listing

those species that resemble your specimen, again considering its condition. Examine Table 2 to find the

species that conforms most closely in all characteristics.

Consider habitat, depth, and geographic range, remembering that some species have not been collected frequently and known ranges are probably smaller than real

ranges. However, for shallow-water species, it would be

very unlikely to find a species known only from Maine and

the Arctic in orth Carolina, and vice versa. If a convincing match is not found by this method, return to the

branching key. The correct species may have been missed

because the specimen did not resemble the drawing.

The third method is to go directly to Table 2 and

scan the columns fOI' unique characteristics ofthe specimen or list all species with the same generic characteristics. such as depth, geographic range, base type, and

number of tentacles, Examining the drawings of these

species may help eliminate a few choices. Be careful not

to exclude a species on the basis of depth or geographic

range if, for example, the specimen was collected only

slightly outside the reported range. The correct species

call be overlooked by any of these methods if some

characteristic of the specimen is not recognized.

For example, a hand section may not show a sphincter

when a diffuse mesogleal sphincter is present. Finally,

Table;) should be used if characteristics of the cnidae

are needed to distinguish among two or more related

species. If a species has not been found that fits

the specimen's characteristics, one should go back to

each characteristic to see if there was a possibility for

misin terpretation.
(For deep water areas, the coastal point at the same latitude is given.)

Primary sources: These are the two or three publications where most of the tabulated data were found. Individual entries may be from another source (see Annotated Systematic List) or

from studies of museum specimens made by the author for this guide.

Table 2

Morphological and ecological characteristics of the species. Abbreviations are given below for entries in each column, left to right.

Maximum length, maximum width: measures (in em) are the maximum published; alive (l), or preserved (P). Width refers to column or base, whichever is appropriate.

Color: Color of tentacles (TenL), oral disc (0. disc), column, and lips including the upper portion of the actinopharynx (LIPS) of live specimens unless denoted by *blue (B), black (BK),

brown (BR), clear (Cl), cream (CR), flesh (F), green (G), gray (GR), maroon (M), orange (0), pink (P), purple (PR), red (R), salmon (S), tan (T), violet (V), white (W), yellow (Y).

Patterns in table include spots (s), lines (I), and bands around tentacles (b) next to color of that pattern.

Reproduction: Monoecious or hermaphrodite (both sexes in one individual (M)), dioecious or gonochoric (separate male and female animals (0)), (S) sexually reproducing type

unknown, brooding internally to planula larva stage (BP), brooding to tentacled juvenile stage (BJ), brooding externally to tentacled juvenile stage (BE). Asexual reproduction includes

longitudinal fission (F), pedal (basal) laceration (L), and transverse fission (TF).

Tentacles: Tentacle number (no.) is given for an average large specimen, usually close to the maximum. The second number (below) is the number of cycles of tentacles. N = nonretractile

tentacles, never fully concealed by margin. For cerianthids, tentacle number refers to marginal tentacles. l=length.

Column: Texture: Periderm on part of column (PE), thin mucus sheath with adherent particles (MS), smooth column (S), vertically striated (ST), with conical, flattened or hemispherical

tubercles (TB), with tenaculi (TE) , v.ith a sand encrusted tunic (zoanthids) or periderm (SA)), with adhesive verrucae in vertical (longitudinal) rows, at least on the top half of the column (V),

nonadhesive vesicles in vertical rows (W), small scattered nonadhesive vesicles (SY) , tightly packed nonadhesive vesicles sometimes tending to form rows near the margin (VP), vertical wrinkles

(WV), horizontal wrinkles (WR), sunken pits (PT). Note that wrinkles typically form on contracted and preserved specimens but disappear on expansion. Texture on top only (T), on bottom only

(B). Division: Column not divided, (S), column forms scapus and scapulus (SS), column forms scapus and capitulum (SC), forms carcinoecium to house hermit crab (CA).

Base type: Base folded around tubular substrata, meeting itself and often adhering to itself (amplexicaulous base, (AM)), broad adhesive base, wider than the column just above the base

(AB), small adhesive base, narrower or equal to the column just above the base (SB), broad adhesive base formed into an inverted cuplike cavity enclosing mud (CB). Physa of burrowing

forms (P). Round or bulblike base of cerianthids (R), with a pore (RP). Base with a chitinous cuticle (CU).

Number ofsiphonoglyphs: A single number is given if that is the common condition. When it is variable (0, I,2,3,4), often as a result of asexual reproduction, this condition is denoted as ·V'.

Sphincter type: No sphincter (N), alveolar (A), diffuse (D), circumscribed (C), pinnate (P), palmate (Pl), reticular (RS), transversely stratified (TS). Tissue location denoted as: mesogloeal

(M), endodermal (E).

Mesenteries: "Top" is just below oral disc. "Bottom" isjust below proximal end of actinopharynx. Numbers are pairs for a typical large specimen, usually close to maximum size. Perfect

mesenteries (P), connected to actinopharynx, total number of mesentery pairs (A), number ofsterile mesenteries in the first, second, or third order (S), number of directives (D) given

if not same as number ofsiphonoglyphs. If the boltom of the column Uust below actinopharynx) has the same number of mesenteries it is denoted 'same', otherwise only the differences

are noted. When mesenteries are not paired (cerianthids, zoanthids), number of individual mesenteries is given (I).

Substrate type: Cobble or gravel (C), gorgonians (G), hydroid stems (H), mud, silt and clay (M), solid rock or boulders (R), sand (S), shell (SH), hermit crab-occupied shell, or crab

carapace (SHH), hermit crab abdomen (SHA), sponge (SP), Sargassum plants (SR), worm tubes (W), algae (A).

Geographic range: The range is given from the northernmost point to the southernmost. If the species occurs elsewhere than the east coast of North America, the general range is given as

its northern limit

NJR

DEL

VlR

NCA

SCA

General ranges:

NHM northern hemisphere, shallow water «30 m)

ARC circumarctic boreal, shallow or deep water

EUR European artie, Scandinavia, British Isles, shallow or deep water

DSA deep water (>30 m) North and South Atlantic

DSN deep water (>30 m) North Atlantic only

LIS long Island Sound

NOV Nova Scotia, ocean side

BFD Bay of Fundy

MNE northern Maine

MNN southern Maine

New Jersey

Delaware

Virginia

North Carolina

South Carolina

lvlAS

FLA,

CCD

CAR

GRN

northern Massachusetts

Florida

Cape Cod

Caribbean

Greenland

Vl

III

c"

III

=

6"

SO

o"

;..-

::;.

5'

5;'

...

ji'

o

...

S'

o

...

"0

c"

...

F

III

... ;'

=S-

...

F

P. =

N

o

=Sis:

III

Data begins on next page.

<t:>
Table 2 (continued) o

Tentacle

S S

S S

Texture Di".

z

a::

"!j

Vl

z

""3

III

/")

:r

::: ;:;'

e:.

....

01>-

....

:;z:l

III

"0

o

;:;.

Manuel 1981

Chia et al. 1989

Verrill 1864-1866

Widersten 1976

Primary

sources

'vleMlirrich 1893, 1898

Sebens (un pub!.)

Carlgren 1921

Seben' (unpubl.)

Verrill 1922

186-!-1866

DEL

DSA

ARC

MAS

EL'R

MNE

NOV

DEL

M!\E

MAS

Geogr

range

0-100 SH,A

W

0-110

Depth Substrate

(m) type

87-117 M,S

C

700-816 R

350-2284 M

110.

14A

61'

36A

4P,A

4S

Bouom

4P,A

-1S

6P,A

~eselltery Pairs

Top no.

10P-68A

2D-OS

N

'"

'"

N

2 :'-:

;\0. of

Siphono- Sphincter

glyphs type

I'

SB

SB

AB

Base

type

SS

Column

WR

SI' S5

WR

SV,WR

PE

51'

31'

lL

0.21'

0.6L

<0.51'

I'

Length

(em)

16

70

3N

16

~o.

<120

2N

24-36

21\

D?

D

S,TF

Reproduction

W

o

R

BR

Lips

Y,W

BR W

PR 0

Colors

Y Y.P

I' BR

W W

YGR

CL,W CL.W

I' I'

O. disc. Column

BR

PR

Y,P

W,S

Tellt.

W

CL.W

I'

0.41'

41'

O.IL

101'

O.5L

0.41'

Nlax.

width

(em)

12L

4.51'

41'

7.51'

81'

O.6L

Max.

length

(em)

Edwardsia

sulcata

Edwardsia

sipunculoides

Actinernus

nobills

Gonactinia

prolifera

Halrurias

pilalus

Species

Edwardsia

elegans

15L

31'

0.5L

II'

P,CL

b

Y,BR Y 16

N

0.11' SV.PE

WR

SS I' N 4P.A 4P,A 0-117 S

C

BFD

SCA

Verrill 1922

PaTaedwardsia

aTenana

61' II' D 16

2

<0.5.1' PE,SA

TE

SC I' N 81'

16A

666-2000 DSN Carlgren 1921

Fagesia

lineata

30L

1.21'

O.oL

0.-11'

CR

W

CR

BR

CR,BR

I'

CR

BR

5

TF

-10

3:\

0.5L S SS I' N -IP,A II',A 0-23 R

W

CCD

"'CA

Verrill 18114-1866

Nematostella

vectensis

2L

II'

0.2L

0.11'

CL,W

b

CL

W

CL

W

CL

W

S

TF

9-18

2:'-:

IL S

MS

SS N 41' SAME

8A-OS

+2-0 M

C

ELR

CCD

ManllcllY81

Hand 1957

Drillaclis

pallida

10L 6L W,GR

G

W,GR

Y

W,GR Y 12-24

:-J

1.6L S

SV

SS I' N S CCD \'will 1922

Haloclava

produeta

20L

71'

3L

21'

GR,Y

CL,W

Y,GR

II'

YGR

W

20 2L VT

ST

SC !\ 101' SA.ME

lOA

+1-15 S.M

SH

CCD

SCA

l'erriIl1864-1866,1898

Widersten 1976

Hargill 1912

Peachia

parasitica

8L 1.4L P,BR

F

P,BR

F

P,BR

F

12

2

1.2L SV,WR

0.51' WV

I'

RP

:-J 6P-IOA SA.ME

6S

0-91 S

C

:vI"E

CCD

Verrill 1864-1866, 1922

Wildersten 1976

Halcampa

duodecimcirrata

3.8L

1.21'

0.5L

0.31'

S,BR

s,b

W 12

2:'-:

0.5L S

:viS

SC I' 2? N? 6P-12A SA.ME

OS

10-600 M

S

BFD

CCD

Verrill 1922

Widersten 1976

Pseudaclinia

melanaste,.

2.0L 4.0L BR.Y BR,Y

M M,W,I

BR,Y

',1

W F 200

>3

1.5L S SB D 24P-60A SAME

'S

0-' SR

R

:--JCA

CAR

Field 1949

den Hartog 1985

Bunodosoma

cavernata

5L 5L Y,G

BR.b

Y,G

BR

Y,G

BR,s

911

5

1.51' VI' AB C,E 12P-44A SAME

12S

+1-8 R "'CA

CAR

Verrill 186-1-1866

Field 1949

Carl. & Hedg. 1952

conlinued
Table 2 (continued)

Tentacle Mesentery Pairs

Species

Max.

length

(em)

Max.

width

(em) TenI.

Colors

0. disc. Column Lips

Repro.

duction No.

Column

Length

(em) Texture Oiv.

Base

type

No. of

Siphono- Sphincter

glyphs type Top no.

Bottom

no.

Depth

(m)

Substrate Geogr.

type range

Primary

sources

Bolocera

tuediae

25L 25L CR,Y

0,1'

CR,Y

0,1'

CR,Y

0,1'

R,O

M

o <260

51\

81' S

WR

SB D,E >6P-?A SA\>IE 20·2000 R

>2S SH

EUR

~CA

Carlgren 1921, 1922

\>Ian uel 1981

Epiaclis

fecunda

5.61' 6.21' BE 90 II' PT,T

WR,B

SC AB C,E 36p·44A S~\1E 25·400

OS

R

S

NWF

MAS

Verrill 1899b, 1922

6P-48A SAME 54-600

6S

481' 12p·192A 111·400

192A 2S

2S

PL 24A 48A

C,E 40-801' SA:>IE

>80A,10S fJJ

re

0-

re

=

s:-

O

N

o

?

>

::. s' ;'

...

ji'

o

...

S'

o

...

"t:I

=-

...

F

re

... ;'

=s:-

... ,F

Q. =

N

o

=s:-

o:

re

Verrill 1883, 1899b

Widersten 1976

Hand 1955b,

Widersten 1976

Carlgren 1934, 1921

1949

Verrill 1882

Carlgren 1934

Carlgren 1921

Widersten 1976

Verrill 1864-1866

Verrill 1922

Carlgren 1942

Verrill 1899b

Widersten 1976

Carlgren 1921

Widersten 1976

Carlgren 1921

Sebens (unpub!.)

Verrill 1864-1866

Carlgren 1921

Verrill 1922

ECR

I\CA

LIS

CAR

DSA

~CA

NCA

CAR

CCD

CAR

ECR

CCD

ARC

ceo

ARC

CCD

CCD

VIR

ARC

ceo

ARC

\>11\1\

R

W

\>I

M

S

R

C

R,S

SH

R

SH

R,C

SH

M,S

W

C,G

SH,R

0-'

tl-I

tl·12

10·400

t2·366

80-872

90·1296

699-4700

1239-2908

24P,A SAME

12S

16·241' 16-241'

144A 120A

16-24S

16p·34A 67A

16S

241'->48A SAME

OS

6P->160A SAME

6S

o

M

RS

D,M

O,M

A,RS

:>I

A,RS

TS,M

A,D,\>I,

SB

SB

SB

SB

AB

AB

SB

AB

AB

CB

AB

AB

S

S

SS

SC

S

VS

WR

\'

V,T

WR,B

S,WR

WV

WR,B

\V,TB

TB,WR

PE

31'

II'

4L

II'

21'

II'

2,5L

4.5L

181'

3,51'

2,5L

<21'

2L

151'

2.5L

0,31'

72

70

3~

>80

4·5

160

5

96

400

61'-:

180

<110

<160

2N

<300

6·7N

600·900

201\

o

o

BJ

S

BJ

BR

R

I'

GR

BR

R

o

R

o

S,O

BR

1',0

BR,O

F,P

GR,Y

W,BR

YO

BR

R,G

T

W

S,O

CR

W

y

BR

R

o

1',0

BR,G

GR

G,BR

F

BR,O

F,P

R

o

W

PR

1',0

R,s

BR

S,O

BR

CR,P

B,G

Y,PR

BR

BR,G

GR

G,BR

W,F,I

BR,O

F,P

GR,BR

W,I

W

PR

S,O

BR

CR,P

B,G

R

BR

1',0

b

BR

Y,O

BR,b

G,BR

W,F

BR,O

F,P

GR,W

BR

BR,GR

G,W,s

10L

25L

91'

6L

4L

31'

121'

4L

10L

51'

25L

61'

25L

II'

10.5

I'

5L

5L

8L

41'

61'

15L

51'

41'

81'

6L

20L

IUL

31'

10L

81'

Paranthus

rapiformis

Sieyonis

obesa

Lif)onema

multicornis

Aclinoseyphia

saginata

Stomphia

(ornnea

Actinostola

callosa

Urtirina

crassicornis

Aulaclinia

stella

Aulactinia

calJitata

Hannatllia

nodosa

Antholoba

perdix

A(linauge

verrillii

15L

71'

10L

51'

S,BR

I'

O,R

BR

S,F

I'

96 21' TB

pE

SS AB

CB

TS

:>I

6P-'A

6-8S

61'

6-8S

128-2840 :>I,S

W,R

GR~

DEL

Carlgren 1942

Dunn 1983

conlinued

--
Table 2 (continued) N>

Ten,acle

0-30 SH 1\CA

SHH CAR

2-1448 SHH ARC

'-'1'1:\

200-4000 SH H CCD

SHA VIR

z

o

0-3

n

:r

::s;:;0

e:.

z

>'!j

en

....

,j>.

....

:;r;l

"0

o

=- McMurrich 1893

Primary

sources

C"rlgren 1921, 1942

Widers,en 1976

Carlgren 1921

Verrill 1864-1866

MeMurrich 1898

I\OV

CCD

CCD

CAR

C

'VI

W

Depth Substrate Geogr

(m) 'ype range

200-600

238-366

Mesellter~ Pairs

M 6P->48A 48.\

6S

'VI 61'-48A SAME

6S

A 12P.99A

'VI 8-12S

TS >16P->48A 48A,6S

M 6S

TS 6P-48A SAME

M 6S

1\0. of

Siphono- Sphincter BOllom

glyphs 'ype Top no. no.

AB

CB

AB

AB

AB

AB

CC

Base

,ype

Column

TB 5S

PE

SS

V.T 5

S

WV,TB SS

WB

TeXLUre Di\'.

II'

1.4L

0.51'

0.51'

0.51'

6.5L

II'

Length

(em)

96

3

96

No.

96

200

190

4

D

Reproduction

I'

\'

o

PR

Lips

O,BR

R

R Y,BR

R

B BR

GR GR,I

Colors

S,P S,P

W.I' W,P

PR

0. disc. Column

PR

CR

S,P

TenL

I',PR

B,f,O.R

GR,VI'.b

51.

21'

91'

91.

31'

41'

51'

Max.

width

(em)

I~L

61'

31'

8L

31'

101'

61'

Max.

leng,h

(em)

Calliactis

tricolor

Allantaclis

parasilica

Paracalliactis

involvms

Phelliaclis

americana

Aclinauge

longicornis

Species

Adamsia

sociabilis

I.4L

0.51'

IL

II'

I' CL,I' CL

I

D 0.3L

0.21'

AB

CC

M 12P->300A 96A

12S m

136-738 SHA NOV

SHH

\'errill 1882. 1883

'1JR

Chondrophellia

(oranata

4.41' 21' o ° *O,GR

BR.Y

96 0.41' WV.TB

PE

SS AB 'VI

A

6P-48A 6P-<48A 600-2448 R DS'I Verrill 1883

Carlgr", 1942

Amphianlhll'

nitidus

~I' 21' R P,I'R

F

P,R

G,W

L 150

4-5

lUI' S.lB ,1M M

I)

12P)A SAME

OS

2~4-fi05 H,SI'

W.G

NOV

DEI,

Verrill 1864. 1899b

Widemell 1976

Manuel 1981

Amphianthus

mirabilis

0.41' 1.21' S.A? 96

4-5

0.31' SS AM M 6P,I~A

OS

360-620 GW 1\UI

,CA

Verrill 1~~3, IHY9b.

1922

Slephanallge

nexilis

2.51' 51' 5

BR

BR

R,O

S

F

S 202

6

0.61' S,TB SS AM A

M

6P,I72A 128A

0-2S

150-547 I',SP

VI'

.-IRC

CCD

Carlgren 1942

Widers,en 1976

Stephanauge

acanellae

1.51' 2.51' S,O

BR

S,O

P,BR

S,O

I'

S 1-10

N

1.51' S

TB

SS AM 2 A

M

6P,100A 48A 443-2430 G

OS

ARC

'IJR

Carlgren 1942

Verrill 1883

Slephanauge

spongicola

2.5L

1.21'

2.0L

1.11'

I'

F

I'

F

I'

F

S 110

4-6

0.31' S

I'E

SS AM

AB

1-4 RS 8-12P,34A SAME

M OS,2-4D

129-578 I-LSP

R,W

'10V

NCA

Verrill 1883

Widersten 1976

Sagarliogelon

venilli

4L

1.51'

3.5L

21'

O,S

F,PR

o

PR

S

F

D 96 3L

0.91'

5

TE,PE

SS A.\1

AB

A,D 12-181', 124A SAME

RS,M OS

135-1416 W,SH

R

ARC

DEL

Carlgren 1921,1942

Widers,en 1976

Aiplasia

erujJtaurantia

2.5L

II'

1.5L

0.51'

BR,G

GR

PR,BR

G,Y,I

F,P,G

R,s(R)

170 1.5L. 5 SS AB M 61'.-I8A

OS.4D

SAME 0-' R

SH

:--JCA Field 1949

Carlgren 1952

continued
Table 2 (continued)

Tentacle Meselilery Pairs

Species

Max.

length

(cm)

Max.

width

(Clll) Tenl.

Colors

O. disc. Colnmn Lips

Reproductioll :-10.

Colullln

Lenglh

(Clll) Textnre Dil'.

Rase

t"pe

No. of

Siphono- Sphincter

glyphs type Top no.

BOHom

no.

Depth

(Ill)

Substrate Geogr

type ranRe

Primaq

sources

Acti110lhoe

modesta

6.4L I.5L GR

I

II'

Y

VI'

F

100 IL S

MS

SS AB M O-? S

C

LIS

CCD

Verrill 1864-1866

Hargitl1912

Daolllesia

pmelonga

2.21' 1.31' Ll 50 0.61' TE

CL'

SS AB

CB,CL'

A

M

121',48A SA:-1E 1041-1960 M

OS

EUR

DEL

Carlgren 1942

70 0.-11'

>201.

71-120

i\

150-200 15L

:-< 51'

SO

N

o

;I;-

:i";'

...

F'

C"'l

o

...

S'

o

...

'"0

::r

...

F'

C"'l

... ;'

:l

S-

...

ji'

:l

eN

o

:l

Sc:

[FJ

r:1'

:l

Widerslell 1976

Shepard CI al. 1986

lIand 1956

Widerslell 1976

Shepard et al. 1986

Species A

Verrill 1873b. 1922

Widersten 1976

Shepard <t al. 1986

species B

Shepard el al. 1986

Specie> C

Shepard et al. 1986

Species 0

Hand 1956

Hecker (unpubl.)

Verrill 1864. 1883

Verrill 1882

Manuel 1981

Carlgren 1952

den Hartog 1985

Carlgren 1942

I-land 1956

\-Ianuel 1981

DSt\

MAS

SC.\

DSN

DS:-I

DSt\

DEL

DSt\

DEL

DSt\

DEL

DSN

ARC

\;JR

CCD

FLA

:-ICA

CAR

:\HM

CAR

R

M

M

R

R.SI'

A.SH

0-70

0-' R

SH

2-234

+3-16

>1000

+1-22

57-7i4 SHH

101'

200-300 M

200-300 M

454-1152 SHH

1600-1900 M

lOA 0-500 M

S.C

921 921

OS

61'->96,1 >48,1

OS

\;

'\

N

l'

N

N 3-91'.>96A >48,1

0-6S

M 5-61'.24A SAME

40

N

M 3-151'.>190A >96,1

A 0-15S 61'.6S

N

OS

1-3

AB 0-3

AB

AB

RI'

RI'

AB

RP'

RI',

RI',

RP'

SS

SC

SC

SC

CA.S

CA.S

SA

S

S

S

S

11'\'

S

101'\-

S.I'\'

<1L

2L

0.41'

3L

0.31'

1.51'

IL

0.51'

96

i\

>48

100

<40

\;

<200

6

>1000

D

F

VI

D

L

D

L

\-1

D,L

CR

BR

I'

G,W

II'

BR.Y

1\'

BR

b

BR

F

CI<

BR

GR

IlR

PK

BK

BR.G

I'.GR

W,CR

O.BR

I'

CR.II'

C

BR

BR

Y

G

CR

BR

GR

BI<

I'

11'.1

W.CR

O,BR

BK

R

C.I'

I'R

BK

°

S

C

CL

BR.S

W.b

II'

PI<.,

II'

Cl..G

W.CR

O.BR

BR.II'

PR

CR.W

Bits

51'

2L

21'

15L

81'

2L

IP

>2L

1.21'

2.5L

0.61'

0.8L

0.81'

3L

II'

2~1'

61'

3L

4L

30L

101'

36L

201'

1.51'

2.01'

>15L

C(lr;fl ntit us

borpnli~

Epizonnl.ll1ts

inrruslatus

lWel1idium

senile

Haliplanelln

luciae

Aiptasia

pallida

(.'Pl'ial1lltid /I

(undescribed)

Viadumenf

lftLrolfl10

Cninnthpopsis

nmf',.;rfl1l10

I:.pizoflntlm~

!Hlguriphil",

C'l'inl1lhid I

(undescribed)

C",-;al1lhid [II

(undescribed)

C",.;(wlhid /\'

(ulldescribed)

COI)'f/(I(II\

ddawal'f,j

0.81' II' BR 90 0.5 AB 61'-60A >200 R_II'

SH

DSN Widersten 1976

"'"
14 NOAA Technical Report NMFS 141

Table 3

Nematocyst and spirocyst size ranges and distributions in anthozoan tissues, compiled from several sources. Only lengths

(llm) are presented, as ranges from smallest to largest obseryed 10 that tissue. No information is available on the

nematocysts of a few species. Abbreviations are as follol'.s: * rare nematocyst type in that particular tissue (mayor may not

be contamination from other tissue surfaces), ± size I ange unknown, CL whole column or distal part, SP scapus, SC

scapulus, S spirocysts. Nematocysts are categorized as follows: B, basitrich; H, holotrich (includes atrichs); MA, microbasic

or macrobasic amastigophore; MB, microbasic b-mastigophore; MP, microbasic p-mastigophore; MU, microbasic

mastigophore of undetermined type; PT, ptychocyst; , any nematocyst where type was not specified. Some authors do not

report abundance of cnidac. Small basitrichs (<15 llm) have been omitted by some authors. Species for which no data are

available are omilled from this table.

Species

Iialruria.)

pilalll'

Arlinrrnus

nobili,

Edwflrdsin

sipunruloidfs

Edwardsia

elegans

Telltacles

B 18-43

S 16-48

B 22-29

S 67

:\ :\8-61

B 19-26

S 10-25

B 22+'

S 5-15*

Column

B 16-45*

S 17-44

B 24-38

S 31-48

:\ 25-36

SP/MB 42-71,90-110

SC/B 14-16

CLiB 37-87,10+*

ML' 62-97*

-\clinopharynx

B 18-42

MP 17-32*

S 1/-42*

B 34-41

S 67

B 16-26

MP 24±*

B 15-32

MP 17-37

:vIB 22-32*

Mesent.

filaments

B 26-42*

MP 18-36

S 22-33*

B 27-43

:VIP 22-42*

S 25-49*

B 19-24

MP 21-33,23-32

lype 2

B 17-2Y

MP 16-25

:vIB 20-22*

Other

tissues

t.conlia

MP 19-35

B 26-39*

S 19-46*

Sources

Sebens (lInpllb1.)

(5 spec.)

Carlgren, 1921

this study (mes.) (5 specs.)

Widerslell,1976

Sebens (lInpllb1.)

(4 spec.)

~ematosomes

,Vfmalostl'lla

uft(fnsis

IlalorlflIlfl

/Jrodllrla

Pearhia

!J{lmsilira

Halrampa duoderimrirrala

PSl'udarlinia

melilnast"r

Bunodosoma

ravernala

Bolorera

fuediae

Ej,iarli,

f,cundn

L'rtirina

rrassirornis

Aulartinia

'I,lla

Aulflrlinia

rapilata

Siryoni'

abesa

Pamnlhus

rnpiformis

B 10-13,20-25

S 11-21

B 11-20

MB 50-112

S nOlle

B 27-39

S 23±*

B 12-20*

S 14-25

B 16-34

B 16-20

S ?

B 21-37,52-87

S 31-74

B 25-29

S 27-38

B 10-14,20-37

S 17-71

B 17-31

S 13-31

B 12-26.35-45

S 9-27

N 37-49

S 70

B 22-25

:viP 22-27

S 15-26

B 10-13

B 20-25

MB 50-112

B 27-34

SP/B 22* SC,S 25-49

S 28+*

B 16-21

H 34-40

B 13-21

H lX-23,34-47

B 16-21,33-63

B 16-24

B 6-37,79-83

S 22-69

B 17-31

S 27±*

B 10-39

:VIP 17-24*

S 12-23*

24-28

B 16-26

MP 16±*

S 33-60

B 14-18

Ml' IX-25

B 14-17.38-57

S 43±*

B 40-46

S 19-23

MP 24-32

S 17-25

B 16-28

MP 1X-20

B 12-23

MP 17·19

B 50·79

MP 23-33

B 33-39

S 29-39*

B 12-26,49-109

YIP 23-30

S 28-41 *

B 24-38

B 16-38

:viP 14-20

S 12-211*

N 26-29

34-36

B 25-30

\1P 19-30

B 14-18

Ml' 18-25

B 14-25,55-75

B 70-83

B 27-45

MP 28±*

B 12-14

MP 22-23

B 16-37

MP 18-23

B 13-J 6,24-35

MP 18-20

B 20-22,50-74

MP 20-35

B 26+*

S 30+*

B 11-34,49-91

MP 20-64

B 18-44,9±*

MP 25-'16

S 15±*

B 16-41

MP 15-20

S 12-20*

B 22-32

\1P 13-15

20-26

S 49-57

B 14-17

ML' 17-22

Hand. 1955

Widersten, 1976

Sebens (lInpub1.)

(lent.,5 spec.)

Widersten, 1976

Widersten, 1976

Carlgren & Hedg., 1952

Carlg-ren & Hedg., 1952

Widersten,1976

Sebens (lInpub1.)

(l spec.)

Widerslen, 1976

Sebens (lInpub1.) (3 spec.)

Carlgren, 1921

Sebens (unpub1.)

(5 spec.)

Sebens (unpub1.)

(5 spec.)

Carlgren. 1934

Widersten, 1976

Carlgren & Hedg., 1952

continued on nexl page
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 15

Table 3 (continued)

ACLino- Mesent. Other

Species Tentacles Column pharynx filaments tissues Sources

AClinoseyphia B 19-38 B 17-35,45-77 B 22-36 B 26-31 Carlgrell, 1934

saginala S 68±* S 21-60 Sebells (ullpub!.) (3 spec.)

LijJonema B 22-60 B 19-28 B 31-47 B 18-44 Carlgrell,1921

Inulticomis N 96-192 S 25-37 Sebells (ullpub!.) (1 spec.)

S 22-60

Aclinoslola B 26-41 B 19-35 B 21-32 Widerstell, 1976

callosa \1B 43-56 S 25-57 MP 22-28 MP 20-29

S 27-64

Siomphia B 14-25 B 12-20 B 14-23 B 10-22 Widerslell,1976

coccinea MB 31-53 31-38 MP 18-27 MP 17-29 Carlgrell,1921

S 19-50 S 22-57

Antholoba B 15-16, 23-36 B 20-28 B 14-27 B 14-28 Widerslell. 1976

IJeTdix S 19-46 MP 15-31 \1P 11-31

Acolllia

Honnathia B 17-34 SP/B 8-11 B 16-35 B 14-16 B 32-40 Wirlerstell,1976

nodosa S 23-56 21-24 MP 23-33 28-31 Carlgrell, 1942

S 17-25 MP 21-23

Acolltia

AClinauge B 23-36 SP/B 11-13 B 14-19,26-42 B 13-18 B 14-22 Carl~rell, 1942

uel1illii S 560 MP 24-28 MP 21-36 MP 24-31 31-43

SC/B lJ-18

Acolllia

Aclinauge B 19-27 SC/B 16-21 B 19-31 B 11±* S 28-35 Carlgrell, 1942

longicornij S 536 MP 21±* MP 17-20

Acolltia

Phelliaclis B 34-43,17-21 * SP/B 25-44,14+* B 37-42,16+* B 12-22,33-48 B 16-23 Widerslell,1976

americana S 38-75 S 27-60 \1P 30-39 MP 28-34 32-52

Paracalliarlis B 13-42 B 18-27 B 21-42 B 18-41,9-10' Sebells (ullpub!.)

involuens S 16-41 :vIA 12-22 MP 19-23* MP 14-24* (5 spec.)

B 8* MA 546* S 13-24*

ACOtltia

Allanlaclis B 26-40 B 24-34 B 26-42 B 13-17 B 16-23 Carlgrell, 1942

parasitica MP 21-34 31-44

S 560

Acolllia

Calliaclis B 21-24 B 12-14 B 21-27 B 11-13 B 21-27 Carlgrell & Hedg. 1952

tricolor MP 19-24

Adamsia B 8-19 B 12-22 B 10-15* B 8-14* Sebells (ullpub!.)

sociabilis S 9-15* S 12-18' MP 9-22* MP 10-14 (5 spec.)

MP 12-13* MP 13±* MB 14-19* MB 12-18

ACOlllia

Cho1ldrophellia B 22-32 SC/B 18-23 B 22-34 B 16-18 B 30-53 Carlgrell,1942

coronata S 19-50 MP 26-29

Acolllia

Amphianthus B 19-30 B 8-13,29-35 B 24-25 B 9±* B 42-57 Widerslell, 1976

nilidus MP 19-29 MP 17-27 MP 23-26 MP 22-28 14±* Carl~rell 1934

S 19-47 S 29-61 MP 27±*

Acontia

Stephanauge B 9-20 SP/B 12±* B /5-18 B 14-28 B 31-37 Widerslell, 1976

nexilis MP 17-23 MP 12-20 MP 17-24 MP 16-27 MP 43-93

S 25-49 Carlgrell, 1942

Slephanauge B 12-16, S 36-41 B 13-14 B 16-20 B 10-16,20-23 Carlgrell. 1942

acanellae MP 20-21 MP 17-24 MP 18-23 MP 17-21

Acontia

Stephanauge B 11-33 B 21-27,31-39 B 14-32 B 12-15 B 13-16,33-45 Widerslell, 1976

Spo1lgicola H 39-49* H 19-20,39-45 MP 17-26 MP 13-26

S 17-34

conlinurd on nexl page
16 NOAA Technical Report NMFS 141

Table 3 (continued)

Actino- Mesenl. Olher

Speci,.s Tentacles Column pharynx filaments tissues Sources

Acomia

Sagartiogt"on B 13-29 SP/B 9-12,16-22 B 24-32 B 27±* B 37-43 Widersten, 1976

unrilli MA 16-44 MA 15±*,24-35 MP 23-31 MP 26-31 MA 57-64

S 22-36 S 22-34 Carlgren, 1942

Acoillia

Aiplasi" B 20-26* B 13-20 B 18-20* B 16-17 B 3J-40 Carlgren, 1952

fl1iptauml/iia MA 30-44 YlA 24-29 MP 14-16 MP 14-17.32-49 MP 59-73

S 15± MA 21-31 MU 21-31

Acolllia

Daonlesifl B 26-34 C/B J8-22 B H-31 B 12-17*.21-28* B 45-58 Carlgren,1942

pradon/!,fI S 24-60 MP 24-'\4 MP 21-31

Acolllia

1\Jll'lridiuUl B 12-31 B 10-30 B 17-43 B 11-21 B 8-14 Carlgren, 1942

5pnill' YlA 16-36 MP 23-31 YIP 17-53 MP 8-48 MA 2H-Y2 Hand, 1955b

MB 12-32 MA 11-35 MB 26-39 MA 17-22 MB 32-80

S 12-30 S 1J-27 MA 31 t' S ~29

AcoIltia

Aiptasin B 13-20 (3 types) B 10-20 (3 lypes) B 9-18 (2 l"pes) B 10-17* B 17-35 (2 types) Carlgren &

pallida MP 24-35 (3 types) MP 17-25 (2 types) MA J7-31 (3 types) MP 10-18 (2 lypes) MP 4H-75 (3 types) Hedg., 1952

30-40

Acontia

Hnlipltlllrllll B 11-20 B 9-17,19-23 B 12-32 B 9-19 B 14-21 Hand,1955b

luriaf MP 19-27 MP/MA 10-24 MP 19-27 MP 9-28 MP 29-56 Widersten, 1976

MA 10-15 MP/MA 21-23 MA 17-28 MA 13-20

S 14-23 S 13-17'

Acolllia

Dindumfm' B 13-22 B 9-14 B 15-~4 B 8-13 B 11-20 Hand,19:;:;b

lpurolmfl MA 19-31 MA 13-22 MA 22-31 MP H-31 MP 22-52

S 10-23

Epizonnllllu MB 22-23. H 22-2-1 H 22-24 MP 22±* YlB 11-21 Widerslen, 1976

inrruslt,lll.\ MP 22-33. II 34-40 II 21-25.38-41 H 23-26

MP 20-28

F.pizonlllhlls H 17-30' H 20-31* MB 16-26 B 19-22* Sebens (unpubl.)

l)flgl",p"ilus II 14-29* YlB 17-23* (3 spec.)

H 18-32*

MP 26±*

Marginal Labial Tenl.

CFria"t!lU!i MB 21-61 H 22-44*,61+* MB 29-54 YlB 17-57 MB 18-64 Widersten. 1970

bOffa/is H/PT 10-36 S 22-60' s 2'1-13 H/PT 55-64* Sebens (unpubl.) (2 spec.)

S 20-56 S 18-54

Marginal Labial Tent.

Cnlflflt/ll'opJis MB 16-19 MB 20-33*,S* MB 14-18,21-23 yIB 19-21.26-33 :VIB 16-33 Widerslen, 1976

(ll/leriranlls H/PT 9-13 II/PT 29-36.59-75 MB 21-32 H.·PT ~1±* S 14-27 Ylariseal et aI., 1977

S 12-27

Aerospheres

COI)"lflrlis MB 19-24 H 38-53 MP 22±*. H 12±* \'IP 21-44 MB 31-34,b9-72 Widersten, 1976

delnware; MP 36±* S 19-27 H 28-46 H 31-35.37-78 MP 33-82

H 15±* S 17-23 S 31+* H 69-85

S 19-38
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 17

Identification Key _

Class: Anthozoa

Subclass: Hexacorallia (Zooantharia)

I Colonial form, or with bases of at least some polyps connected 112

I ot colonial form (some species in aggregations); column without sand or other foreign material

actually embedded in it, although such material may adhere to the outside III

11(1) Column rough and rigid due to sand and other foreign material embedded in it; tentacles without

acrospheres; one siphonoglyph; sphincter present order Zoanthidea

11(1) Column smooth without embedded sand; tentacles with acrospheres; siphonoglyph absent; sphincter

weak or absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. order Corallimorpharia

111(1) Tentacles of two distinct kinds and cycles (oral and marginal); one siphonoglyph; inhabiting permanent

tube; mesenteries unpaired order Ceriantharia

111(1) Tentacles of one or more kinds and cycles; permanent tube usually absent3 ; mesenteries paired

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. order Actiniaria

Order Zoanthidea

1 Inhabits deep water (60-700 m), encrusts shells or rocks; polyps small (to 1.5 cm tall, 0.6 cm wide

contracted) Epizoanthus incrustatus

Figure 7

Epizoanthus incrustatus. Side view of

colony, contracted (preserved) specimen. Scale: 1 em.

2 Many Corallimorpharia, and a few Actiniaria, have polyps connected only temporarily following fission and are otherwise separate, not

colonial.

3 One or more species of burrowing Actiniaria produce a permanent tube.
18 NOAA Technical Report NMFS 141

1 Inhabits deep water (500-1,100 m), encrusts shells with hermit crabs; polyps large (to 2.0 em tall,

1.2 em wide contracted) . . . . . . . . . . . . . . . . . . . . . . . . . . .. Epizoanthus paguriphilus

- Figure 8

Epizoanthus pagu·riphilus. Dorsal view, contracted (preserved)

specimen, redrawn from Verrill (1883). Scale: 1 em.

Order Corallimorpharia

1 Single polyp or attached polyps in groups of two or more; acrospheres present; tentacles retractile;

inhabits deep water (>200 m) Corynactis delawarei

Figure 9

Corynactis delawarei. Connected

polyps on shell fragment, contracted specimen. Scale: 1 em.

Order Ceriantharia

1 Polyp with ::;80 marginal tentacles (depth ~1,500 m) 2

1 Polyp with >80 marginal tentacles (depth ::;500 m) 3
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 19

2(1) Polyp with :::;50 tentacles in large specimens; tube opening flush with, orjust below, substrate

.................................................................... cerianthid 1 (undescribed)

Figure 10

Cerianthid I. Expanded tentacle crown, from in situ photograph hy B.

Hecker (Lamont-Doherty Geol. Obs., NY). Scale: 5 em.

2(1) Polyp with >50 tentacles in large specimen; tube protrudes several cm above substrate

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. cerianthid II (undescribed)

Figure 11

Cerianthid II. Expanded tentacle crown, [rom In situ photograph (Shepard et aI., 1986). Scale: 5 em.
20 NOAA Technical Report NMFS 141

3(1) Polyp with marginal tentacles of equal length in uniplanar array; tube opening up to 20 cm above

substrate (depth ~200 m) cerianthid III (undescribed)

3(1) Marginal tentacles of unequal length, in multiJ::lanar array 4

Figure 12

Cerianthid III. Expanded tentacle crown, from in situ

photograph by B. Hecker (see Fig. 10). Scale: 5 em.

4(3) Tentacles greenish yellow; tube Hush with substrate; tentacle number unknown, but may be >80

(depth 200-400 m; no photographs available) cerianthid IV (undescribed)

4(3) Tentacles dark colored or banded; tube flush or above substrate (depth 0-500 m). . . . . . . . . . . . . . . . . . .. 5
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 21

5(4) Marginal tentacles number to ] 25 in large specimens, unbanded; spirocysts of marginal tentacles 27 )..lm

length; Cape Cod south to Caribbean (depth .,;70 m) . . . . . . . . . . . . . . . . . . . . . . .. Ceriantheopsis americanus

I

Figure 13

Ceriantheopsis americanus. (A) Contracted (preserved) specimen. (B) Expanded

tentacle crown, redrawn from Field (1949). Scale: 5 cm.

5(4) Marginal tentacles number 150-200, banded; spirocysts of marginal tentacles to 56)..lm length; Cape

Hatteras north to Arctic (depth 0-500 m) Cerianthus borealis

I

Figure 14

Cerianthus borealis. (A) Contracted (preserved) specimen. (B) Expanded tentacle crown,

from in situ photograph by B. Hecker. (see Fig. 10) Scale: 5 cm.
22 NOAA Technical Report NMFS 141

Order Actiniaria

1 Polyp with rounded or pointed nonadhesive base (not disc-like); one siphonoglyph 2

1 Polyp with flattened or cuplike adhesive base; one or more siphonoglyphs . . . . . . . . . . . . . . . . . . . . . . . .. 11

Suborder Nynantheae

Tribe Athenaria: one siphonoglyph; no adherent base; burrowing in mud, sand, or gravel

2(1) Scapus with hard sand-encrusted cuticle; also with tenaculi Paraedwardsia arenaria

2(1) Scapus without hard sand-encrusted cuticle; with or without tenaculi 3

Figure 15

Pamedwarsia arenaria. Expanded specimen with sandencrusted scapus. Scale: 3 em.

3(2) Tentacle number ~20 4

3(2) Tentacle number <20 7
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 23

4(3) Two distinct body regions (scapus, scapulus); scapus with periderm Fagesia lineata

4(3) Single body region 5

A

Figure 16

Fagesia lineata. (A) Aggregation of five contracted specimens. Scale: 1 em. (B)

Expanded tentacle crown and scapulus. Scale: 1 em.

5(4) Tentacle number 20; 20 rows of papillae; adhesive on column. . . . . . . . . . . . .. Haloclava producta

5(4) Tentacle number >20 6

B

Figure 17

Haloclava producta. (A) Expanded tentacle crown, redrawn from Hargitt

(1914). Scale: 5 mm. (B) Partically contracted specimen. Scale: 5 mm.
24 NOAA Technical Report NMFS 141

6(5) Tentacle number 20-36; body >5 cm long when preserved; column with periderm..Edwardsia sipunculoides

A

\

B

-

Figure 18

f:dwaTdsia sipunculoides. (A) Contracted specimen. Scale: 5 mm. (B) Expanded tentacle crown. Scale: 5 I:lm.

6(5) Tentacle number 12-24; column smooth; periderm absent; body <5 em long preserved

................................................ Drillactis pallida or Halcampoides sp. (Field, 1949)

Figure 19

Drillactis pallida. Expanded individual, redrawn

from Verrill (1922). Scale: 5 mm.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 25

7(3) Column with papillae (tenaculi); tentacle number 8-12 Halcampa duodecimcirrata

7(3) Column without papillae, tentacle number ~12 8

A

B

Figure 20

Halcampa duodecimcin-ata. (A) Expanded individual. Scale: 5 mm. (B) Partially contracted specimen, same scale, both redrawn from Verrill (1922).

8(7) Tentacle number 12; cinclides present; body <3 cm long contracted; mouth with canchula

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peachia parasitica

8(7) Tentacle number ~12; column smooth; cinclides absent; conchula absent. 9

B

Figure 21

Peachia parasitica. (A) Contracted specimen. Scale: 5 m. (B) Expanded tentacle crown,

redrawn from Verrill (1922). Scale: 1 em.
26 NOAA Technical Report NMFS 141

9(8) Tentacle number 9-18; column smooth; no periderm, but with some loosely adherent material or thin

sheath on scapus; tentacles usually banded; nematosomes (40 mm diameter) present in coelenteron

............................................................. , . . . . . . . . . .. Nematostella vectensis

9(8) Tentacle number 16; periderm on scapus; nematosomes absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10

- Figure 22

Nematostella vectensis. Expanded individual, drawn

from life. Scale: 1 mm.

10(9) Tentacle number 16; 8 longitudinal grooves on column; periderm on scapus Edwardsia elegans

I

Figure 23

lidwardsia elegans. Expanded individual, redrawn from

Verrill (1922). Scale: 1 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 27

10(9) Tentacle number 16; 12 body ridges; periderm on scapus Edwardsia sulcata

Figure 24

F:dwardsia sulca/a. Expanded individual. Scale: 1 em.

11(1) Acontiapresent 12

11(1) Acontia absent 31

Tribe Thenaria: Families with Acontia Present

12(11)' Base attached to hermit crab shell or forms enclosure around hermit crab abdomen ., . . . . . . . . . . . . .. 13

12(11) Base not attached to hermit crab shell or enclosing hermit crab abdomen. . . . . . . . . . . . . . . . . . . . . . .. 16

13(12) Base forms enclosure around hermit crab abdomen (taking the place ofa shell); base with cuticle .... 14

13(12) Base attached to existing shell inhabited by hermit crab; base without cuticle. . . . . . . . . . . . . . . . . . . . .. 15
28 NOAA Technical Report NMFS 141

14(13) Column folded around hermit crab abdomen to form a conical base; tentacles oriented downward;

:::::300 mesen tery pairs; moderate depths «250 m) Adamsia sociabilis

Figure 25

Adamsia sociabilis. (A) Partially contracted specimen on hermit crab

abdomen, tentacles downward. (8) Dorsal view. Scale: 1 em.

14(13) Column not folded to form a cone; deeply concave base forms cavity for hermit crab abdomen;

tentacles oriented laterally; :::::50 mesentery pairs; deep water (200-4,000 m) Paracalliactis involvens

Figure 26

Paracalliactis involvens. Contracted (preserved) specimen. Scale: 1 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 29

15(13) Tentacle number :'0:96; north Atlantic south of Cape Hatteras in deep water (:2:1,450 m)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Allantactis parasitica

Figure 27

Allantactis parasitica. Contracted (preserved) specimen. Scale: 1 em.

15(13) Tentacle number :'0:200; south of Cape Cod in shallow water (:'0:10 m) Calliactis tricolor

-

B

Figure 28

Calliactis tricolor. (A) Contracted specimen, base. (B) Contracted specimen, side. (C) Expanded

tentacle crown. Scale for all: 1 em.

16(12) Column wall thick (:2:4 mm at mid-column on large specimen); column tubercles present . . . . . . . . . .. 17

16(12) Column wall thin «4 mm at mid-column on large specimen); column tubercles absent 20
30 NOAA Technical Report NMFS 141

17(16) Pedal disc not cuplike; tubercles in rows Hormathia nodosa

17(16) Pedal disc cuplike (strongly concave); scapulus distinct and ridged on contracted specimens '.. 18

B

Figure 29

Horrnathia nodosa. (A) Contracted specimen with moderately developed tubercles.

Scale: 1 em. (B) Contracted specimen with well-developed tubercles. Scale: 1 em.

18(17) Scapus with rounded tubercles in up to 48 vertical rows; tentacles number up to 190. . .. Phelliactis americana

18(17) Tubercles irregularly arranged on scapus; tentacles number <100 19

Figure 30

Phelliactis americana. Contracted (preserved) specimen.

Scale: 1 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 31

19(18) Tubercles low and irregularly arranged on scapus Actinauge longicornis

c

Figure 31

Actinauge longicomis. (A) Contracted specimen. Scale: 5 em. (B) Expanded individual, from B. Hecker photo (see Fig. 10). Scale:

5 em. (C) Section through margin showing tentacles. Scale: 1 em.

19(18) Tubercules on scapus prominent, covering most of column Actinuage verrilli

B

- Figure 32

Actinauge verrilli. (A) Contracted specimen. Scale: 5 em. (B) Expanded individual, from B. Hecker photo (see Fig. 10). Scale: 5

em. (C) Section through margin showing tentacles and sphincter.
32 NOAA Technical Report NMFS 141

20(16) Mid-column with vesicles or with cinclides having raised rims 21

20(16) Mid-column without vesicles or cinclides with raised rims . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 22

21(20) Tentacle number ~64; cinclides with raised edges in 10-12 rows of 2-5 per row; red spots mark

cinclides when alive , , . . . . . . . . . . . . . . . . . . . . .. Aiptasia eruptaurantia

B

Figure 33

Aiptasiogeton eruptaurantia. (A) Expanded individual and (B) contracted specimen, both redrawn from Field (1949), Scale: 1 em.

21(20) Tentacle number 60-100; small vesicles irregularly arranged on column; distinctive double-lined

pattern on tentacles Actinothoe modesta

., """'J~'. • ..,I.~" #',

,.::~~,~,,:

" ..~.:.. '

B

Figure 34

Actinothoe modesta. (A) Expanded individual. Scale: 1 em. (B)

Single tentacle showing lined pattern.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 33

22(20) Column smooth, without periderm or vesicles; base not amplexicaulous . . . . . . . . . . . . . . . . . . . . . . . . .. 23

22(~0) Column with periderm or with weak vesicles at margin; base amplexicaulous or not amplexicaulous. .. 27

23(22) Inner tentacles less than twice as long as outer ones; shallow water (usually <100 m); margin plain or

sculptured; acontia often numerous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24

23(22) Inner tentacles more than twice as long as outer ones; deep water (>100 m); no sculpturing at margin;

few acontia (occurs on sponges, rocks, hydroids) Stephanauge spongicola

Figure 35

Stephanauge spongicola. (A) Contracted specimen and (B) expanded specimen.

Scale: 1 em.

24(23) Tentacle number to ~1,000; oral disc lobed in large individuals (note distinct cnidom of acontia,

Table 3) Metridium senile

24(23) Tentacle number <100; oral disc not lobed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25

c

I I

.'1,./,'

-

Figure 36

Metridium senile. (A) Expanded (living) individual. Scale: 5 em. (B) Expanded juvenile. Scale: 1 em. (C) Partially contracted

specimen. Scale: 1 em.
34 NOAA Technical Report NMFS 141

25(24) Cinclides inconspicuous, scattered; marginal sphincter not distinct; ~96 mesentery pairs; vertical

orange, yellow, or white stripes (rarely absent) on green or brown column when alive (note distinct

cnidom of acontia, Table 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Haliplanella luciae

25(27) Cinc1ides conspicuous, in vertical or horizontal rows; distinct marginal sphincter present or absent;

~96 mesentery pairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 26

Figure 37

Haliplanella luciae. (A) Contracted and (B) expanded individual. Scale: 5 mm.

26(25) Cinclides in 1-2 bands at mid-column; marginal sphincter weak to moderately strong; ~24 mesentery

pairs; column and tentacles uniform cream to brown (note distinct cnidom of acontia, Table 3)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aiptasia pallida

Figure 38

Aiptasia pallida. Expanded (living) individual.

Scale: 1 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 35

26(25) Cinclides in distinct vertical rows on scapus; distinct marginal sphincter absent; paired white lines

mark mesenteric insertions on column; ~96 mesentery pairs; one or more white tentacles almost

twice as long as others may be present (note distinct cnidom of acontia, Table 2)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diadumene leucolena

Figure 39

Diadumene leucolena. Expanded (living) individual. Scale: 5 mm.

27(22) Hard tubercles in two rows of 12 each at margin, forming crown on contraction; lower column with

dark periderm; base not amplexicaulous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Chondrophellia coronata

27(22) Tubercles at margin weak if present (>12), not hardened; lower column with or without periderm .... 29

B

Figure 40

Chondrophellia coronata. Top (A) and side (B)

views of contracted specimen. Scale: 1 em.
36 NOAA Technical Report NMFS 141

29(27) Margin without sculpturing; acontia numerous; inner tentacles about twice the length of outer ones;

column with periderm; base attached to worm tubes (Hyalinoecia anifex); base not fully amplexicaulous

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Sagartiogeton verrilli

29(27) Margin with some sculpturing; acontia present but not numerous; inner tentacle length twice that of

outer ones; column without periderm; base attached to pennatulid or gorgonian axis. . . . . . . . . . . . . .. 30

-

B

Figure 41

Sagartiogeton verrilli. (A) Expanded individual on worm tube, redrawn from Verrill (1883). (B) Expanded

specimen. (C) Contracted specimen. Scale on all: 1 em.

30(29) Low tubercles in one row on ridges at margin; base attached to pennatulid (Balticinae) denuded axis;

base amplexicaulous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Stephanauge nexilis

Figure 42

Stephanauge nexilis. Contracted specimen. Scale:

1 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 37

30(29) Vesicles weak, in one row at margin; acontia poorly developed or absent; base attached to gorgonian

(Acanella arbuscula) axis: base amplexicaulous Stephanauge acanellae

Figure 43

Stephanauge acanellae. Partially contracted specimen. Scale: 1 em.

31(11) New mesenteries arise in endocoels of existing pairs, so musculature appears reversed from normal

plan (longitudinal muscles of directives face apart, those of other pairs face each other, Fig. 3)

(suborder Endocoelantheae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 33

31(11) Mesenteric musculature follows normal plan; new mesenteries arise in exocoels 32

32(31) Column and tentacles structurally similar, without demarcation; eight perfect mesenteries present;

two directive pairs present; only one member of each non directive pair perfect (suborder Protantheae)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Gonactinia prolifera

32(31) Column and tentacles differ in structure, with clear demarcation; four or more perfect mesenteries

present; one or two directive pairs present, both of which are perfect except after fission (suborder

Nynantheae, tribe Thenaria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34

Figure 44

Gonactinia prolifera. Individual undergoing transverse

fission, from photo by author. Scale: 1 mm.
38 NOAA Technical Report NMFS 141

33(32) Single siphonoglyph present; more mesenteries present just below oral disc than in rest of column:

sphincter absent; column 3-4 times as tall as wide; oral disc somewhat wider than pedal disc

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halcurias pilatus

-

Figure 45

Halcu'fias pilatus. (A) Contracted specimen; note collar raised above tentacle bases.

(B) Expanded individual, from B. Hecker photo (see Fig. 10). Scale: 1 em.

33(32) Two siphonoglyphs present; sphincter absent; equal number of mesenteries proximally and distally;

mesenteries after first 20 pairs have members of pair unequal; column twice as wide as tall; oral disc

lobed much wider than pedal disc. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Actinernus nobilis

A

c

Figure 46

Actinernus nobilis. Top (A) and side (B) views 0:: contracted specimen. (C) Expanded individual, from

B. Hecker photo (see Fig 10). Scale: 5 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 39

Tribe Thenaria: Acontia Absent

34(32) Entire column, or lower half, smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 35

34(32) Verrucae obvious; tubercles or tenaculi on column. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 44

35(34) Column elongate, more than twice as tall as wide; tentacle number :s:180; fine horizontal grooves on

column; burrowing habit Paranthus rapifarmis

35(34) Column less than twice as tall as wide; tentacle number variable; not burrowing. . . . . . . . . . . . . . . . . . .. 36

A

-

8 c

Figure 47

Paranthus rapiformis. Two conformations of preserved specimens (A and B). Scale: 1 em. (C) Expanded

tentacle crown redrawn from Verrill (1899a). Scale: 1 em.

36(35) Actinostola rule applies to mesenteric arrangement (see definitions and Fig. 3) . . . . . . . . . . . . . . . . . . . .. 37

36(35) Other mesenteric arrangement applies 40

37(36) Column wall very thick (>5 mm at mid-column in large specimen); tentacles cannot be fully retracted ..38

37(36) Tentacles can be fully retracted; column wall thin «3 mm at mid-column in large specimen); pedal

disc wide; tentacle number :S:96, in 2-3 cycles Stomphia coccinea

B

A

- Figure 48

Stomphia coccinea. Two conformations of preserved specimens (A and B). Scale: 1 em. (C) Expanded tentacle crown redrawn

from Verrill (l899a). Scale: 1 em.
40 NOAA Technical Report NMFS 141

38(37) Upper column with large square tubercles; (column twice as tall [or greater] as wide, preserved)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Actinostola callosa

38(37) Column without tubercles; column about twice as wide as tall, preserved. . . . . . . . . . . . . . . . . . . . . . . . .. 39

A

B

Figure 49

Actinostola callosa. (A) Contracted specimen. Scale: 5 em. (B) Partially expanded oral disc, redrawn from

Verrill, 1883. Scale: 5 em. (C) Section through margin; note tentacle size and length increases toward

mouth. Scale: I em.

39(38) Tentacles at margin wide and conical, ~60 in number . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Sicyonis obesa

Figure 50

Sicyonis obesa. Partially con tracted (preserved) specimen. Scale: 5 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 41

39(38) Tentacles at margin fine, filamentous, ::;160 in number. . . . . . . . . . . . . . . . . . . . . . .. Actinoscyphia saginata

Figure 51

Actinoscyphia saginata. (A) Expanded individual. Scale: 5 em. (B) Contracted (preserved)

specimen. Scale: 5 em.

A

40 (36) Column flattened; oral disc much wider than base; tentacles easily detached (deciduous); basal

sphincters cover the convex oral disc . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Liponema multicornis

40 (36) Column height less than or equal to width, preserved 41

A

Figure 52

Liponema multicornis. (A) Specimen with tentacles shed. Scale: 5 em. (B) Expanded (living)

specimen, redrawn from Verrill (1922), same scale.

41 (40) Tentacles less than half as long as column, preserved. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 42

41 (40) Tentacles as long as column or longer, preserved 43
42 NOAA Technical Report NMFS 141

42 (40) Anemone small «3 cm diameter base); base attached to sponges, gorgonians (e.g. Paramuricea,

Primnoa) or hydroid stems; folds at parapet; acontia evident Amphianthus nitidus

Note: a similar small anemone «4 cm diameter base) with a row of weak vesicles at the margin, with acontia poorly

developed or absent, is found attached to the gorgon ian Acanella (Stephanauge acanellae, see line 30 in key, Fig. 43)

Figure 53

Amphianlhus nilidus. (A) contracted specimen on gorgonian branch. Scale: 1 em. (B) Expanded

individual. Scale: ] em.

42(40) Anemone small «3 cm diameter base); base attached to gorgonians (Paragorgia, Paramuricea);

acontia reduced or absent; cinclides absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Amphianthus mirabilis

Figure 54

Amphianlhus mirabilis. Contrac.ted specimens on gorgonian branch.

Scale 1 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 43

43(41) Anemone large (to 5 cm diameter base); tentacle number :<:;260; tentacles with basal sphincters;

tentacles easily detached (deciduous) and longitudinally ridged Bolocera tuediae

8 A

Figure 55

Bolocera tuediae. (A) Expanded tentacle crown. (B) Contracted

specimen. (C) Expanded individual, from B. Hecker photo (see

Fig. 10). Scale in all: 5 em.

43(41) Anemone small (to 3 cm diameter base); tentacles as long as column; tentacles without basal

sphincters, not deciduous or longitudinally ridged Anemonia sargassensis

B

Figure 56

Anemonia sargassensis. (A) Expanded individual on Sargassum, redrawn from Hargitt

(1914). Scale: 1 em. (B) Partially contracted specimen. Scale: 1 em.
44 NOAA Technical Report NMFS 141

44(34) Scapus covered by tenaculi; scapulus smooth; deep water (>1,000 m) Daontesia praelonga

44(34) Verrucae or vesicles present, but tenaculi absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 45

Figure 57

Daontesia praelonga. Contracted specimen

with damaged base. Scale: 1 em.

45(44) Adhesive verrucae present and scattered, or in longitudinal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 46

45(44) Vesicles or pits present, covering most of column. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

46(45) Vesicles small, scattered, nonadhesive; column twice as tall as wide; tentacles number ::;160 in 5 cycles;

mesenteric arrangement decamerous; 10-20 oldest mesenteries sterile. . . . . . . . . . . .. Urticina crassicornis

46 (45) Verrucae distinct and adhesive (some may appear as pitlike cavities) scattered on column or in rows .. 47

A

.'~

~: .. ,

Figure 58

Urticina crassicornis. (A) contracted specimen. (B) Expanded individual. Scale: 5 em.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 45

47(46) Verrucae present and adhesive; tentacle number 48-96; verrucae color usually contrasts with column

color when alive Aulactinia stella

Figure 59

Aulactinia stella. (A) Contracted specimen with adhered shell gravel. (B) Tentacle crown of expanded individual. Scale: 1 em.

47(46) Verrucae adhesive and in distinct rows on upper third of column only Aulactinia capitata

-

B

Figure 60

Aulactinia capitata. (A) Partially expanded oral disc of preserved specimen. (B) Contracted specimen. Scale: 1 em.
46 NOAA Technical Report NMFS 141

48(45) Nonadhesive vesicles cover column 49

48(45) Pits on column low and flat; column twice as wide as tall (preserved); tentacles number ~90

......................................................................... " Epiactisfecunda

A

Figure 61

l:-piaclis Jecunda. (A) Contracted specimen showing brood pits on column. (B) Partially

contracted specimen, both redrawn from Verrill (1922). Scale: 1 em.

49(47) Acrorhagi in fosse below outer tentacles; tentacle number :0::96; oral disc as wide as base; column

covered by small vesicles or papillae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Bunodosoma cavernata

A

. " '., , ,~ J :.

-

B

Figure 62

Bunodosoma cavernata. (A) Expanded individual. (B) Contracted (preserved) specimen. Scale: 1 em
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 47

49 (47) Acrorhagi absent; tentacle number S;400; oral disc twice as wide as base; column covered by small

vesicles or papillae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Antholoba perdix

A

B

c

Figure 63

Antholoba perdix. Side (A) and top (B) views of contracted expanded specimen. Scale: 5 em.
48 NOAA Technical Report NMFS 141

Table 4

Species that may be encountered at the borders of the area covered by this key. The most recent complete description is

given at right: (see Carlg-ren, 1949, for synonymies and generic characteristics for each species).

Species bordering this area to the north (Greenland, Davis Straits, Canada);

Acthelmis intestinalis (Fabricius, 1780)

Actinostola groenlandica Carlgren, 1899

Amphianthus margaritaccus Danielssen, 1890

Amphianthus venniwlatus Carlgren 1942

Anthosactisjan-mayeni (Dan ielssen, 1890)

Boloura maxima Carlgren, 1921

Bunotlactis spetsbergensis Carlgren, 1921

C:actosoma abyssO'Tum Danielssen, 1890

Cerianthus sp. llCfJdi Gosse, lR59

CribrinojJsis similis Carlgren, 1921

f:dw(ll'dsia andresi Danielssen, 1890

Epiactis no1'timanni Carlgren, 1921

t:'JJizoanthus danielseni Carlgren, IY13

t:pizoanthus erdmanni Danielssen, 1890

t:pizoanthus glacialis Danielsson, 1890

Epizoanthus lindahli Carl~ren, 1913

Halcampoides purpurea Danielssen, 1890

Haliactis arctica Carlgren, 1934

Honnathia digitata (Mlliler, 1776)

lsozoanthus bulbosus Carlgren, 1913

lsozoanthus davisi Carlgren, 1913

lsozoanthus ingolfi Carlgren, 1913

Kadosactis rosea Danielssen, 1890

Phelliactis rolntsta Carlgren, 1928

Parasiryonis groenlandica Carlgren, 1933

Ptychodactis patula Appelliiff, 1893 (order Ptychodactiaria)

Pycnan.thus laevis Carlgren, 1921

Sicyonis tuberculata Carlgren, 1921

Siryonis ingolfi Carlgren, 1921

Tealidium jungerseni Carlgren, 1921

(Carlgrl"n,1942)

(Carlgren, 1921)

(Carlgren, 1942)

(Carlgren, 1942)

(Carlgren, 1921,1933)

(Carlgren, 1921)

(Carlgren, 1921)

(Carigren,1Y21)

(Carlgren, 1942)

(Carlgren, 1921)

(Carlgrl"n, 1921)

(Carlgrl"n, 1933)

(Carlgren, 1933)

(Carlgrl"n, 1933)

(CarlgTl"n,1933)

(Carlgrl"n, 1933)

(Carlgren, 1921)

(Carlgrcn, 1934)

(Carlgren, 1933, 1942)

(Carl~ren, 1933)

(Carlgren, 1Y33)

(Carlgren,1933)

(Carlgren, 1933, 1942)

(Carigren,lY42)

(Carlgren, 1933)

(Carlgren, 1921, 1934, 1942)

(Carlgrl"n, 1921)

(Carlgren, 1921)

(Carlgrl"n,1921)

(Carl~ren, 1921)

Species bordering this area on the south; (South Carolina, Geor~ia, florida, Bermuda).

Bunodosorna granuliferum (Lesueur, 1817)

Cond)'lactis gigantea (Weinland, 1860)

Bunodeopsis antilliensis Duerden, 1897

Bunodeopsis globuliJem (Verrill, 1899b)

Actinia bermudensis (McMurrich, 1889)

Phymanthus crucifer (Lesueur, 1817)

Aiptasia tagetes (Duchassaing and Michelotti, 1866)

ActinostellafloswliJem (Lesueur, 1817)

Bal'tholomea annulata (Lesueur, 1817)

Anthoplrum catenulata (Verrill, 1900)

Anthopleum cameola (Verrill, 1907)

Anthopleura varia-armata Watzl, 1922

Corynactis pamula Duchassaing & Michelotti, 1861

Species bordering this area to the east, in the central

Bathyphellia margaritacea Dan ielssen, 1890

Corallimorphus rigidus Moseley, 1877

C:orallimorphus ingens Gravier, 1918

Amphianthus michaelsarsi Carlgren, 1934

Phelliactis hertwigii Simon, 1892

Actinauge abyssorum Carlgren, 1934

Paracalliactis michaelsarsi Carlgren, 1918

Stephanauge inomata Gravier, 1918

(Correa, 1964, den Hartog, 1985)

(McMurrich, 1889, Verrill, 1907, Correa, 1964,Jennison,

1981, den Hartog, 1985)

(den Harto~ 1985)

(Carlgren 1950, Correa 1964)

(Verrill 1898a, 1907, Correa 1964, den Hartog, 1Y85)

(den Hartog, 1985;Jennison, 1981)

(Verrill, 1907; Carlgren and Hedgepeth, 1952)

(Verrill 1907, dl"n Hartog 1985)

(Verrill 1907, Correa 1964,Jennison 1983, den Hartog

19H5)

(Vl"rrill, 1907, den Hartog, 1985)

(den Hartog, 1985)

(Carlgren, 1950; den Hartog, 1985)

(den Hartog, 1980)

orth Atlantic (>1,000 m depth);

(Carlgren, 1942)

(Carlgren, lY28)

(Carlgren, 1934)

(Carlgren, 1934)

(Carlgren, 1928, 1942)

(Carlgren, 1975)

(Doumenc,1975)

(Carlgren 1942)
Annotated Systematic List

Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 49

Family Actinernidae

This classification scheme follows that ofCarlgren (1949),

the most recent complete compilation of genera and

species of the orders Actiniaria, Ptychodactiaria, and

Corallimorpharia. Full synonymies of most species listed

here appear in Carlgren (1949) and in the more recent

studies listed; only the most commonly encountered synonyms and the most recent complete descriptions of each

species and its biology are given in this account.

Order ACTINIARIA

Suborder Protantheae

Family Gonactiniidae

Gonactinia prolifera (Sars, 1850, p. 142) (see also

Stephenson, 1935, p. 25; Carlgren, 1949, p. 18; Manuel,

1981, p. 92)

This small anemone has been reported from scattered

locations in the Gulf of Maine: Cashes Ledge, Eastport,

Maine (Sebens), New Brunswick, Canada (Logan et aI.,

1984), low intertidal to 100 m. This is a very small

anemone «8 mm tall, expanded) with 16 or fewer

tentacles and no marginal sphincter. Tentacles and

column are translucent pink, cream, or white. Asexual

reproduction occurs by transverse fission; new tentacles

arise part way down the column. Two or three individuals

can remain attached for indefinite periods. This anemone

can swim by tentacle movement and can walk by alternating tentacle and base contact with the substrate. Reproduction has been described by Chia et al. (1989).

Suborder Endocoelantheae

Family Halcuriidae

Halcurias pilatus McMurrich, 1893, p. 142 (see also

McMurrich, 1898, p. 227; McMurrich, 1901, p. 155)

This is a tall slender white anemone with brilliant orange lips and actinopharynx, and a single siphonoglyph.

It occurs in large aggregations on vertical cliffs and on

boulders at 160-307 m depth along submarine canyons

off Maryland and Delaware where it was recently photographed and collected by B. Hecker. It was previously

known only from the South Atlantic (McMurrich, 1893).

A distinctive feature is that the second and third cycles

of incomplete mesenteries (microcnemes) end just below the oral disc. New mesentery cycles occur in

endocoels of the first cycle, and there are no basilar

muscles in anemones ofthis suborder. The coelenteron

(gastrodermal) side ofthe actinopharynx is often bright

orange even in preserved specimens.

Actinernus nobilisVerrill, 1879, p. 474 (see also Carlgren,

1918, p. 32; Carlgren, 1921, p. 14)

This is a relatively large, short and wide anemone with

two siphonoglyphs and a very thick-walled white column formed into [our large lobes at the oral disc.

Purple tentacles begin wide at the margins oflobes and

become narrow distally to form thin filaments. This

species is found on rock and shells in deep water (350-

2884 m) from Delaware north to at least Nova Scotia.

New mesenteries arise in the endocoels ofthe first cycle

and there are no basilar muscles in anemones of this

suborder.

Suborder Nynantheae

Tribe Athenaria

Family Edwardsiidae

Edwardsia sipunculoides Stimpson, 1853, p. 7 (see also

Verrill, 1922, p. 131; Carlgren, 1931, p. 22; Williams,

1981, p. 346)

A deepwater (87-117 m) elongate burrowing anemone

found in Alaska, in the arctic, and south to Massachusetts on the east coast; this species has 24-36 tentacles

and eight prominent longitudinal ridges on the column. Nemathybomes are present except in the physa.

E. sipunculoides is similar in form to E. sulcata, but bmastigophores of its nemathybomes and column are

much larger, and there are more tentacles in large

specimens.

Edwardsia sulcata Verrill, 1864-1866, p. 29 (see also

Widersten, 1976, p. 860; Williams, 1981, p. 346)

This elongate and very large burrowing anemone has

15 tentacles and 12 prominent longitudinal ridges on

the column. It is found from the shallow subtidal zone

to 110 m and is known from two specimens collected at

Chelsea and Gloucester, Massachusetts, (in USNM collection) and six specimens from Mt. Desert Island, Maine

(Widersten, 1976). Nemathybomes are numerous except in the physa and there is a strong yellow-brown

periderm which is easily removed.

Edwardsia elegans Verrill, 1869, p. 162 (see also Verrill,

1922, p. 129; Carlgren, 1931, p. 17; Williams, 1981, p. 346)

A very large elongate burrowing anemone with 16 tentacles and eight prominent longitudinal ridges on the

column, this species occurs in the low intertidal to 117

m depth from the Bay of Fundy south to Cape Cod.

Nemathybomes are present except in the physa.
50 NOAA Technical Report NMFS 141

ParaedwardsiaarenariaCarlgren, 1921, p. 69 (in Carlgren,

1949) (see also Carlgrcn, 1921, p. 69)

This is a relatively large burrowing anemone from very

deep water in northern Europe and along the east coast

of North America south to Cape Hatteras (one specimen, 2,100 m, ident. by K. Sebens, unpub!.). Sand

grains completely encrust the cuticle (periderm).

Nemathybomes are absent, the scapus develops tenaculi,

and there are more mesenteries in the upper than in

the lower half of the column.

Fagesia lineata Verrill, 1874, p. 379 (=Edwardsia leidyi

Mark, 1884, in Verrill, 1898a, p. 496)

This species is found from Northern Massachusetts south

to North Carolina and adults form low intertidal to

shallow subtidal (to 23 m) tight aggregations by means

of transverse fission (Crowell and Oates, 1980). Larvae

andjuveniles are parasitic on the ctenophore Mnemiopsis

leidyi (Verrill, ] 898a; Crowell, 1976). There are up to 40

t.entacles; physa and nemathybomes are absent, and a

thick cuticle covers the scapus.

Nematostella vectensis Stephenson, 1935, p. 44 (=Nematostella pellucida Crowell, 1946, p. 58 (see also Hand,

1967, p. 411; Williams, 1976, p. 51, p. 257; Williams,

1979,p.69)

This very small brackish-water, estuarine and salt marsh

species (Williams, 1975b; Williams, 1976) is found in

northern Europe and as far south as Cape Cod on the

Atlantic coast of orth America. It is colorless except

for numerous white bands around the 16 tentacles.

Nematosomes in the coelenteron are unique to this

genus. emathybomes are absent. Asexual reproduction is by transverse fission.

Drillactis pallida (Verrill, 1879, p. 198), = Edwardsia

pallida, notE. pallidaCarigren, 1921; see Williams, 1981,

p. 351 (see also Verrill, 1922, p. 133, Drillactis)

This low intertidal sand-burrowing species is known

only from Verrill's (1879) incompletely described specimens taken at Provincetown, Massachusetts. It has a

long, slender, smooth column, 24 tentacles, with the

inner ones shorter than the outer, and mesenteries

divisible into macrocnemes and microcnemes. The aboral end is tapered, there are two siphonoglyphs, and

nemathybomes are absent. A specimen recently collected in South Carolina appears to be this species. The

anemone described by Field (1949) as Halcampoides sp.

may also be this species, although her specimens had

only] 2-16 tentacles.

Family Haloclavidae

Haloclava producta Stimpson, 1856, p. llO (see also

Verrill, 1864-1866, p. 30; Carlgren, 1921, p. 107, =

Halcarnpa albida Verrill, 1864-1866, p. 29; Widersten,

1976, p. 863)

This elongate sand-burrowing anemone has 20 tentacles, broadest at their tips, and 20 prominent rows of

raised adhesive verrucae on the distal two thirds of the

column. It is found from the low intertidal to shallow

subtidal (15 m). [t has no marginal sphincter, a single

siphonoglyph, and is reported to be nocturnally active.

Mesarmaea laevis and M. chloropsis (Verrill, ]864-1866,

p. 27 [Ilyanthus]).

These are incompletely described species, possibly rare,

and are not included in Carlgren's (1949, p. 31) discussion of the genus. Alesacrnaea chloropsis was collected

from South Carolina, M. laevis from Eastport, Maine.

Both species have elongate columns, tapering toward

the narrow base, which may have a terminal pore and

have up t.o 36 ten tacles of moderate length. The margin

is crenulated with rounded projections. Mesacmaea laevis

reaches 2.5 cm in length preserved; M. chloropsis reaches

R cm when expanded. These species are not included

in the key because of their uncertain status.

Peachia parasitica (Agassiz, 1859, p. 23, Bicidium) (see

also Verrill, 1922, p. 124 (Siphonactinia); = Bicidiopsis

anica, Verrill, 1922, p. 127; = B. tubicola, Verrill, ]922, p.

126; Widersten; 1976; p. 863)

This species occurs from Cape Cod to northern Maine

and probably farther north. It is a burrowing anemone

wit.h eight to 18 (usually 12) tent.acles and 10-20 (often

12) ridges on the column, only twice as long as wide

when preserved. It is found in gravel from the low

intertidal to 91 m depth in silt clay. The prominent

raised conchula on the oral disc is a good means of

identification. Juveniles and small adults are parasitic

on the scyphyzoan medusa Cyanea anica.

Family Halcampidae

Halcampa duodecimcirrata (Sars, 1851, p. 142, Edwardsia).

(Verrill, 1869, p. 162, Edwardsia farinacea, see also

Carlgren, 1921, p. 119; Verrill, 1922, p. 122; Widersten,

1976, p. 862)

This elongate burrowing anemone has 12 conical

banded tentacles with no obvious column ridges, but

with a periderm. It has been found in mud from 10
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 51

to150 m depth, from Cape Cod north to at least the Bay

of Fundy. A specimen collected recently from 600 m

was identified by the author. Prominent tenaculi on the

column are often red even on preserved specimens.

Family Actiniidae

Pseudactinia melanaster (Verrill, 1901, p. 51; Verrill, 1907,

p.257) (=Anemonia elegans Verrill, 1901, p. 50; Verrill,

1907, p. 261; = A. sargassensis Hargitt, 1908, p. 117; = A.

antilliensis Pax, 1924, p. 99 (see also Field, 1949, p. 8;

den Hartog, 1985, p. 175)

This small anemone is found throughout the Caribbean and north to North Carolina, intertidally and on

floating Sargassum plants, and is often cryptically colored. The 30-200 relatively long tentacles are not retractile. The column is smooth and has light colored

acrorhagi on the parapet. There are no siphonoglyphs

or directive mesenteries, and asexual reproduction occurs by longitudinal fission.

Bunodosoma cavernata (Bose, 1802, p. 22, Actinia) (see

also McMurrich, 1887 (Phymactis), p. 61; Verrill, 1899a

(Bunodosoma) , p. 45; Carlgren and Hedgepeth, 1952,

p.149)

This is a stout, low intertidal to shallow subtidal Caribbean anemone, found north to North Carolina, with

close-packed nonadhesive vesicles covering the column

and acrorhagi at the parapet. There are up to 96 tentacles of moderate length and two siphonoglyphs. It

may be confused with B. granuliferum (Lesueur, 1817)

farther south (but see McCommas and Lester, 1980).

Bolocera tuediae (Johnston, 1832, p. 163, Actinia) (see

also Carlgren, 1921 p. 141; Stephenson, 1918, p. 112,

116; Carlgren, 1891, p. 241; = B. occidua McMurrich,

1893, p. 154; = B. longicornis, Stephenson, 1935, p. 130;

Widersten, 1976, p. 863; Manuel, 1981, p. 104)

This widely distributed species is found from North

Carolina to the arctic and northern Europe, from 20-

1,000 m depth, rarely to 2,000 m. It is also found in the

southwest Atlantic (subspecies occidua, RiemannZurneck, 1980,60-545 m depth). The column is completely smooth, hidden by the tentacles when expanded.

Column and tentacle color is uniform orange, cream,

or pink, sometimes with inner tentacles a darker color.

Tentacles often drop off during collection and on preserved specimens.

},piactisfecunda (Verrill, 1899b, p. 378, Epigonactis). (Verrill,

1922, p. 11; = E. regularis, Verrill, 1899b, p. 380)

This is a small stout deepwater anemone (25-360 m)

from the northwest Atlantic, collected from Massachusetts to Newfoundland, with obvious external pits in the

column in which embryos, larvae, and juveniles are

brooded. The tentacles are stout and number at least

90, and the column has prominent tubercles on the

upper half. This species may be synonymous with },piactis

marsupialis Carlgren, 1901 (p. 482), from the Pacific

arctic region.

Urticina crassicornis (Johnston, 1847 p. 226, Actinia

crassicornis) (see also Carlgren, 1921, p. 168 (Urticina) ,

Stephenson, 1935, p. 139; Hand, 1955, p. 72 (Tealia); =

Rhodactinia davisiiAgassiz, 1847, p. 677; Verrill, 1883, p.

18; = Urticina eques (Gosse, 1860, p. 351, Bolocera; see

Manuel, 1981, p. 109)

This large circumarctic-boreal anemone is found on

mid-intertidal rock and on hard substrata to >300 m

depth, from just below Cape Cod to the Arctic. The

small scattered verrucae are not adhesive. It broods

juveniles to the 12-24 tentacle stage. The morphology

and nematocyst distributions are equivalent to U.

crassicornis on the west coast of North America (Hand,

1955). This species is probably also equivalent to Urticina

eques (= Tealia felina var. lofotensis Stephenson, 1935) in

northern Europe. Manuel (1981) determined the genus name Urticina Ehrenberg, 1834, to have priority

over the more commonly used Tealia Gosse, 1858.

Aulactiniastella (Verrill, 1864-1866, p. 16, Bunodes) (see

also Verrill, 1899a, p. 43 (Bunodactis); Verrill, 1922 p.

429 (Tealiopsis); Carlgren, 1921, p. 148; Carlgren, 1942,

p. 74 (Cribrina)

This small intertidal and shallow subtidal anemone

broods itsjuveniles internally and occurs from Maine to

Greenland on rocky shores. Dunn, et. al. (1980) referred this species, certain other Bunodactisspecies, and

a newly described west coast (Pacific) species (Aulactinia

incubans) to the genus Aulactinia.

Aulactinia capitata Verrill, 1864-1866, p. 20 (see also

Dunn et al., 1980, p. 2078)

This southern east coast (Atlantic) and Gulf of Mexico

species occurs from low intertidal to shallow subtidal

habitats, buried to the tentacles in sand. This species is

usually somewhat elongate with a very narrow lower

column and wide upper column when preserved. The

upper half to third of the column has prominent vertical rows of tightly packed adhesive verrucae. The lower

half to two thirds lacks verrucae and forms deep horizontal fissures in preserved specimens. The first verrucae below the tentacles are trilobed.
52 NOAA Technical Report NMFS 141

Family Actinostolidae

Sicyonis obesa (Verrill in Carlgren, 1934, p. 7, Aetinemus

obesus) (see also Carlgren, 1949, p. 81)

Known only from specimens collected off Chesapeake

Bay and Martha's Vineyard, this is a large very deepwater

(>1,200 m) species. One specimen studied by Verrill

was placed in the USNM as the type, but Carlgren

(1934) gives the first full description of the species,

listing Verrill as the author and noting that Actinemus

obesa seems to have been a "museum name" and was

never published. Possibly the largest anemone found

along this coast (to >12 em basal diameter, preserved);

the thick body wall, short thick tentacles, and smooth

vertically flattened column are distinctive.

Paranthus rapiformis (Lesueur, 1817, p. 171, Aetinia)

(see also: Verrill, 1899b, p. 213 (Arnmophilaetis); Carlgren,

1934, p. 17; Widersten, 1976, p. 868)

This is a small to moderate-size intertidal to shallow

subtidal sand-dwelling anemone from the southern east

coast and the Gulf of Mexico. Specimens often swell to

spherical or goblet-shaped forms with the base much

smaller than column diameter and remain that way

when preserved. Newly collected and inflated preserved

specimens resemble peeled onions.

Aetinoscyphia saginata (Verrill, 1882, p. 225, Aetinemus

saginatus) (see also Stephenson, 1918, p. 128;

Stephenson, 1920, p. 541 (Aetinoscyphia); Carlgren, 1934,

p.8).

An arctic-boreal, deepwater (::=:700 m) North Atlantic

species found from the arctic to N. Carolina, this species has threadlike tentacles and a vertically flattened

smooth column, usually with a small base. The base

may enfold worm tubes or sponge (Hyalonema sp.) spicule bundles. Ecological observations and photographs

of a similar species (A. aurelia) are given by Aldred et al.

(1979) .

Liponema multicornis (Verrill, 1879, p. 198, Boloeera) (see

also Verrill, 1922, p. 117 (Euboloeera); Carlgren, 1921, p.

143 (Boloeera); Carlgren, 1928, p. 148 [Liponema]).

This flattened deepwater (>80 m) North Atlantic

anemone is found as far south as Cape Cod, has a

convex oral disc and is completely covered with several

hundred short papilliform tentacles. The base is much

smaller than the oral disc and is hidden by it in life.

Aetinostola callosa (Verrill, 1882, p. 224, 315, Urtieina;

(see also Carlgren, 1891, p. 227 (Aetinostola); =

Catadiomene atrostorna Stephenson, 1918, p. 118; Riemann-Zurneck, 1971, p. 169; Widersten, 1976, p. 865)

This large deepwater (>90 m) circumarctic-boreal species is found in large numbers as far south as Cape Cod.

It is taller than wide, with a very thick tubercled column

and short, thick, striated tentacles beginning at the

margin, increasing in length toward the mouth. The

concave base is as wide or wider than the upper column, sometimes enfolding a mud bolus. The column

tubercles are thick and flattened, most prominent on

the upper third of the column.

Stomphia coeeinea (Muller, 1776, p. 231, Aetinia) (see

also Carlgren, 1921, p. 234 (Stomphia); Carlgren, 1942,

p. 75; Verrill, 1922, p. 1189; Siebert, 1973, p. 363-373;

Widersten, 1976, p. 866)

This moderate size deepwater (5-400 m) circumarcticboreal anemone has a smooth column, about as tall as

wide or less so, with flaring base, sometimes with a

mound formed in the center of the base. It sometimes

occurs with Urtieina erassieornis, which it resembles, except for the broader base, flatter body, and totally

smooth column. It is able to detach easily and swims in

response to attack by predatory seastars.

Antholoba perdix (Verrill, 1882, p. 223, Urtieina) (see also

Verrill, 1883, p. 49; Verrill 1899b, p. 210 (Paraetis,

Archaetis); Widersten, 1976, p. 869)

This flattened deepwater (>100 m) anemone has several hundred small tentacles covering most of the oral

disc, increasing in size toward the mouth. This anemone

is able to capture large (30 em) swimming fish in aquaria.

The body is flattened vertically. The column is smooth

when alive, and is deeply wrinkled vertically and horizontally when preserved. The fosse is not distinct and

tentacles begin at the margin. This species is often

found on shells from Cape Cod south into the Caribbean sea.

Family Hormathiidae

Hormathia nodosa (Fabricius, 1780, p. 350, Aetinia) (see

also Carlgren, 1928, p. 294 (Hormathia); Carlgren, 1933,

p. 26; Carlgren, 1942, p. 46; = Aetinauge rugosa Verrill,

1922, p. 95; =A"borealis p. 98; = Chondraetinia tubereulosa,

p. 102; Widersten, 1976, p. 871).

A common aggregating deepwater (>60 m) anemone

found from the Canadian Arctic and Greenland to

Cape Cod, this species has a thick column with a wide

range of variation in texture, from a few longitudinal
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 53

rows of raised tubercles near the margin to rows of

large hooked tubercles (var. tuberculosa) on the entire

column. A periderm forms on the scapus whereas the

scapulus is smooth. The base is as wide or wider than the

column. This species may form hybrids with Harmathia

digitata where they overlap, according to Carlgren (1942).

This anemone sometimes attaches to shells, crab carapaces, gorgonians, and other hard objects.

Actinauge longicornis (Verrill, 1882, p. 222, Urticina) (see

also Verrill, 1882, p. 53 (Actinauge); Carlgren, 1942, p.

36; Riemann-Ziirneck; 1973).

This is a very common large deepwater (>200 m)

anemone found from Cape Cod to the Caribbean, taller

than wide, whose expanded oral disc is not bilobed.

Living specimens have a distinct dark-colored scapulus

(brown, purple) against a bright white column and

tentacles. The thick column wall has very small low tubercles, or the entire column is almost completely smooth

(some wrinkles when preserved) and lacks a periderm.

This species has fewer mesenteries than does A. venilli,

with 16 or more perfect and six sterile in the upper

column, 'lnd six pairs near the base. It is found south of

Cape Cod to the south Atlantic (Riemann-Zurneck, 1973).

Actinauge verrilli (McMurrich, 1893, p. 184) (see also

Verrill, 1922, p. 94; Carlgren, 1933, p. 28; == Urticina

nodosa Verrill, 1873a, p. 440; Widersten, 1976, p. 872;

Dunn, 1983, p. 48)

This large and common deepwater (>120 m) anemone

is found from the arctic south to Delaware. It is taller

than wide, and its expanded oral disc forms two large

lobes resembling the leaves of a Venus Dytrap plant.

The tentacles and oral disc retract fully on preservation. The column wall is thick and sculptured into large

square tubercles, larger in the upper half, whose form

and number vary widely. A periderm is usually evident

on the scapus. This species has more mesenteries than

does A. longicomis, with 40 perfect and one sterile in the

upper column, and six pairs near the base. This species

also has been found in the south Atlantic (McMurrich,

1893; Dunn, 1983).

Phelliactis americana (Widersten, 1976, p. 872)

Only a few specimens of this deepwater (to 1,000 m)

species have been collected, on gravel, south of Nova

Scotia and east of Cape Cod. The thick column has a

distinct ridged scapulus and tubercled scapus and is

asymmetrical, one side being somewhat larger than the

other. Tubercles are low hemispheres in up to 48 regular vertical rows, and there may be traces of periderm.

Acontia are abundant and the tentacles are short and

numerous (to 190). This species resembles Hormathia

nodosa but has more tentacles, a more prominent scapus, and tubercles in very regular vertical rows.

Paracalliactis involvens (McMurrich, 1893, p. 182,

Adamsia) (see also Carlgren, 1947, p. 15; == Urticina

consorsVerrill, 1882, p. 225)

A deepwater species (>200 m) found from Cape Cod

south to at least Virginia. This anemone forms a

carcinoeCium housing a hermit crab (Sympagurus pictus).

The column is smooth and there are numerous small

tentacles. The base secretes a cuticle and completely

replaces the original shell that it once covered.

Allantactis parasitica Danielssen, 1890, p. 20 (see also

Carlgren, 1933, p. 30; Carlgren, 1942, p. 34; == Calliactis

kroyeri Danielssen, 1890, p. 36)

This deepwater circumartic-boreal anemone (6-1,448

m) lives on shells (e.g. Sipho, Neptunea) inhabited by

hermit crabs (Pagurus bernhardus). The anemone does

not replace the shell by forming a cuticle. The column is

smooth and often extremely flattened vertically when preserved, although Carlgren (1942) illustrated one specimen, probably preserved, about three times as long as

wide. This species is more northerly in distribution than

Paracalliactis involvens, which is similar, and it possesses a

column that is not divided into scapus and scapulus.

Calliactis tricolor (Lesueur, 1817, p. 171, Actinia) (==Actinia

bicolor, Lesueur, 1817, p. 171; == Cereus sol Verrill, 1864-

1866, p. 24; == Adamsia egletes Duchassaing and Michelotti,

1866, p. 134) (see also McMurrich, 1898, p. 234

(Adamsia); Haddon, 1898, p. 457 (Calliactis); Duerden,

1902, p. 359; Field, 1949, p. 12; Carlgren and Hedgepeth,

1952, p. 160)

This shallow-water «30 m) anemone, about as tall as

wide when preserved, lives on gastropod shells (Polinices,

Fasciolaria, Rehderia, Busycon) alone or inhabited by hermit crabs (Pagurus policaris, P. impressus, Clibinarius

vittatus, Petrochirus bahamensis) or on crab carapaces

(Pericera cornuta, Hepatus epheliticus) (Cutress et al. (1970);

McLean and Mariscal [1973]; Brooks and Mariscal [1986]).

It is found in the Caribbean, the Gulf of Mexico, and

north along the east coast to North Carolina.

Adamsia sociabilis Verrill, 1882, p. 225 (see also Verrill,

1883, p. 45)

A small deepwater (>150 m) species found from New

Jersey north to at least Nova Scotia, this anemone forms

a conical carcinoecium covering only the abdomen of a

hermit crab (Catapagurus sparreri) , and is found on
54 NOAA Technical Report NMFS 141

mollusk shells (Cadulus, Dentalium, Cavolina) when juvenile. The anemone is carried on the crab with the

mouth and tentacles pointing downward, sweeping the

substrate. Tentacles form two distinct circles around

the protruding mouth.

Chondmphellia coronata (Verrill, 1883, p. 53, Actinauge

nodosa) (see also Carlgren, 1925, p. 6, Chondrophellia;

Carlgren, 1928, p. 215; Carlgren, 1942, p. 32, p. 77; =

Actinaugefastigiata, McMurrich, 1893, p. 187; = Hormathia

elongata Gravier (1918, p. 14).

This distinctive and rarely collected deepwater (>600

m) orth Atlantic anemone is found as far south as

Virginia. It is about two to three times as long as wide

when preserved and has a "crown" comprising two rows

of 12 tubercles each at the margin, completely hiding

the retracted scapulus and tentacles when contracted.

A dark brown periderm covers the lower half of the

column. The base can be broad, concave, and folded

around a mud bolus.

Amphianthus nitidus (Verrill, 1898b, p. 144, Raphactis)

(see also Carlgren, 1934, p. 10, Amphianthus; Widersten,

1976, p. 873)

This small deepwater (128-605 m) anemone has an

amplexicaulous base that surrounds gorgonian branches

(Paramuricea, Primnoa) or, less commonly, hydroid stems

or sponges. This species has been collected from Delaware north to at least Nova Scotia. Acontia are yellow

and numerous. Inner tentacles are longer than outer,

and the scapus is smooth or has up to eight low tubercles. The broad and distinctly ridged scapulus is

visible when the anemone is contracted. This species

closely resembles A. mirabilis but differs in size, column,

and scapulus texture, and in the presence of acontia.

Amphianthus mirabilis (Verrill, 1879, p. 74, Synanthus)

(see also Verrill, 1883, p. 48, Verrill. 1899b, p. 205;

Carlgren, 1942, p. 7)

This is a small deepwater anemone (375-620 m) collected from the fishing banks off Nova Scotia and south

to South Carolina. It has a smooth column and a broad

amplexicaulous base wrapped completely around gorgonian branches (e.g. Paragorgia arborea, Paramuricea

grandis) often with bases of several anemones fused.

Acontia and cinclides are absent, contrary to Carlgren's

(1949) diagnostic characters of the genus, and pedal

laceration probably occurs.

Stephanauge nexilis (Verrill, 1883, p. 55, Actinauge) (see

also Verrill, 1899b, p. 145 (Stephanauge), 1922, p. 99;

Carlgren, 1942, p. 37; Widersten, 1976, p. 875)

A circumarctic deepwater (150-547 m) species found

as far south as Cape Cod, this anemone's amplexicaulous

base grasps the denuded axis of a pennatulid (e.g.

Balticina). The upper scapus forms up to 26 flattened

square tubercles, and the scapulus is distinctly ridged.

Tentacles are numerous, to 200, but usually <l00; acontia

are few or possibly absent; there are few cinclides, and

those arc confined to the directive chambers. The base

is elongated along one axis, causing lateral flattening of

the scapus near the base. Some specimens may have

more than six pairs of perfect mesenteries. See S.

acanellae for comparison.

Stephanauge acanellae (Verrill, 1883, p. 46, Sagartia) (see

also Carlgren, 1942, p. 49, Stephanauge abyssicola

(Moseley); Verrill, 1898b, p. 145; Verrill, 1922, p. 101,

Raphactis; Carlgren, 1928, p. 300)

This is a small deepwater (>487-2328 m) anemone with

an amplexicaulous base grasping branches of the gorgonians Acanella aTbuscula or Mopsea sp. It is a

circumarctic species, found south to New Jersey and

possibly to the Antilles (Verrill, 1883). The column is

smooth and laterally flattened especially near the base,

sometimes with one or more rings of low tubercles.

Acontia are absent. It resembles S. nexilis but differs in

column texture, color, depth, and substrate type. Morphological similarity and a nearly undistinguishable

cnidom suggests that S. nexilis could be a subspecies or

a sister species of S. acanellae.

Stephanauge spongicola (Verrill, 1883, p. 47, Sagania) (see

also McMurrich, 1898, p. 238; Widersten, 1976, p. 875)

This is another small deepwater (144-578 m) species

found from Nova Scotia south to N. Carolina. It has a

flat or amplexicaulous base, a smooth column sometimes with thin periderm, and 70-80 tentacles increasing in size from margin to inner oral disc. It occurs on a

variety of substrata including hydroid stems, sponges,

rocks, and worm tubes. Acontia are numerous, and

cinclides are rare. The irregular number of siphonoglyphs (1-4) suggests asexual reproduction by longitudinal fission. It differs from S. acanellae and S. nexilis by

having fewer tentacles and mesenteries and abundant

acontia, and by having atrichous nematocysts in the

tentacles and column.

Family Sagartiidae

Sagartiogeton verrilli Carlgren, 1942, p. 18 (=Sagania

abyssicola Verrill, 1882, p. 314; = Sagartiogeton abyssicola

Carlgren, 1928, p. 260)
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 55

Sagartiogeton verrilli is a small deepwater (139-1,416 m)

anemone found on worm tubes (Hyalinoecia artifex) or,

more rarely, on rock. The base is Oat or amplexicoulous,

acontia are numerous, tentacles are red to maroon

when alive; the actinopharynx is deep maroon when

preserved. The column has a periderm that peels off in

patches. The 96 tentacles increase in length from the

margin to the inner oral disc. This species resembles

Stephanauge acanellae and S. nexilis but has fewer tentacles, a larger number of perfect mesenteries, numerous acontia, and tenaculi on the scapus. It also resembles S. spongicola but has more mesenteries, more

perfect mesenteries, tenaculi, and lacks atrichous nematocysts on the tentacles and column ectoderm. It differs

from all three similar species by having microbasic

amastigophore nematocysts in the tentacles, column,

and acontia.

Actinothoe modesta (Verrill, 1866, p. 337, Sagartia) (see

also Hargitt, 1914, p. 1240)

This elongate low intertidal to shallow subtidal burrowing anemone has an adherent base attached to cobbles,

shell, or rock below sand. Two distinctive brown stripes

occur on each tentacle when alive. It has been found

only in Long Island Sound and southern Cape Cod to

date. Acontia are extruded through the column and

there is a mucous sheath on the column containing

adherent sand grains when the anemone is first collected. This is the only burrowing acontiate anemone

along the east coast of North America. The following

three species are incompletely described and may be

synonymous with A. modesta or with other North Atlantic Actinothoe species: Actinothoe gracillima (McMurrich,

1887, p. 61, Sagartia); Actinothoe pustulata (McMurrich,

1887, p. 60, Sagartia); Actinothoe bradley (Verrill, 1869, p.

484, Sagartia).

Family Metridiidae

Metridium senile (Linnaeus, 1767, p. 1088, Actinia) (see

also Stephenson, 1935, p. 214; Carlgren, 1942, p. 59;

Hand, 1956, p. 192; = Metridium dianthus Carlgren 1933,

p. 22; = Sargartia dianthus Gosse, 1855, p. 274; = Actinia

rnarginata Lesueur, 1817, p. 172; = M. fimbriatum Verrill,

1865, p. 150; Widersten, 1976, p. 850)

This very common and large shallow-water (mid-intertidal to 166 m) circumarctic-boreal anemone occurs

south to New Jersey and is distinguished by a lobed

crown of up to thousands of small tentacles in large

specimens. The column is smooth and has a distinct

collar, not present in small individuals. Asexual reproduction by pedal laceration produces clones of anemones all with the same color pattern (Parker, 1897;

Hoffmann, 1976; Hoffman, 1986; Shick and Hoffmann,

1980); clones interact agonistically by inflating elongate catch-tentacles with specialized nematocysts

(Purcell, 1977; Sebens, 1985). Acon tia are numerous

and are easily discharged through the column cinclidae.

This anemone preys on small to relatively large zooplankton (Sebens and Koehl, 1984). Color is extremely

variable. Common substrates include rock, shell, and

wood.

The reproductive cycle has been described for the

west coast population whose individuals have some mature gametes throughout the year (Bucklin, 1982). J uveniles and small intertidal adults lack a lobed oral disc

and have elongate tentacles of roughly equal size

(Rawlinson, 1934; Carlgren, 1942). Although large

adults are easily identified, small individuals could be

confused with Diadumene leucolena, the only other epifaunal acontiate anemone with a partially overlapping

range. Diadumene has prominent cinclides in vertical

rows, and many differences in the cnidom (Table 3,

also Hand, 1956; Westfall, 1965). Bucklin (1985) identified two ecophenotypes, M. senile dianthus and M. senile

pallidum (Stephenson, 1935) from British shores (by

electrophoresis). Genetic distance between British and

North American east coast Metridium indicates a subspecies- to species-level of differentiation. See recent

discussion in Fautin et al. (1989).

Family Aiptasiidae

Aiptasia pallida (Verrill, 1864-1866, p. 26, Dysactis) (see

also Verrill, 1864-1866, p. 322, Paranthea; McMurrich,

1887, p. 59, Aiptasia; Andres, 1883, p. 391; = Diadumene

leucolena in Field, 1949, p. 13, according to Hand, 1956,

p. 229; den Hartog, 1985, p. 175)

A small mid-intertidal to shallow subtidal Caribbean

anemone found as far north as North Carolina. The

inner tentacles are long, slender, and of roughly equal

length; those of the outer cycle are only about one

fourth the length of the innermost ones. The most

distinctive feature of this species is the two (rarely one

or three) prominent horizontal rows of cinclides, without raised edges when preserved, girdling the

midcolumn; the column is smooth otherwise. This

feature distinguishes it from the co-occurring A.

eruptaurantia.

This species can be distinguished from its Caribbean

congener, Aiptasia tagetes, by the lack of distinct rows of

cinclides in the latter. Both species harbor zooxanthellae (Clark andJensen, 1982) giving the otherwise clear,

white, or white speckled anemone a brown color. Common substrata include rock, algae, sponges, and man-
56 NOAA Technical Report NMFS 141

grove roots. Pedal laceration is common, resulting in

clones of a single sex. Adults produce large oocytes

(250 ~m) in spring and summer and spawn by October,

although some populations spawn in April as well (in

Florida,Jennison, ]983).

Aiptasia eruptaurantia (Field, ] 949, p. 16, Artinothoe)

(see also Carlgren, 195~, p. 376)

This moderate-size intertidal anemone is known only

from North and South Carolina. It has distinctive red

spots (surrounding cinclides) on the column and lines

on the oral disc. Cinclides occur in one or more, coomonly several, horizontal bands on the upper column

and have distinct raised rims. Tentacles decrease in

length away from the mouth; the outer ones are about

two thirds as long as the innermost. This species can be

distinguished from A. pallida by the arrangement of

tentacles and cinclides, by the presence of microhasic

amastigophore nematocysts in the tentacles and column of A. eruptaurantia, and by several other differences in the cnidom (Table 3).

Family Bathyphelliidae

lJaontesia praelonga (Carlgren, 1928, p. 259, 302,

Sagartiogl'lonpraelongus) (see also Carlgren, 194~, p. 5)

A rarely encountered deepwater (>1,000 m) anemone

known previously from Scandinavia and south of Iceland, this species was collected recen tly in 1,] 62 m

depth off Delaware (by B. Hecker, K. Sebens ident.).

The scapus has tightly packed elongate papillae (tenaculi)

and is smooth to longitudinally ridged. This species

differs from all others in this region by the presence of

these numerous densely packed tenaculi on the scapus.

Family Aiptasiomorphidae

Haliplanella luciae (Verrill, ] 89Ha, p. 493, Sagartia) (sec

also Stephenson, 1935, p. 197; Hargitt, 1914, p. 241;

Hand, 1956, p. 210 (Haliplanplla); = Sagartia lineatn

Verrill, ]869, p. 57; = HalijJlanella lineata (Verrill) in

Manuel, 1981, p. 134)

This small intertidal anemone has spread, possibly from

Asia (Uchida, ] 932; Williams, 1954, Williams, 1973)

throughout the world. The smooth column is green to

brown, frequently with vertical orange, yellow or white

stripes. This anemone reproduces asexually by longitudinal fission (Davenport, 1900; Parker, 1902; Davis,

] 909; Davis, 1919; Minasian, 1976; Shick and Lamb,

1977; Minasian and Mariscal, 1979) and occasionally

forms elongate catch-tentacles, probably used in agonistic encounters (Williams, 1975a; Watson and Mariscal,

1!J83) resulting in spacing between individuals.

Family Diadumenidae

lJiadumene leucolena (Verrill, 1864-1866, p. 366, Sagartia)

(see also Carlgren, 1949, p. 109, Diadumene, Carlgren,

1952, p. 25; Hand, 1956, p. 223; Vassallo, 1969, p. 121;

not lJ. leucolenaField, 1949, p. 13)

A small low intertidal to shallow subtidal anemone oftcn with symbiotic algae (zooxanthellae), especially in

the actinopharynx and oral disc. This species is found

from South Carolina to northern Massachusetts (Salem

Harhor; K. Sebens, unpub!.). Acontia are numerous,

and the column usually has vertical rows of low vesicles

along the top half. This anemone resembles small

l'vIetrirlium senile but can be distinguished by column

structures and by several differences in the cnidom

(Table 3).

Order ZOANTHARIA

Family Epizoanthidae

Epizoanthus inCTustatus (Duben and Koren, 1847, p. 268,

iHamillifera inrrustata); (= loanthus couchii Gosse, 1860,

p. 267; = 1. rubricornis Haddon and Shackleton, 1891, p.

652; = Epizoanthus amencanus Verrill, 1864-1866, p. 34;

see also Haddon and Shackleton, 1891, p. 636; Manuel,

1981, p. 73)

This moderately deepwater (>60 m) zoanthid (large

colony with 15-20 polyps) forms a carcinoecium housing a hermit crab (e.g. Eupagurus pubescens). Small colonies (2-5 polyps) are found loose on worm tubes

(Hyalinoecia artifex) or on small debris. Polyps are up to

1.5 em tall by 0.6 em wide preserved, with sand-encrusted columns and upper colony surfaces. Polyps cover

only the upper (dorsal) colony surface.

1:.,pizoanthus paguriphilus Verrill, 1882, p. ] 37, 316 (see

also Haddon and Shackleton, 1891, p. 64]; =E. abyssorum,

Manuel, ]981, p. 77; USNM specimens ident. by

K. Sebens)

This deepwater (>500 m) zoanthid (colony <] 2 polyps)

forms a carcinoecium housing a hermit crab (e.g.

Pampagurus pilosimanus). Polyps are up to 2 em tall by

1.2 em wide preserved, with sand-encrusted columns

and upper colony surfaces. There is a large polyp-free

area on the central, upper (dorsal) surface. This and

the larger polyp size distinguish it from E. incrustatus.
OrderCE~T~

Family Cerianthidae

Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 57

tacles. This species has been photographed In Baltimore Canyon by B. Hecker.

Cerianthus borealis Verrill, 1873b, p. 5 (see also Verrill,

1883, p. 534; Kingsley, 1904, p. 345; Shepard et a!.,

1986; Table 3; = cerianthid B, Carlgren, 1927, p. 444)

This large shallow to deep subtidal (0-500 m) cerianthid

has 150-200 solid colored or distinctly banded marginal tentacles of unequal length and multiplanar arrangement. In large deepwater specimens, the tube

protrudes 5-15 em above the substrate but may be at or

near the sediment surface in very shallow water. This

species is found from Delaware north to the arctic.

Spirocysts of the marginal tentacles are up to 56 )..tm in

length, distinguishing this species from Ceriantheopsis

americanus. Kingsley (1904) presented figures of the

larva of this species (termed Arachnactis larva).

Ceriantheopsis americanus (Agassiz, 1859, p. 24, Cerianthus)

(see also Verrill, 1864-1866, p. 32; McMurrich 1890, p.

131-150, Ceriantheopsis; Carlgren, 1912, p. 41; Field,

1949,p.19)

A large low intertidal to shallow subtidal (0-70 m)

species found south of Cape Cod to the Caribbean Sea,

this anemone has up to 125 marginal unbanded tentacles of approximately equal length held in a

multiplanar array. The tube opening does not protrude

much above the substrate, if at all. Spirocysts of the

marginal tentacles reach only 27 )..tm in length, half the

size of those in Cerianthis borealis. Mariscal et al. (1977)

provided information on cnidae and tube construction.

Cerianthid I (undescribed species, see Table 3 in

Shepard et a!., 1986; = cerianthid A)

This is a small very deepwater (1,600-1,900 m)

cerianthid found in dense aggregations. Tentacles are

held close to the sediment surface and the tube does

not protrude above the substrate. Preserved specimens

are <6 em long. Marginal tentacles are uniformly dark

red to black or purple in living specimens, numbering

about 30. One specimen was collected in Baltimore

Canyon (1,806 m) by B. Hecker. Shepard et a!. (1986)

reported the known range from Oceanographer to

Veatch Canyons along the continental slope.

Cerianthid II (undescribed species)

This large very deepwater (1,500-3,000 m) species has

marginal tentacles of equal length and multiplanar arrangement, dark, purple to almost black and unbanded.

The tube extends only a few em above the sediment.

Photographs indicate there are about 60 marginal tenCerianthid III (undescribed species, see Table 3 in

Shepard et aI., 1986; = cerianthid C)

This deepwater species (200-400 m) is found in canyons on the continental slope (Lydonia and Oceanographer to Block Canyons). The tube protrudes up to 20

em above the sediment surface. Marginal tentacles are

white with purple marks in living specimens and are of

unequal length, held in a uniplanar parabolic arrangement. No specimens have been collected, but it has

been photographed in situ by Shepard et al. (1986).

Photographs indicate there are approximately 80-100

marginal tentacles in large specimens.

Cerianthid IV (undescribed species, see Table 3 III

Shepard et aI., 1986; = cerianthid D)

A deepwater species known only from photographs

taken in Hudson Canyon (200-300 m). The greenishyellow marginal tentacles are held flush to the sediment surface, are of unequal length, and are arranged

in a multiplanar array. No specimens have been collected, and its identification, even as a cerianthid, is

thus only tentative.

Order CORALLIMORP~

Family Corallimorphidae

Corynactis delawarei Widersten, 1976, p. 860

This small, deepwater (>200 m) corallimorpharian has

been collected south of Cape Cod. Polyps are solitary or

connected in rows ofseveral individuals formed by longitudinal fission, with smooth unornamented columns. Tentacles bear acrospheres on their tips, and inner tentacles

are longer than outer ones. Column and base are bright

red to reddish brown or white in life. Acrospheres are

distinct and white. Only one indistinct siphonoglyph is

present. Found on rock, onuphid polychaete tubes, and

on shell fragments (specimens in USNM).

Glossary _

Definitions (modified and expanded from Carlgren,

1949; Manuel, 1981; for a more complete set of terms

specific to cerian thids, see Arai, 1965).

Aboral: Part of polyp or structure remote from mouth,

toward base.

Acontioid: Acontium-like organ ofsome Ceriantharia.
58 NOAA Technical Report NMFS 141

Acontium: Thread-like nematocyst-bearing organs attached to mesenteries, usually below mesenteric filament, in certain Actiniaria.

Acrorhagus (pI. acrorhagi): Verruca-like organ, with

holotrichous nematocysts, in the parapet or fosse

region of some anemones (Actiniidae), used for intra- and interspecific aggression. Also termed "marginal spherule."

Acrosphere: Spherical tentacle tip, clearly demarcated

from stalk, often bearing dense nematocyst batteries.

Actinopharynx: Tubular "throat" of an anthozoan polyp,

usually compressed laterally.

Actinostola rule: Principle that mesentery pairs of 3rd

and later (youngest) cycles have one member of pair

much smaller.

Basal disc: Flattened base of many solitary anthozoans

(=pedal disc).

Budding: Asexual reproduction manifested by growth

of new polyps from body wall, or growth of tissue

between polyps (in colonial Anthozoa).

Capitulum: Delicate uppermost region of the column

directly adjacent to tentacle bases in some Actiniaria.

Carcinoecium: Structure containing a hermit crab, formed

by some colonies of Zoanthidea (also formed by the

actinarians Adamsia, Stylobates, and Paracalliartis).

Catch-tentacle: Modified inner tentacle with numerous

holotrichous nematocysts (in some acontiate

Actiniaria); used in in tra- and in terspecific aggression (=fighting tentacle, sweeper tentacle).

Chamber: Area of coelenteron enclosed between adjacent mesenteries (includes endocoels, exocoels).

Ciliated tract: Lateral strand of trilobed mesen teric

filament in most Actiniaria and Zoantharia.

Cinclis (pI. cinclides): Perforation or organized weak

spot in column wall ofsome Actiniaria.

Circumscript muscle: Well-defined endodermal muscle,

usually pronounced, borne on a single mesogleal

lamella above the level of the body wall or mesentery.

Cnida (pI. cnidae): (nematocyst, spirocyst, ptychocyst).

Microscopic capsule containing a long explosively

evertible tube, sticky or penetrative, used for prey

capture, defense, adhesion, burrow lining construction (cerian thids); characteristic of all Cnidaria.

Cnidoglandular tract: Central or only strand of the

mesentric filament.

Cnidom: Complement of cnida categories in an anthozoan.

Cnidorhagus: Globular projection from a mesentery,

with numerous nematocysts (Ceriantharia).

Coelenteron: Body cavity of anthozoans (=gastric cavity).

Collar: Strongly marked circular fold of body wall just

below margin (=parapet).

Colony: Attached polyps produced by asexual budding

from a common sexually produced ancestral polyp,

retaining permanent organic connection between

its constituent individuals.

Column: Cylindrical body of an anthozoan polyp (often

refers only to external surface of body wall).

Complete mesentery: Equivalen t to "perfect mesentery";

see definition.

Conchula: Raised ear- or shell-shaped extension of lip

around siphonoglyph opening in Peachia.

Contractile: Capable of being shortened by muscular

action (not equivalent to retractile).

Couple: Two equivalent mesenteries on opposite sides

of the body across the directive axis.

Cuticle: Any hard outer covering of column or pedal

disc in Actiniaria or Zoantharia, usually chitinous or

sand-encrusted.

Cycle: Set of mesentery pairs of similar size and development time (in relation to those of other cycles)

and more or Jess equally spaced in the radial plan.

Also, a ring of tentacles of equal size arising from

chambers between mesenteries of the equivalent

cycle.

Decamerous: Arranged in multiples of ten, applying to

number of tentacles or pairs of mesenteries in a

cycle.

Diffuse muscle: A muscle composed of a number of

separate mesogleallamellae.

Directive axis: Axis of bilateral symmetry in an anthozoan pOlyp, passing through the siphonoglyph (s)

and between pairs of directive mesenteries (long

axis of mouth) in cross section.

Directive mesentery: One of a pair straddling the directive axis and connected to a siphonoglyph when

present. In Hexacorallia, retractor muscles are on

the exocoelic side, as opposed to all other mesentery

pairs.

Distal: Part or region away from the base (=oral).

Dorsal: Side of body containing the siphonoglyph in

cerian th ids.

Ectoderm: Outer cell layer of the body wall (Cnidaria).

Ectomesogleal muscles: Muscles embedded primarily

in the mesoglea on ectodermal side.

Endocoel: Space between a pair of mesenteries.

Endoderm: Inner cell layer of the body wall (Cnidaria).

Covers both sides of mesenteries (=gastrodermis).

Endomesogleal muscles: Muscles embedded primarily

in the mesoglea, on endodermal side.

Exocoel: Space between two adjacent pairs of mesenteries.

Fertile mesentery: Mesentery bearing gonads somewhere

along its length.

Filament: = Mesenteric filament.

Fosse: Groove around the top of the column, enclosed

by the parapet or collar in some Actiniaria.

Gastric cavity: Alternate term for Coelenteron.

Hexamerous: Arranged in multiples of six (pairs of mesenteries or tentacles), the fundamental arrangement

in Actiniaria, Corallimorpharia, and Scleractinia.
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 59

Imperfect mesentery: One that does not attach to the

actinopharnyx.

Inner cycle: Cycle oftentacles nearest mouth (primary).

Labial tentacles: Short tentacles arising from oral disc

near mouth (Ceriantharia).

Lacuna: Cavity in mesoglea of zoanthids.

Lamella: Planar protrusion of mesoglea around which

portions of muscles are organized.

Larviparity: Brooding of larvae in the paren tal coelenteron.

Limbus:Junction of column and basal disc (Thenarian

Actiniaria) .

Longitudinal fission: Asexual reproduction in

Corallimorpharia and Actiniaria, by vertical (longitudinal) division of the polyp into two parts.

Macrocoele: Space between two large macrocnemesin which

other smaller macrocnemes and microcnemes occur.

Macrocneme: Perfect mesenteries with well-developed

musculature, filament, and gonads.

Margin: Junction of column wall and oral disc, sometimes applied to the parapet.

Marginal pseudospherules: Vesicles situated at the margin, resembling acrorhagi, often with an aperture, containing same cnidae as column (=pseudoacrorhagi).

Marginal spherules: Alternate term for acrorhagi.

Marginal tentacles: Tentacles of outer or marginal series in cerianthids. Tentacles arising from the margin in Corallimorpharians.

Marginal tooth: Small protuberance on the parapet in

zoanthids.

Mesenteric arrangement: Mesenteries are arranged in

pairs. Directives are situated in the directive axis and

have their longitudinal muscles (retractors) turned

towards the exocoel. Ordinary pairs have the retractor muscles turned towards the endocoel (facing

each other). The directives are always perfect, other

pairs may be perfect or imperfect. Usually the partners of the ordinary pairs are equally developed

(equal), but sometimes they may differ in size (unequal). Usually the arrangement of the pairs is by

multiples of six, hexamerous, sometimes of eight,

octamerous, or of ten, decamerous. Irregularities in

the arrangement are fairly common in connection

with displacement or absence of the directive or

dislocation of tentacles or after asexual reproduction. In elongate mud- or sand-dwelling forms, the

perfect mesenteries are often strongly differentiated

from the imperfect ones. Perfect mesenteries, or

macrocnemes, have very strong retractors, gonads,

and filaments. Imperfect mesenteries, or microcnemes, lack these organs. Intermediate mesenteries may occur. The arrangement of mesenteries in

some families may be different from normal arrangement (e.g. Minyadidae, Endocoelantheae, Exocoelactiidae, and some Actinostolidae).

Mesenteric filament: Thickened structure attached

along inner free edge of most mesenteries.

Mesenteric muscles: One side of each mesentery is

occupied by a longitudinal muscle, the other by a

transverse and a parietobasilar muscle; the latter

usually run obliquely from the column to the pedal

disc. In most Actiniaria, basilar muscles also run

along both sides of the base of the mesentery, close

to the pedal disc. The longitudinal muscles form

more or less strong retractors. When the muscles are

very strongly concentrated and only one mesogleal

lamella (or a few main lamellae close to each other)

issues from the main lamella of the mesentery, the

muscle is termed circumscript. Restricted retractors

show about the same degree of concentration, but a

number ofmore scattered muscle lamellae arise from

the mesentery. If the retractor is not strongly concentrated, it is termed diffuse. Sometimes, especially

in elongate Athenaria, the longitudinal muscles are

very weak except where they form the retractor. Close

to the body wall, the longitudinal muscles increase

in size and, together with the parietobasilar muscles,

form a parietal muscle.

Mesenteric insertion: Attachment of mesentery to column wall or disc, usually externally visible as a line.

Mesenteric stoma (pI. stomata): Aperture in a mesentery connecting adjacent chambers of coelenteron

(distal or oral end). Marginal stomata are just below

margin and oral stomata occur near mouth.

Mesenteries: Internal walls, arranged radially and arising from the column wall and oral disc or base.

Mesoectodermal muscles: Muscles that are primarily ectodermal, with a small part embedded in the mesoglea.

Mesoendodermal muscles: Muscles that are primarily endodermal, with a small part embedded in mesoglea.

Mesoglea: Middle supporting layer ofthe body wall and

other structures in Cnidaria, consisting of collagenous elastic connective tissue (alt. mesogloea).

Mesogleal muscles: Muscles wholly embedded in the

mesoglea.

Metacnemes: Mesentery pairs arising after the first 12

mesenteries.

Microcneme: Small imperfect mesentery, lacking welldeveloped musculature, gonads, and filament; used

only in opposition to macrocneme.

Multiplication chamber: Ventral chamber of cerianthids

in which new mesentery couples arise.

Muscle process: Raised ridge of mesoglea bearing a

layer of muscle fibers.

Nemathybome: Externally opening pocket in the mesoglea, containing a nematocyst battery, in certain

Edwardsiidae. Often arranged in eight rows.

Nematocyst: Major category of cnidae usually possessing

an armament ofspines on the tube, arranged in three

right-hand spirals. Types are given in introduction.
60 NOAA Technical Report NMFS 141

Nematosome: Ball of flagellated cells and nematocysts

occurring free within the coelenteron of Nematostella.

Oral disc: Transverse plate of tissue beginning at distal

end of column with mouth at its center and tentacles

arising from its periphery or from the entire disc.

Papillae: Very small, nonadhesive protrusions of column, usually solid. Sometimes used to refer to very

small vesicles.

Parapet: Infolded or upright distal rim of scapus, enclosing a circumferential groove (fosse) in Zoantharia

and Actiniaria (= collar).

Parietal muscle: See mesenteric muscles.

Parietobasilar muscle: See mesenteric muscles.

Pedal disc: Basal disc.

Pedal laceration: Asexual reproduction in Actiniaria by

fragmen tation from the edge of the pedal disc.

Perfect mesentery: A mesentery reaching from the column wall and attaching to the actinopharynx.

Periderm: An external layer of secreted material of

variable texture and appearance often incorporating sand, shell, and detritus.

Physa: Rounded aboral end of many Athenaria

(Actiniaria) .

Planula: Ciliated, sexually (sometimes asexually) produced larva of Anthozoans.

Polyp: Single anthozoan individual.

Primary mesentery or tentacle: One belonging to the

first cycle to arise during development: in mesenteries, usually the most fully developed, in tentacles,

usually the inner cycle.

Protocnemes: The first 12 mesenteries that arise as

couples in Actiniaria.

Proximal: Pertaining to the attached or basal end, :)f

the end of a polyp nearest the colony mass (=aboral).

Pseudocycle: One of several concentric rings of marginal tentacles in the Ceriantharia (not a true cycle

because mesenteries are unpaired).

Ptychocyst: Category of cnidae produced by ccrianthids,

used in tube construction and distinct from spirocysts

or nematocysts.

Radius: Area on the oral disc between adjacent mesenteric insertions.

Retractile: Capable of withdrawing oral disc, tentacles,

and often distal part of column into the remainder

of the column or into burrow.

Retractor muscle: See mesenteric muscles.

Ruga (pI. rugae): A tiny, usually microscopic, adhesive

spot or papilla on the column (Actiniaria).

Scapular ridge: Longitudinal ridge on the distal part of

the scapus, terminating in a marginal tooth on the

parapet (Zoantharia).

Scapulus: Naked region of the column above the scapus, typically (but not only) present when the latter

bears periderm (Actiniaria).

Scapus: Major and most extensive region of column;

immediately above basal disc or physa. May bear

periderm, verrucae, vesicles, tubercles (Actiniaria).

Septum (pI. septa): Vertical, radially oriented plate attached to the theca in the scleractinian coral skeleton. Also a widely used synonym for mesentery in

Anthozoa, but should be used only for skeletal septa

of corals.

Shaft: Proximal division of tube in heteronemic

nematocyst.

Siphonoglyph: Ciliated groove at one or both corners

of actinopharynx, often with thickened wall. Pumps

water into the coelenteron from the exterior.

Sphincter muscle: Circular muscles situated near distal

end of the column in many Actiniaria and

Zoanthidea. The endodermal circular muscles of

the column are often accumulated at or near the

margin, froming a sphincter, is either endodermal

or embedded in mesoglea (mesogleal sphincter). A

third form, an endomesodermal or mesoendodermal

sphincter is transitional. The endodermal sphincter

is either elongate and broadly attached to the column (diffuse sphincter), or more concen trated (restricted). The most concentrated endodermal sphincter is termed "circumscribed" of which there are two

kinds: "pinnate" with only one main lamella and

"palmate" with few main lamellae. In the genus

B%eera and some allied genera, the base of each

tentacle is provided with an endodermal sphincter. A

weak endodermal sphincter may also be present to

close the cinclides. An oral sphincter closes the mouth.

Spirocyst: Major category of cnidae possessing a tube

devoid of spines (except in some large forms) that

bears, when discharged, a left-hand spiral of glutinous substance, which is not penetrative. Spirocysts

are unique to the Hexacorallia.

Stoma (pI. stomata): See mesenteric stoma.

Stomodaeum: = Mouth and/or actinopharynx, obsolete term.

Sucker: Adhesive spot on the column of anemones,

histologically similar to a verruca but not protuberant. Often used for verruca.

Sulcus: = Siphonglyph, obsolete term.

Tentacle: Hollow appendage arising from oral disc,

usually slender and more or less prehensile, used for

prey capture.

Tenaculi: More or less solid papillae situated on column, tall and sometimes lobed; ectoderm is partly

chitinized and with an usually strong, sometimes

stratified cuticle, to which grains of sand or detritus

may adhere.

Thread: Distal division ofthe tube in heteronemic nematocysts.

Transverse fission: Asexual reproduction by division of

a polyp horizontally, across the column (Fagesia,

Gonaetinia, some cerianthids).
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 61

Tubercle: Solid mesogleal excrescence on the column,

covered by ectoderm but nonampullaceous (Actiniaria).

Ventral: Arbitrary orientation ofsingle siphonoglyph in

Actiniaria.

Verruca (pI. verrucae): Hollow, wartlike, adhesive excrescence of the column in certain Actiniidae.

Vesicle: Ampullaceous, nonadhesive evagination of the

column, simple or compound; with few or numerous

nematocysts of various categories.

Viviparity: Production of fully formed juveniles after a

period of brooding within coelenteron.

V-notch: A constant feature atjunction ofshaft and thread

in undischarged p-mastigophore nematocysts, caused

by final turn ofspines on shaft. Often funnel-shaped.

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Systematic Index

(not including obsolete synonyms) _

Acthelmis intestinalis . . . . . . . . . . . . . . . . . . . 48

Actinauge . . . . . . . . . . . . . . . . . . . 53

Actinauge alryssorum . . . 48

Actinauge longicornis . 12,15,31,53

Actinauge verrilli. . . . . . . . . . . . . . . . . . . . . . .. 53

AcGnernidae 49

Actinernus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 49

Actinernus nobilis 10, 14, 38, 49

Actinia 48,51,52,53

Actinia bermudensis 48

Actiniaria 1,2,3,5,17,22,49

Actiniidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 51
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 65

Actinoscyphia .... : . . . . . . . . . . . . . . . . . . . . . . . . .. 52

Actinoscyphia saginata 11, 15,41,52

Actinostella flosculifera . . . . . . . . . . . . . . . . .. 48

Actinostola . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4, 39, 52

Actinostola callosa 11,15,40,52

Actinostola groenlandica . . . . . . . . . . . . . . . . . . . . . . . 48

Actinostola spetsbergensis . . . . . . . . . . . . . . . . . . . . . .. 48

Actinostolidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 52

Actinothoe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 55

Actinothoe bradley . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

Actinothoe gracillima . . . . . . . . . . . . . . . . . . . . . . . . .. 55

Actinothoe modesta 12,32,55

Actinothoe pustulata . . . . . . . . . . . . . . . . . . . . . . . . . . 55

Adamsia 53

Adamsia sociabilis 12, 15,28,53

Aiptasia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

Aiptasia eruptaurantia . . . . . . . . . . . . . .. 12, 16, 32, 56

Aiptasia pallida 13, 16, 34, 56

Aiptasia tagetes . . . . . . . . . . . . . . . . . . . . . . . . . . . 48, 55

Aiptasiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 55

Aiptasiomorphidae . . . . . . . . . . . . . . . . . . . . . . . . .. 56

Aiptasiogeton 56

Allantactis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 53

Allantactis parasitica. . . . . . . . . . . . . . . .. ] 2, ] 5, 29, 53

Amphianthus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

Amphianthus margaritaceus 48

Amphianthus michaelsarsi . . . . . . . . . . . . . . . . . . . . . . 48

Amphianthus mirabilis ]2, ]5,42,54

Amphianthus nitidus ]2, ]5,42,54

Amphianthus vermiculatus 48

Anemonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5]

Anemonia sargassensis. . . . . . . . . . . . . . . . . . . . .. 43, 5]

Antholoba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 52

Antholoba perdix ] 1, 15,47,52

Anthopleura carneola. . . . . . . . . . . . . . . . . . . . . . . . . . 48

Anthopleura catenulata 48

Anthopleura vario-armata 48

Anthosactisjan-mayeni . . . . . . . . . . . . . . . . . . . . . . .. 48

Antipatharia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5

Aracnanthidae 5

Athenaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 22, 49

Aulactinia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Aulactinia capitata . . . . . . . . . . . . . . . . .. 11, 14, 45, 51

Aulactinia incubans . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Aulactinia stella 11, ]4,45,51

Bartholomea annulata. . . . . . . . . . . . . . . . . . . . . . . .. 48

Bathyphelliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56

Bathyphellia margaritacea . . . . . . . . . . . . . . . . . . . . .. 48

Bolocera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5], 52

Bolocera maxima. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Bolocera tuediae . . . . . . . . . . . . . . . . . . .. 11, 14, 43, 51

Bunodactis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

Bunodactis spetsbergensis . . . . . . . . . . . . . . . . . . . . . . . 48

Bunodeopsis antilliensis 48

Bunodeopsis globulifera 48

Bunodosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

Bunodosoma cavernata 10, ]4,46,49,51

Bunodosoma granuliferum. . . . . . . . . . . . . . . . . .. 48, 51

Cactosoma abyssorum 48

Calliactis 53

Calliactis tricolor 12,15,29,53

Ceriantharia 1,5,17,18

Ceriantheopsis 57

Ceriantheopsis americanus 13, ]6,21,57

Cerianthid I . . . . . . . . . . . . . . . . . . . . . . . . .. 13,19,57

CerianthidIl ]3,19,57

Cerianthid Ill 13,20,57

Cerianthid IV. . . . . . . . . . . . . . . . . . . . . . . .. 13,20,57

Cerianthidae 5

Cerianthus borealis 13, 16, 21, 57

Cerianthus lloydi . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Chondrophellia . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 54

Chondrophellia coronata 12,15,35,54

Condylactis gigantea . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Corallimorpharia 1,5,17,18,57

Corallimorphidae . . . . . . . . . . . . . . . . . . . . . . . . . .. 57

Corallimorphus ingens . . . . . . . . . . . . . . . . . . . . . . . .. 48

Corallimorphus rigidus .. . . . . . . . . . . . . . . . . . . . . .. 48

Corynactis 57

Corynactis delawarei 13, ]6, 18,57

Corynactis parvula . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Cribrinopsis similis . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Daontesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Daontesia praelonga . . . .. ] 3, 16, 44, 56

Diadumene . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

Diadumene leucolena. . . . . . . . . . . . .. 13, 16,35,55,56

Diadumenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56

Drillactis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 50

Drillactis pallida . . . . . . . . . . . . . . . . . . .. 10, 24, 50, 56

Edwardsia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 49

Edwardsia andresi. . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Edwardsia elegans . . . . . . . . . . . . . . . . . .. ] 0, ] 4, 26, 49

Edwardsia sipunculoides . . . . . . . . . .. 10, 14,24,26,49

Edwardsia sulcata 10,27,49

Edwardsiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 49

Endocoelantheae 4, 37

Endomyaria 5

Epiactis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Epiactisfecunda . . . . . . . . . . . . . . . . . . .. 11, 14, 46, 51

Epiactis marsupialis . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Epiactis nordmanni . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Epizoanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 56

Epizoanthus 5, 17,56

Epizoanthus danielseni . . . . . . . . . . . . . . . . . . . . . . .. 48

Epizoanthus erdmanni. . . . . . . . . . . . . . . . . . . . . . . . . 48

~pizoanthusglacialis. . . . . . . . . . . . . . . . . . . . . . . . .. 48
66 NOAA Technical Report NMFS 141 _

Epizoanthus incrustatus. . . . . . . . . . . . . .. 13, 16, 17,56

.cpizoanthus lindahli . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Epizoanthus paf5Uriphilus 13, 16, 18,56

Fagesia 10.50

Fagesia lineata . . . . . . . . . . . . . . . . . . . . . . . . . .. 23, 50

Gonactinia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

Gonactinia proliJera 10, 37, 49

Gonactiniidae 49

Halcampa 50

Halcampa duodecimcirrata . . . . . . . . . . .. 10, 14,25,50

Ha1campidae . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25,50

Halcampoides . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24. 50

Halcampoides purpurea 48

Halcurias 4

Halcurias pilatus .. . . . . . . . . . . . . . . . .. 10, 14, 38, 49

Ha1curiidae 49

Haliactis arctica . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Haliplanella 56

Haliplanella luciae 13, 16,34,56

Haloclava 50

Haloclava producta 10, 14,23,50

Haloclavidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 50

Hexacorallia 17

H onnathia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 52

Honnathia digitata 48, 53

Honnathia nodosa 11,15,30,52,53

Hormathiidae 52

Isozoanthus bulbosus . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Isozoanthus davisi . . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Isozoanthus ingolfi . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Kadosactis rosea ... . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Liponema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Liponema multicornis 11, 15,41,52

Mesacmaea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 50

Mesacmaea chloropsis ... . . . . . . . . . . . . . . . . . . . . .. 50

Mesacmaea laevis . . . . . . . . . . . . . . . . . . . . . . . . . . .. 50

Metridiidae 55

Metridium 55

Metridium senile 13, 16,33,55

Nematostella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 50

Nematostella vectensis 10, 14,26,50

lynantheae . . . . . . . . . . . . . . . . . . . . . . . . .. 22,37,49

Octocorallia 5

Palythoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5

Paracalliactis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 53

Paracalliactis involvens . . . . . . . . . . . . . . . . . . . . . . . . 53

Paracalliactis michaelsarsi . . . . . . . . . . . . . . . . . . . . .. 48

Paraedwardsia . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 50

Paraedwardsia arenaria. . . . . . . . . . . . . . . . .. 10, 22, 50

Paranthus 52

Paranthus rapiJonnis 11, 14, 39, 52

Parasiryonis groenlandica . . . . . . . . . . . . . . . . . . . . .. 48

Peachia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25, 50

Peachia parasitica 10, 14,25,50

Phelliactis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 53

Phelliactis americana 12, 15,30,53

Phelliactis hertwigii . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Phelliactis robusta . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Phymanthus crucifer . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Protantheae . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 37, 49

Pseudactinia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

PSP'udactinia melanaster. . . . . . . . . . . . . . . . .. 10, 14, 51

Ptychodactis patula . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Pycnanthus laevis . . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Sagartiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 54

Sagartiogeton ..... . . . . . . . . . . . . . . . . . . . . . . . . .. 54

Sagartiogeton verrilli 12, 16,36,54

Scleractinia 5

Sicyonis 52

Sicyonis ingolfi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Sicyonis obesa 11, 14, 40, 52

Sicyonis tuberculata . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

Stephanauge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 54

Stephanauge acanellae. . . . . . . . . . . .. 12, 15, 37, 42, 54

Stephanauge inornata . . . . . . . . . . . . . . . . . . . . . . . . . 48

Stephanauge nexilis. . . . . . . . . . . . . .. 12, 15,36,54,55

Stephanauge spongicola . . . . . . . . . . .. 12, 15,33,54,55

Stomphia 39, 52

Stomphia coccinea 39, 52

Tealia 51

Tealia felina val'. lofotensis . . . . . . . . . . . . . . . . . . . . . 51

Tealidium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 48

Tealidium jungerseni 48

Thenaria. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 27, 39

Urticina 7,51,52,53

Urticina crassicornis 11, 14,44,52

Urticina eques . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 51

Zoantharia 1,3,17,56

Zoan thidea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5, 17

Acknowledgments _

Preparation of this manual was supported in part by a

grant from the National Science Foundation to the

Editorial Board of the "Marine Flora and Fauna of the
Sebens: Anthozoa: Actiniaria, Corallimorpharia, Ceriantharia, and Zoanthidea 67

Eastern United States." The author thanks Frederick

M. Bayer and Steven Cairns for allowing access to the

USNM collections; ArdisJohnston for help with specimens and curating at the Museum of Comparative Zoology, Harvard University; Kathy Durante for nematocyst determinations; Mel Carriker for encouragement

and editing; Ronald W. Hardy and Sharyn Matriotti for

careful editing; J. C. den Hartog and D. G. Fautin for

careful reviews of the manuscript; and Elaine Cole,

Susan Thornton, Julia Miles, Cecile Villars, Karen van

Dersall, Trish Dahill, and Ellie Valminuto for preparation of the manuscript. Barbara Hecker (LamontDoherty Geological Observatory) supplied an excellent

recent collection of anemones procured under the auspices of the U.S. Bureau of Land Management. Ruth D.

Turner made valuable suggestions on the manuscript.

Drawings were done by Tom Siebert from specimens at

the MCZ and at the USNM and from photographs

supplied by Barbara Hecker and the author. Figures 1,

2, 3, and 30 are by the author.

Coordinating Editor _

Melbourne R. Carriker, College ofMarine Studies, University of Delaware, Lewes, DE ] 9958.

Editorial Advisers _

A. Ralph Cavaliere, Department of Biology, Gettysburg

College, Gettysburg, PA ] 7325,

Arthur G. Humes, Boston University Marine Program,

Marine Biological Laboratory, Woods Hole, MA

02543.

David L. Pawson, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian

Institution, Washington, D.C. 20560.

Kenneth P. Sebens, Department of Biology, University

of Maryland, College Park, MD 20742.

Ruth D. Turner, Museum of Comparative Zoology,

Harvard University, Cambride, MA 01002.

Robert T. Wike, Department of Botany, University of

Massachusetts, Amherst, MA 01002.

In addition to establishing the format for the "Marine

Flora and Fauna of the Eastern United States," the

Board invites systematists to collaborate in the preparation of manuals, reviews manuscripts, and advises the

Scientific Editor of the National Marine Fisheries Service.

Coordinating Editor's Comments _

Publication of the "Marine Flora and Fauna of the

Eastern United States" is most timely in view of the

growing universal emphasis on work in the marine

environment and the critical need for precise and complete identification of organisms related to this work. It

is essential, if at all possible, that organisms be identified accurately to species. Accurate scientific names of

plants and animals unlock the great quantities of biological information stored in libraries, obviate duplication of research already done, and often make possible

prediction of attributes of organims that have been

inadequately studied.

Kenneth P. Sebens is Professor of Biology and Director

of the Marine, Estuarine, Environmental Sciences

Graduate Program at the University of Maryland, College Park, Maryland. He began research on the biology

of sea anemones in 1973 as part of his Ph.D. dissertation research at the University of Washington in Seattle, where he received his degree in 1977. During

1974 to1976 he also conducted research on sea anemones and zoanthids in Panama and along the coast of

Chile. From 1977 to 1984, Dr. Sebens was on the faculty

of Harvard University and was Associate Curator of

Invertebrates at Harvard's Museum ofComparative Zoology where (with Prof. H. Levi) he effected a major

relocation, reorganization, and computer cataloging of

the coelenterate collections. In 1985 he became Director of the Marine Science Center, Nahant, Massachusetts, and then Professor at Northeastern University.

His field research from 1977 to 1998 has focussed on

the community ecology ofrocky subtidal habitats in the

GulfofMaine, the biology oftemperate zone octocorals,

and on coral reef communities in St. Croix, Belize,

Florida, and in Jamaica.
68 NOAA Technical Report NMFS 141

Published Manuals

NOAA Tech Rep.

NMFS Circular no.

Marine Flora and Fauna of the Northeastern United States

NTIS no.

Annelida: Oligochaeta Cook, David G., and Ralph 0. Brinkhurst

Protozoa: Ciliophora BOlTor, Arthur C.

Higher Plants of the Marine Fringe Moul, Edwin T.

Pycnogonida McCloskey, Lawrence R.

Crustacea: Stomatopoda Manning, Raymond B.

Crustacea: Decapoda Williams, Austin. B.

Tardigrada Pollock, Leland W.

Cnidaria: Scyphozoa Larson, Ronald j.

Higher Fungi: Ascomycetes, Deuteromycetes, and Basidiomycetes Cavaliere, A. R

Copepoda: Harpacticoida Coull, Bruce C.

Sipuncula Cutler, !':dward B.

Echinodermata: Holothuroidea Pawson, David L

Copepoda: Lernaeopodidae and Sphyriidae lJo,ju-Shey

Copepoda: Cyclopoids Parasitic on Fishes Ho,ju-Shey

Crustacea: Branchiura (;ressey, RogPr F.

Protozoa: Sarcodina: Amoebae Bovee, Eugene c., and Thomas K. Sawy"

Crustacea: Cumacea Watling, Les

Arthropoda: Cirripedia Zullo, Victor II.

Cnidaria: Scleractinia Cairns, Stephm D.

Protozoa: Sarcodina: Benthic Foraminifera Todd, Ruth, and Doris Low

Turbellaria: Acoela and Nemertodermatida Bush, Louise F.

Lichens (Ascomycetes) of the Intertidal Region 7aylor, Ronald M.

Echinodermata: Echinoidea Serafy, D. Keith, and F. Julian Fell

Echinodermata: Crinoidea Messing, Charles G., andjohn rf. Dearborn

Erect Bryozoa Ryland, John s., and Peterj. Haywm'd

Marine Flora and Fauna of the Eastern United States

Cephalopoda Vecchione, Michael, Clyde F. E. RO'per, and Mirhael j. SWl'eney

Copepoda, Cyclopoida: Archinotodelphyidae, Notodephyidae, and Ascidicolidae

Dudley, Patricia, and Paul L. Jllg

Dicyemida Short, Robert B.

Platyhelminthes: Monogenea Hendrix, Shennan S.

Acanthocephala Amin, Omar M.

374

378

384

386

387

389

394

397

398

399

403

405

406

409

413

419

423

425

438

439

440

446

NMFS no.

33

91

99

73

96

100

121

135

COM 73 50670

COM 73 50888

COM 74 50019

COM 74 50014

COM 74 50487

COM 74 51194

PB 257 987

PB 261839

PB 268 036

PB 268 714

BP 273 062

BP 274 999

BP 280 040

BP 281 969

BP 222 923

BP 285 538

BP 296 460

BP 297 676

BP 124520

BP 225 053

BP 219387

PB 124 735

PC A03/MF AOI

PB 86 ]56 395

PB9] 173013

PB 89 189583

PB 91 154179

PB 92118884

PB 95 136222

PB 98159809