MARINE FAUNA AND FLORA OF BERMUDA A Systematic Guide to the Identification of Marine Organisms

Edited by WOLFGANG STERRER Bermuda Biological Station St. George's, Bermuda

in cooperation with Christiane Schoepfer-Sterrer and 63 text contributors

A Wiley-Interscience Publication JOHN WILEY & SONS New York Chichester Brisbane Toronto Singapore ANTHOZOA 159

sucker) on the exumbrella. Color vari­ many Actiniaria and Ceriantharia can able, mostly greenish gray-blue, the move if exposed to unfavorable condi­ greenish color due to zooxanthellae tions. Actiniaria can creep along on their embedded in the mesoglea. Polyp pedal discs at 8-10 cm/hr, pull themselves slender; strobilation of the monodisc by their tentacles, move by peristalsis type. Medusae are found, upside­ through loose sediment, float in currents, down and usually in large congrega­ and even swim by coordinated tentacular tions, on the muddy bottoms of in­ motion. shore bays and ponds. Both subclasses are represented in Ber­ W. STERRER muda. Because the orders are so diverse morphologically, they are often discussed separately. In some classifications the an­ Class Anthozoa (Corals, anemones) thozoan orders are grouped into 3 (not the 2 considered here) subclasses, splitting off CHARACTERISTICS: Exclusively polypoid, sol­ the Ceriantharia and Antipatharia into a itary or colonial eNIDARIA. Oral end ex­ separate subclass, the Ceriantipatharia. panded into oral disc which bears the mouth and Corallimorpharia are sometimes consid­ one or more rings of hollow tentacles. ered a suborder of Scleractinia. Approxi­ Stomodeum well developed, often with 1 or 2 mately 6,500 species of Anthozoa are siphonoglyphs. Gastrovascular cavity compart­ known. Of 93 species reported from Ber­ mentalized by radially arranged mesenteries. muda, 76 are included here; the remain­ Mesoglea a mesenchymal or fibrous connective ing 17 are deep-water or rare shallow­ tissue. Adult body sizes range from water species. Scleractinia of 2 mm diameter to whip-like Antipatharia 7 m long. Colonies of OCCURRENCE: Throughout the world Scleractinia may also reach several meters oceans from the Arctic to the Antarctic, in diameter and weigh several tons. Most from the intertidal to hadal depths (10,700 anthozoans, however, measure between 3 m), and in temperatures of - 1° to 30°C. In and 50 em. Most orders possess some kind general, more species occur in the shallow­ of inorganic supporting structure, such as water tropical and subtropical regions as­ a calcareous or horny axis or microscopic sociated with coral reefs, but virtually all calcareous sclerites distributed throughout benthic marine habitats are exploited. Full the tissue. Some gorgonians and pen­ oceanic salinity is usually required, but natulids have both supporting axes and some scleractinians and gorgonians can sclerites, whereas the Actiniaria, Coral­ tolerate brackish water of 1.7% salinity. limorpharia, Zoanthidea and Ceriantharia Some zoanthids and actinians thrive in pol­ have no such supporting structure. Antho­ luted areas. Substrate type is an important zoans are among the most colorful animals factor governing distribution; some in the reef environment. Unfortunately species require mud, others sand or hard the pigments are rarely retained in pre­ substrates, and others are epizoic or served specimens and, although distinctive epiphytic. Anthozoans are usually patchy in some species, color is not usually a reli­ in distribution, often abundant in coral able taxonomic character. Anthozoans are reef habitats where conditions of substrate, predominantly sedentary animals, often light and current combine to create highly firmly attached to the substrate. The only productive areas. Densities of 27-30 col­ locomotion involved in most life cycles is onies of Scleractinia or Gorgonacea per in the planktonic larval stage. However, square meter are not unusual in reef envi- 160 KINGDOM ANIMALIA

ronments, whereas antipatharians rarely identification relies on a great number of exceed 2-3 colonies/m~. characters and methods. Most intertidal and shallow-water An­ The basic structural unit of alllhozoans is the thozoa can be collected with the aid of a polyp, which may exist alone or in a colonial aJTan~e­ hammer and chisel while snorkeling or ment, all contiguous polyps tracing their ancestry through asexual budding to a single founder polyp. SCUBA diving. A wooden spatula or flat The polyp usually has a short, squat, cylindrical body bivalve shell is good for detaching (column) with a flattened oral end (oral disc). The anemones. Burrowing actinians, cerian­ mouth occupies the center of the oral disc and is sur­ thids and pennatulids may be obtained by rounded by several rings (cycles) of hollow tentacles. sieving of sand and mud, preferably after The mouth leads through a short, longitudinally ridged tube (stomodeum) into the ~astrovascularcav­ the animals have been spotted in the ex­ ity (coelenteron). The hollow tentacles are continuous panded condition. It is best to collect with this cavity. Sometimes the stomodeum is flat­ cerianthids at night or at dusk, when most tened, with one or both of the narrow edges modified species expand. Ifkept moist, and the tem­ into flagellated grooves (siphonoglyphs) that propel perature is controlled, anemones may live water into the polyp. The ~astrovascularcavity is ra­ dially partitioned by thin lamellae (mesenteries, for several days and even survive shipment septa) on which the gonads develop. The inner, free by mail. Deep-water (over 70 m) species edges of the mesentel'ies bear distinct, thickened are usually collected fortuitously by trawl­ rims, the (mesenterial) filaments, which perform the ing and dredging, or selectively by sub­ functions of digestion, absorption and excretion. mersible. These filaments generally have the form of a simple cord, often very sinuous in the lower part of the Some anthozoans are commercially or polyp. In Ceriantharia the filaments in the upper part pharmacologically important. Some gor­ of the polyp are bilobed, in Actiniaria and Zoanthidea gonians and zoanthids contain compounds trilobed. Mesenteries (except in Ceriantharia) are ar­ (prostaglandins and palytoxin, respec­ ranged in cycles of different order. They are termed tively) that are biologically very active perfect or complete if they are attached to the col­ umn, oral disc, base and stomodeum; imperfect or and of pharmaceutical value. Care must incomplete if they do not reach the stomodeum. The be taken when collecting the zoanthid inner, free edges of the imperfect mesenteries Palythoa in late summer because it contains (mesenterial filaments) are sometimes very sinuous a highly toxic substance (palytoxin) which and trilobed. Most anthozoan polyps are bilaterally is very irritating to the skin and open cuts, symmetrical, the axis bein~ determined by the siphonoglyphs or the flattened stomodeum. Two causing severe pain and long-lasting blis­ mesenteries may occur symmetrically on both sides of ters. Actinians, some scleractinians and the axis (coupled) and/or directly adjacent (in pairs). cerianthids are popular aquarium animals. The space between the mesenteries of a pair is called Finally, because anthozoans constitute an endocoel, the space between 2 pairs an exocoel. A most of the mass of coral and patch reefs, directive mesentery pair flanks a siphonoglyph. In some elongate actinians there may be 8 well­ they contribute to storm protection, tourist developed, fertile macromesenteries alternating with recreation and sand production, not to 8 smaller sterile micromesenteries. Nematocysts, tbe mention the habitats provided for innu­ characteristic "stinging capsules" of Coelenterata, are merable other animals, many of which are found on the tentacles, stomodeum, column and eaten by humans. The taking of corals for mesenterial filaments of all Anthozoa. The fine struc­ ture of nematocysts as well as the cnidom (types of souvenirs is strongly discouraged, and pro­ nematocysts in a species or higher taxon) are impor­ hibited by law in certain areas (Coral Reef tant criteria for classification. The body wall of an Preserves). anthozoan is composed of 3 layers: an outer ectoderm (epidermis), an inner endoderm (gastrodermis) and an intermediate mesoglea, composed of a mesenchy­ IDENTIFICATION: Because of the mor­ mal or fibrous connective tissue. The mesoglea of a phological diversity within the class, colonial anthozoan is collectively termed coenen- ANTHOZOA 161 chyme. Many species contain symbiotic zooxanthellae groove (fossa) that occurs between the collar and the (dinoflagellates) that enable many Scleractinia to be base of the capitulum. The scapulus and capitulum reef~building (hermatypic) in shallow, well-lit waters; are invariably smooth, but the scapus may bear a vari­ the other, non-reef-building (ahermatypic) Sclerac­ ety of special structures, such as vesicles, hollow, tinia, i.e., without zooxanthellae, are able to live in nematocyst-bearing outgrowths of the gastrovascular dark, cold deeper waters. cavity; acrorhagi, vesicle-like nematocyst batteries oc­ The following terms pertain to features ofindivid­ curring on the parapet or in the fossa; pseudotenta­ ual orders rather than the entire class. cles, large, branched, inflated, vesicle-like structures All gorgonaceans have a central supporting struc­ bearing acrorhagus-like nematocyst batteries; warts, ture, usually called the axis in Scleraxonia and the adhesive, generally brightly colored, button-shaped central core in Holaxonia. The axis is composed of structures similar to vesicles; and tenaculi, adhesive, horny material and usually fused sclerites; the central solid mesogleal papillae also used to attach foreign core has a horny outer layer (cortex) surrounding a particles to the column. The column may also be chambered core but sclerites are not present. Sur­ pierced by tiny, regularly arranged pores (cinclides) rounding the cortex the inner and outer rinds, con­ that provide for rapid water expulsion during a sud­ centric layers of mesoglea (coenenchyme), make up den contraction and for the emission of acontia. most of the colony. Within the coenenchyme numer­ Acontia are long, spirally coiled threads generally be­ ous tiny calcareous skeletal elements (sclerites) lend low the mesenterial filaments, thought to function in support to the coenenchyme. Sclerites occur in many defense. different sizes and shapes. One of the simplest kinds The calcareous skeleton of scleractinian coral col­ is the spindle, a straight, monaxial structure pointed onies (corallum) is made up of individual units (coral­ at both ends. Scales are thin, flat sclerites and plates lites) produced by individual polyps. The calice is the are thicker scales. Monaxial, elongate sclerites that oral surface of a corallite. Some of the growth f"rms are enlarged at one end (the head) are called clubs. of colonial Scleractinia include cerioid-closely ap­ Leaf-clubs have heads ornamented with foliate pro­ pressed prismatic corallites, which usually share fused cesses whereas wart-clubs have only low, blunt pro­ walls; plocoid-slightly more widely spaced, cylindri­ tuberances on their heads. Capstans are monaxial cal corallites with separate walls; phaceloid-parallel spicules with 2 whorls of tubercles at either end and or nearly parallel, laterally free corallites; and mean­ terminal tufts. Quadriradiates, also called "butterfly droid-meandering rows of confluent corallites with spicules", have 4 terminal processes, usually radiating walls only between rows. In meandroid coralla, rows in the same plane. The portion of the expanded of corallites (valleys) alternate with slightly elevated polyp that projects above the surface of the branch is walls (collines); the colline may be grooved on top by called the anthocodium; it can be retracted into a an ambulacrum. Between each pair of mesenteries is tubular basal neck zone (calyx), which may support a a thin calcareous septum (sometimes called sclerosep­ transverse ring of sclerites (collaret). Directly above tum to distinguish it from a mesentery). Each septum the collaret are groups of vertically arranged sclerites, is composed of numerous radiating rods (trabeculae) peaking at each tentacle. The peaks are called points arranged in a plane. The trabeculae of each septum and the collaret and points are collectively termed the are usually grouped into one or more fan-shaped pat­ crown and points. Above the points are the 8 tenta­ terns (fan systems). Usually the trabeculae are parallel cles. and closely adjacent (solid septum), or there is space Pennatulaceans are composed of I large, axial pri­ between the trabeculae (fenestrate septum). Small mary polyp that consists of an anchoring peduncle rods or bars (synapticulae) may connect opposed and a distal rachis, from which the secondary polyps faces of adjacent septa. If the upper margin of a sep­ arise. Secondary polyps consist ofsiphonozooids, con­ tum projects above the calicular edge it is termed an trolling water circulation, and autozooids, the typical exsert septum. Small vertical lobes or pillars on the food-gathering polyps. lower inner edges of the septa are termed pali or Many species of Actiniaria have a muscle band paliform lobes. The columella is a calcareous axial (marginal sphincter) that encircles the upper part of structure tiJUnd in the calices of many species. It is the column. Each mesentery bears a longitudinal quite variable in shape, including twisted or straight muscular band, the retractor. Basilar muscles are rods, straight or labyrinthiform lamellae, and spongy present in species with a broadly attached pedal disc. or solid masses. In colonial Scleractinia the calcium The column can usually be divided into the scapus carbonate deposited between corallites is called (column proper) and the scapulus, a histologically dif­ coenosteum. It may be solid or vesicular; in the latter ferentiated region at the level of the sphincter. Some­ case, small thin plates called dissepiments form per­ times there is a capitular region above the scapulus, pendicular to the growth gradient, forming small separated by a fold of tissue (collar or parapet), and a cavities in the coenosteum. Dissepiments may also 0(- 162 KINGDOM ANIMALIA cur within the corallites of both colonial and solitary alcoholic suspension of spicules should be corals. pipetted onto a slide, allowed to dry, and In the Corallimorpharia, the tentacles on the oral mounted in balsam or a synthetic resin. disc are sometimes segregated into peripheral mar­ ginal tentacles and more central discal tentacles; the To identify the non-skeleton-bearing latter are often branched. Some of the unbranched Zoantharia (Actiniaria, Corallimorpharia, tentacles have knob-like tips (acrospheres) heavily la­ Ceriantharia, Zoanthidea), both the inter­ den with nematocysts. nal and external anatomy must be exam­ The mesenteries in Zoanthidea are both paired ined. This is generally done on the basis of and coupled, with alternating complete and incom­ plete mesenteries, except for the 4th through 6th preserved specimens. Menthol or mag­ mesenteries on either side of the dorsal directive. nesium chloride may be used to relax Most zoanthids have 2 consecutive imcomplete specimens prior to preservation. The gen­ mesenteries in the 5th and 6th positions (the brachy­ eral internal anatomy is best studied on the cnemic arrangement), but some have 2 adjacent, com­ basis of a transverse cut of the column. To plete mesenteries in the 4th and 5th positions (the determine the character of the (marginal) macrocnemic arrangement). Ceriantharia also have tentacles of 2 kinds: short sphincter a longitudinal cut is required in labial tentacles surrounding the mouth, and larger addition. To study the general anatomy of peripherally arranged marginal tentacles. Mesen­ Ceriantharia it is enough to make a longi­ teries are arranged in couples, not pairs, which one tudinal cut of the body. It may also be im­ after another develop in the multiplication chamber portant to examine the types and distribu­ situated opposite the single siphonoglyph. Successive couples are arranged in a regular bilateral pattern of tion of nematocysts, though this is not repeating duplets (suborder Penicillaria) or quartets necessary for identifying Bermuda species, (suborder Spirularia) of mesenteries. Mesenterial The nematocysts are prepared by making filaments may bear both craspedonemes, thread-like, a squash preparation of a tiny piece of branched, and often bunched processes, or a single (preferably living) tissue in a drop of sea­ aboral acontioid, a short, thick, acontium-like thread. The horny skeleton (axis) of Antipatharia is often water. Examine with a microscope (prefer­ covered by numerous small spines. If a colony con­ ably with interference contrast) under oil sists primarily of a single main stem it is called mono­ immersion (l,000 x ). podial but even monopodial colonies may bear pin­ The classification of the Scleractinia re­ nules (symmetrically arranged, simple or branched ramifications of smaller branch diameter). If the pin­ lies entirely on characteristics of the coral­ nule branches dichotomously once or twice, second­ lum. The tissue should be removed by ary and tertiary pinnules, respectively, result. There soaking the specimen in sodium hypochlo­ are no calcareous elements in the mesoglea. rite, followed by thorough rinsing. It may Characters used in the classification of be necessary to cut through or break a the Alcyonaria include nature of the skele­ corallum to examine its internal structure. ton if any, arrangement of polyps, branch­ Identification of the Antipatharia re­ ing pattern, terminal branch diameter, quires examination of the colony form, sometimes color and, most importantly, axis spination, and external characteristics the shape and distribution of sclerites. The of the polyp. sclerites can be easily separated from the Haematoxylin and eosin and Gomori tissue by dropping a small amount of Trichrome Stain are usually good histo­ sodium hypochlorite (full-strength com­ logical stains for anthozoan tissue. mercial bleach) onto a small piece of tissue Fixation and Preservation: In general, on a glass slide. Within minutes the or­ all anthozoans can be fixed in 6-10% for­ ganic tissue dissolves and the sclerites may malin (mixed with seawater) and pre­ be rinsed, covered with a cover slip, and served in 70% ethanol. Exceptions are Ac­ examined. For more permanent mounts tiniaria and Zoanthidea, which retain their the sclerites should be repeatedly rinsed colors better in formalin. It is usually desir­ with water in a small vial, after which an able to fix the specimen in the expanded ANTHOZOA 163

condition. For gorgonians and zoanthids a rates and longevity of anthozoans are also 7.5-8.0% solution of magnesium chloride poorly known. One tropical shallow-water in fresh water is effective for narcotization. gorgonian has an annual growth rate of The magnesium chloride is repeatedly 0.1-4.0 cm, whereas a deep-water species is substituted for half of the water in the con­ known to average 0.9 cm/yr. Branching tainer until most of the seawater has been_ reef Scleractinia may grow 4.5-27 cm/yr, replaced. The animal is then killed with massive species 0.3-1.9 cm/yr, and deep­ concentrated formalin. Menthol crystals water ahermatypes about 0.6 cm/yr. Anti­ dropped on the water surface are effective patharians can grow 2-10 cm/yr. Some in relaxing many Actiniaria and Cerian­ deep-water gorgonians and antipatharians tharia. Scleractinia are usually preserved reach sexual maturity at 12-13 yr and may dry, as are some gorgonians and anti­ live for over 70 yr. Actinians have been patharians. Bouin's fixative may be used kept in aquaria for over 70 yr, and it is not for histology; however, neither Bouin's unreasonable to surmise that some massive nor formalin should be used in cases colonial corals are centuries old. where calcareous structures are to be pre­ Anthozoans obtain nourishment by a served. variety of methods. Most are suspension feeders, preying on zooplankton and small BIOLOGY: Sexual reproduction is either crustaceans, fish, jellyfish, etc., by using dioecious or hermaphroditic; protandrous their tentacles and extruded mesenterial hermaphroditism is usual in monoecious filaments. Both structures have abundant forms. Gonads are often limited to certain nematocysts that, when discharged, are ca­ mesenteries. Fertilization is either internal pable of immobilizing and adhering to or external. Periods of sexual activity for small animals. Cilia and mucus aid in the most species are unknown. The gorgonian capture and transport of food to the Plexaura homomalla is sexually active in mouth. In this manner many anthozoans June and July; however, some tropical are also known to ingest particulate or­ Scleractinia are fertile year round, but on a ganic matter (detritus), including bacteria, lunar periodicity. All orders except Ac­ in the same manner. Dissolved organics tiniaria and Ceriantharia include species are quickly assimilated, but their role in that produce colonies by asexual repro­ nutrition is not fully understood. Some an­ duction. The original founder polyp, a thozoans may also receive nourishment product of sexual reproduction, may pro­ from a symbiotic relationship. All orders duce colonies consisting of millions of except the Pennatulacea, Ceriantharia and asexually budded polyps. When budding Antipatharia include species that possess occurs the original polyp usually remains symbiotic dinoflagellate algae (zooxanthel­ intact and one additional polyp is pro­ lae) in their endodermal tissue. These sym­ duced, either within the original tentacular biotic species are especially abundant in ring (intratentacular budding) or outside tropical reef habitats. The zooxanthellae of it (extratentacular budding). In the case receive protection and waste products of parricidal budding, however, the origi­ from the anthozoan, whereas the advan­ nal polyp is destroyed, giving rise to tages to the anthozoan are variable and not numerous daughter polyps. Although the fully understood. Zooxanthellae undoubt­ Actiniaria do not produce colonies, they edly promote calcification in Scleractinia; are capable of asexual reproduction by in general, zooxanthellae probably provide pedal laceration and longitudinal and some nourishment for the anthozoan in transverse fission. Transverse fission is also the form of dissolved organics, e.g., vita­ known to occur in Ceriantharia. Growth mins, glucose and amino acids. Some gor- 164 KINGDOM ANIMALIA gonians seem to depend entirely on nutri­ REFERENCES: For general information see HYMAN tion from the zooxanthellae; they die if (1940) and BAYER & OWRE (1968). General refer­ placed in darkness, even if provided with ences to the following orders include: Alcyonacea, Gorgonacea, and Pennatulacea (BAYER 1956; food. BAYER & WEINHEIMER 1974; CAIRNS 1976); Although formidably armed with ne­ Actiniaria and Corallimorpharia (CARLGREN 1949; matocysts for protection, Anthozoa are not DEN HARTOG 1980); Scleractinia (WELLS 1956); immune to predation. Some of the most Zoanthidea (HADDON & SHACKLETON 1891, common predators are gastropods, poly­ CARLGREN 1923); Ceriantharia (CARLGREN 1912; DEN HARTOG 1977); and Antipatharia (OP­ chaetes, echinoids, asteroids, pycnogo­ RESKO 1972). nids and fishes. For a general reference to Bermuda's Anthozoa There are numerous associations, both see VERRILL (1900a, 1901 a, 1907b). Specific refer­ parasitic and symbiotic, of Anthozoa with ences to the Bermudian fauna include Alcyonaria other animals, e. g., parasitic copepods in (BAYER 1961; VERRILL 1900a, 1907b); Actiniaria (VERRILL 1899, 1900a, 1907b); Scleractinia (VER­ most gorgonian gastrovascular cavities; RILL 1900a, 1901 b, c, 1907b; LABOREI. 1966; pycnogonid cysts on gorgonians; and Mil­ WELLS 1972); Corallimorpharia (V ERRI1.1. 1907b, lepora encrustation of branching corals. DEN HARTOG 19S0); Zoanthidea (VERRILL Other less destructive examples include as­ 1900a, 1907b); Ceriantharia (VERRILL 190Ia). Anti­ sociations of Actiniaria with ciliate proto­ patharia are previously unreported from Bermuda. zoans, amphipods, shrimps, crabs and fishes; Zoanthidea with sponges; gorgo­ S.CL. ALCYONARIA (= OCTOCO­ nians with ophiuroids and crinoids; and RALLIA): Colonial Anthozoa gorgonians and scleractinians with poly­ with 8 pinnate (feathered) tenta­ chaetes, which alter the coral branches to cles and 8 unpaired mesenteries. form their tubes. (All orders but the Stolonifera, T elestacea and Coenothecalia are represented in Bda.)

Plate 36

Plate 46 DEVELOPMENT: The fertilized anthozoan zygote undergoes cleavage, resulting in a stereo- or coeloblastula. Gastrulation pro­ O. ALCYONACEA (Soft corals): ceeds by delamination and invagination, or Alcyonaria with thick and encrust­ ingression, producing a ciliated planula, ing, lobate, or erect and arbores­ the common planktonic larva. The planula cent attached colonies consisting may drift in the plankton for weeks, dur­ of very long polyps that usually ing which time it forms its first mesen­ extend from the base of the col­ teries. A ring of tentacles is formed, usu­ ony to the uppermost branches. ally after the planula has settled. The (Of approximately 1,000 species pelagic larvae of zoanthids are called zoan­ only 14 are known from the west­ thella (or Semper's larva) and zoanthina. ern Atlantic; 1 has been collected The former usually develop into species of from Bda.) Palythoa, the latter into Zoanthus. The planulae of Ceriantharia float on the sur­ F. NIDALIIDAE: Alcyonacea with face and are therefore often collected in simple or divided, stiff, cylin­ surface plankton tows. These larvae pass drical branches. Coenenchyme through a cerinula stage of 6 mesenteries. rigid, densely packed with tuber- ANTHOZOA 165

culate spindles. Anthocodial ar­ S.O. HOLAXONIA: Gorgonacea mature in form of crown and with a distinct central axis com­ points. posed of horny material or a mix­ ture of horny and calcareous sub­ Nidalia occidentalis Gray (Dan­ stances. delion coral): Clavate colonies with polyps containing flat scales F. PLEXAURIDAE: Holaxonia with scalloped edges. Granulated with a soft, cross-chambered cen­ platelets abundant in pharyngeal tral core. Cortex of axis loc­ walls. Spindles have diameters of ulated. Sclerites often over 0.20 about 1/6 their lengths. Colonies mm long; clubs usually present. white, pale orange or yellowish Branches stout. (14 spp. from brown. Lower shelf and upper Bda.) slope depths. (Color Plate 5.12.) Plexaura homomalla (Esper) o. GORGONACEA (Sea fans, Sea (=Plexauropsis tricolor Stiasny, whips): Alcyonaria with arbo­ 1935): Genus with stout tree­ rescent (rarely lobate or encrust­ like colonies having thick, bushy ing), attached colonies consisting branches. Outer layer of rind of polyps with uniformly short with sclerites composed mostly gastrovascular cavities. Special­ of spindles and clubs; inner layer ized axial structures usually pres­ of spiny spindles that are usu­ ent. (Of approximately 1,200 ally purple.-Species with end species, 23 are known from Bda; branches 2.5-5.0 mm wide. Inner 20 are included here.) rind contains purple capstans, middle rind composed of spin­ 8.0. SCLERAXONIA: Gorgonacea dles, and outer rind with large with axial structure composed of asymmetric leaf-clubs having free or fused spicules. serrate leaves with distinct transverse collaret. Colonies to F. BRIAREIDAE: Scleraxonia with 35 em tall; dark brown. On patch axis of separate spicules; axis reefs and outer reefs. Most active perforated by gastrodermal ca­ spawning VI-VII. Growth rate: nals throughout the branches 0.13-4.2 cm/yr, average 2.0 cm/ and not separated from cortex by yr. Predators: gastropods Cypho­ boundary canals. (1 sp. from ma gibbosum (Flamingo tongue) Bda.) and Simnia; also labrid fish.

Briareum polyanthes (Duch. & P. flexuosa Lamouroux (= P. ed­ Mich.): Genus with large (up to wardsi sensu Stiasny, 1935): 0.8 mm) tuberculate spindles and Genus as above.-Species with 3-armed bodies.-Species forms end branches 2.5-4.5 mm wide. incrustations on rocks, dead Inner rind contains purple cap­ coral, and other living gorgo­ stans and short rods, middle rind nians. Polyps very large, purplish composed of short spindles, brown. At few nearshore lo­ outer rind large leaf-clubs with calities. serrate folia. Anthocodia without Briareum polyanthes ~~03mm ~ ~ ~

~ ~ 05mm i Plexaura homomalla l)1mm

46 ALCYONACEA, GORGONACEA 1 (Soft corals 1)

166 ANTHOZOA 167

collaret. Colonies to 40 cm tall. rarely over 0.4 mm long, with Color variable: yellow, brown, simple sculpture. Colonies to 100 purple and reddish purple; com­ cm tall; purple. Inner and outer monly purple in Bermuda. reefs, 5-15 m. Primarily a clear-water (patch reef and outer reef) species. P. wagenaari (Stiasny): Genus Spawning VI-VII. The dominant as above.-Species with calycular octocoral species in shallow water aperture separated by distances (to 3 m). (Color Plate 5.8-10.) greater than their own diame­ ters. Apertures 0.5-1.0 mm wide. Pseudoplexaura porosa (Hout­ Leaf-clubs of outer rind rarely tuyn) (= Plexauropsis bieolor Ver­ exceed 0.2 mm and have globose rill, 1907; P. erassa sensu Verrill, heads; spindles stout, usually less 1907): Genus with outer'rind of than 0.5 mm long, with com­ smooth leaf-clubs or smooth­ plicated sculpture. Colonies to 30 headed wart-clubs, spiny spin­ cm tall. Rose, gray, light greenish dles, and capstans, all colorless. gray or purple. Inner and outer Middle rind with white or pur­ reefs, 5-15 m. ple spindles. Polyps lacking scler­ ites and fully retractile, resulting in gaping, elliptical pores on branch.-Species with calycular Plate 47 apertures separated by distances smaller than their own diameters (hence the porous appearance of Eunicea fusca Duch. & Mich.: dry specimens!). Apertures 1.0­ Genus with tree-like colonies and 1.5 mm wide. Leaf-clubs of outer bushy branches with prominent, rind large (to 0.4 mm long) and tubular calyces. Outer rind with coarsely sculptured; spindles also small, colorless clubs and purple, large (to 1 mm long), often uni­ warty spindles; middle rind with laterally spinose. Colonies to 225 large colorless, white or purple cm tall (!), yellow, brownish or spindles.-Species with terminal reddish purple. Extremely com­ branches 3 mm or less in diame­ mon. Near shore, patch and ter. Outer rind contains small outer reefs, 1-15 m. (Color Plate (rarely over 0.1 mm) wart-clubs; 5.3,4.) middle rind has spindles about 1 mm long. Colonies rarely above P. Jlagellosa (Houttuyn) (= Plex­ 50 cm tall; gray. aura esperi Verrill, 1907): Genus as above.-Species with calycular E. tourneforti Milne Edwards & apertures usually separated by Haime (=Eunieeopsis atra Verrill, distances greater than their own 1901 a; Eunieeopsis tourneforti diameters. Apertures 0.5-1.0 mm sensu Verrill, 1907): Genus as wide. Leaf-clubs of outer rind above.-Species with outer rind rarely exceed 0.1 mm in length containing leaf-clubs and wart­ and have r~lUnded folia, usually clubs to 0.15 mm long; middle not globose; spindles slender, rind with stout, long (to 1.5 mm) 0.1 mm 0.3mm 1.5mm E. calyculata

47 Eunicea, Plexaurella (Soft corals 2)

168 ANTHOZOA 169 spindles. Colonies broad, can­ rods only 0.05-0.07 mm long. delabrum-shaped. The typical Rind containing numerous stout form has terminal branches lO­ tri- and quadriradiates both to 15 mm in diameter and well­ 0.5 mm long. Colonies to 100 em developed lower calycular lips. E. tall; yellowish brown. On outer, lourneforli f. alra Verrill, 1901 a, inner and patch reefs. has terminal branch diameters of 6-10 mm and poorly developed P. nutans (Duch. & Mich.): lower calycular lips. Colonies to Genus as above.-Species has 61 em tall; dark gray, blackish strongly armed polyps with stout brown, or black. Most common rods about 0.3 mm long. Rind on outer reefs and reef slope. containing tri- and quadri­ (Color Plate 5.15, 16.) radiates with slender arms, length of sclerites to 0.45 mm. E. clavigera Bayer: Genus as Colony sparsely branched, to 100 above.-Species with outer rind em tall; gray or brown. On inner containing small leaf-clubs about and outer reefs. 0.1 mm long; middle rind with very long (to 3 mm) spindles. Colonies straggly, not candela­ brum-shaped. No collaret. Col­ Plate 48 onies rarely larger than 50 em; brown. Muricea laxa Verrill: Genus E. calyculata s. s. (Ellis & Solan­ with arborescent, densely der) (=E. grandis Verrill, 1900a; branched colonies. Branches Euniceopsis grandis sensu Verrill, hard and prickly, with numer­ 1907): Genus as above.-Spe­ ous, close-set, tubular or shelf­ cies with outer rind containing like, lower calycular nms small wart-clubs, rarely longer (edges).-Species with calyces than 0.15 mm; middle rind with having long spindles that are large (to 2 mm) white spindles re­ spiny proximally and smooth dis­ sembling rice grains. Colonies tally, forming prominent termi­ not candelabrum-shaped. Dis­ nal spikes in the calyx. Branching tinct collaret present. Branches lateral; end branches long, thin long and stout, 8-16 mm in diam­ (2-3 mm in diameter) and flex­ eter. To 1 m tall; yellowish ible. Colonies 25-30 em tall; gray, brown. On patch reefs and near­ bluish white or yellow. Usually shore; never abundant. found in deeper water of outer reef, but sometimes as shallow as Plexaurella dichotoma (Es­ 2 m. (Color Plate 5.5-7.) per): Ge~1Us with rind contain­ ing numerous quadriradiates M. muricata (Pallas): Genus as ("butterfly spicules"). Erect, di­ above.-Species with tuberculate chotomously branched colonies outer rind spindles lacking of thick, furry appearance. Calyc­ smooth terminal spikes. Branch­ ular apertures slit-like.-Species ing lateral in 1 plane; end with weakly armed polyps, the branches short and thick (4.5-6.0 170 KINGDOM ANIMALIA

M mUricata

M. atlantica

48 Muricea (Soft corals 3)

mm in diameter). Axis conspicu­ Pseudopterogorgia americana ously flattened at points of (Gmelin) (= Gorgonia americana branching. Rarely over 30 em sensu Verrill, 1907): Genus tall; white, beige or yellow. Com­ with pinnate branching, twigs mon on patch and outer reefs. round or slightly flattened; branches do not anastomose. M. atlantica (Kiikenthal) (=M. Two types of sclerites predomi­ muricata sensu Verrill, 1907): nate: straight spindles and C­ Genus as above.-Similar to shaped scaphoids.-Species very above species, except that axis is slimy to the touch when alive not flattened at points of branch­ (when dead, the mucus usually ing. Also, outer rind spindles causes the branches to stick to­ sometimes have strong spines on gether). Scaphoid sclerites tuber­ one side. Rarely over 50 em tall; culate on the concave side and white or yellow. Outer reefs. echinulate on the convex side. (Color Plate 5.13, 14.) Colonies to 100 em tall; pale yel­ low or light purple. Common on inner and outer reefs.

Plate 49 P. acerosa (Pallas) (Sea plume): Genus as above.-Species not F. GORGONIIDAE: Holaxonia slimy in life. Scaphoids weakly with a soft, cross-~hamberedcen­ curved, more or less smooth on tral core. Sclerites rarely over the convex side. Colonies to 180 0.15 mm long; no clubs. Cortex em tall. Color of live colony light of axis slightly loculated. (3 spp. purple, purple-red or light yel­ from Bda.) low becoming white upon drying. ANTHOZOA 171

"..oJ '0(0 r\ I· . !"irI' . ~~., T'\ Ellisella' barbadensis,9

6ocm) ~ ~Ii

O.18mm ~\ O.05mm J Pseudopterogorgia acerosa ,~J (:.,j~i.:i.JV· 49 GORGONIIDAE, ELLISELLIDAE (Soft corals 4), PENNATULACEA (Sea pens)

Mostly nearshore; often host to do not anastomose. Slit-like caly­ snails (Cyphoma) , the shrimp To­ ces arranged along the 2 narrow zeuma and brittle stars. edges of the branches. Scaph­ oids, spindles, and anthocodial Pterogorgia citrina (Esper) rods present in coenenchyme. (=Gorgonia citrina sensu Verrill, Colonies to 45 cm tall; yellow, 1900a, 1907): Branching lat­ sometimes with purple edges. eral; end branches slender, stiff, Primarily inshore. (Color Plate and strongly flattened. Branches 5.1, 2.) 172 KINGDOM ANI MALIA

Gorgonia ventalina L. (= G. fla­ F. KOPHOBELEMNIDAE: Sessi­ bellum sensu Verrill, 1907): Ii florae with bilaterally oriented Branches anastomose, forming polyps on rachis, leaving a dorsal uniplanar, reticulate, fan-shaped streak bare. Colonies clavate, colonies. Branches round or with a well-developed axis. slightly compressed in the plane of the fan. Very small calyces Sclerobelemnon ef. theseus Bayer: located in 2 rows along edges of Slender, elongate, slightly clavate branches. Scaphoids, spindles, colonies with aut6zooids ar­ and anthocodial rods present in ranged in about 9 irregular lon­ coenenchyme. Colonies to 180 gitudinal rows. Sclerites are small em tall and ISO em wide; purple, oval or slightly constricted rarely yellow or brown. Re­ platelets with serrated ends, and stricted to outer and patch reefs: flat scales resembling double­ (Color Plate 5.17.) bitted axe heads. Colonies to IS em tall; orange or yellow. At F. ELLISELLIDAE: Holaxonia depths of 50-60 m. (Color Plate with a solid, calcified, flexible, 5.11.) horny core. Spicules characteris­ tically dumbbell-shaped or clubs. S.CL. ZOANTHARIA (= HEXACO­ (3 spp. from Bda.) RALLIA): Colonial or solitary Anthozoa usually with more than Ellisella barbadensis (Duch. & 8 simple (not pinnate) tentacles. Mich.): Whip-like, unbranched Mesenteries both paired and un­ colonies with biserial or multiple paired. (All recent orders of this lateral bands of calyces. Whips to subclass are represented in Bda.) 2 m long and usually less than 8 mm in diameter; vermilion red O. ACTINIARIA (Sea anemones): when alive. Most common on the Solitary Zoantharia with hex­ fore-reef slope (50 m and more). amerously arranged mesentery (Color Plate 14.5.) pairs. After first 6 pairs of mesenteries, additional pairs are O. PENNATULACEA (Sea pens): formed in all exocoels. Each ten­ Alcyonaria without branches and tacle corresponds to one inter­ not firmly attached to substrate. mesenterial space. No skeleton. Primary polyp elongate proxI­ (Of approximately 800-1,000 mally, forming a stalk which an­ species III this order, 17 are chors the colony in mud. Second­ known from Bda, IS are Ill­ ary polyps originate from distal eluded here. Of the 5 super­ rachis. (Of approximately 300 families, the Abasilaria and described species, only I IS Mesomyaria are not represented known from Bda.) in Bda.)

S.O. SESSILIFLORAE: Pennatulacea with secondary polyps arising di­ Plate 50 rectly from rachis, not united near their bases by ridge-like or SUP.F. BOLOCEROIDARIA: Actinia­ leaf-like structures. na without marginal sphincter ANTHOZOA 173

muscle. Column divided into pseudotentacles that bear con­ scapus (often bearing vesicles) spicuous, semiglobular, white to and scapulus. Acontia absent. bluish nematocyst batteries. Scapus and pseudotentacles usu­ ally brown (zooxanthellae). Oral BOLOCEROIDIDAE: Boloce­ F. disc has 96 tentacles, speckled roidaria with tentacles separated with fine white dots (nematocyst from the coelenteron by a thin, batteries). Diameter of pedal disc centrally perforated diaphragm to 5 em, that of the expanded and a sphincter, which cause crown of pseudotentacles often the tentacles to break off easily more than 25 em. Found in reef at this partition. Basilar muscles habitats, the base attached in absent. (2 spp. from Bda.) holes or cracks among coral and rocks. Expands pseudotentacles Bunodeopsis antilliensis Duerden during daytime (photosynthesis), (=B. globulifera Verrill, 1900a): and tentacles at night (feeding). Scapus provided with sessile or A severe stinger. In deeper wa­ stalked vesicles of variable shape ters (10m) often associated with and development (small and sim­ the shrimp Thor amboinensis. ple to large and highly com­ (Color Plate 9.11.) pound), and variegated in color, usually green, brown or white. Scapulus, oral disc, and tentacles SUP.F. ACONTIARIA: Actiniaria with smooth and transparent. Mouth a mesogleal sphincter. Acontia occasionally surrounded by a always present. blue ring. With 20-40 tentacles, variable in length and densely F. AIPTASIIDAE: Acontiaria with speckled with fine white dots a very weak sphincter. Column (nematocyst batteries); usually divided into scapus and scapu­ expanded only at night. Diame­ Ius. Scapus smooth, with cin­ ter of base rarely exceeds 15 mm. clides. Tentacles often long. Six, Common on dead coral, living rarely 8 pairs of perfect, fertile sponges, mangrove roots, and mesenteries. (2 spp. from Bda.) Thalassia leaves. With zooxan­ thellae. Asexual reproduction by pedal laceration common. A Bartholomea annulata (Lesueur) strong stinger. (Color Plate 6.1.) (=Aiptasia annulata sensu Ver­ rill, 1900a, 1907) (Ringed anemone): Tentacles with dis­ F. ALICIIDAE: Boloceroidaria tinct white to faint blue, incom­ with tentacles in open connection plete annulations. Common in with the coelenteron. Basilar bays, inlets and on coral reefs, muscles weak. (1 sp. from Bda.) among rocks, stones and corals. Also common in mangrove areas, Lebrunia danae Duch. & Mich. but usually not on the mangrove (Brown anemone; B Gill-bearing roots. No asexual reproduction. anemone): Scapus smooth, with With zooxanthellae. (Color Plate 4-8 highly expandable, dendritic 6.5.) 176 KINGDOM ANI MALIA

bottoms. Usually found In antilliensis) (B Dark-star anem­ crevices or under stones. (Color one): Column smooth, low, Plate 6.11.) fawn colored to reddish brown. Variable number (sometimes none) ofcream colored acrorhagi on parapet. Oral disc with 30-200 Plate 51 tapering tentacles, usually with acute tips. Oral disc with a dis­ tinct, stellate pattern of alternat­ SUP.F. ENDOMYARIA: Actiniaria ing pale and dark stripes. No with an endodermal, occasionally directive mesenteries or si­ reduced sphincter. Acontia ab­ phonoglyphs. Diameter of base sent. varies from 0.5 to 4.0 cm. Subtid­ ally and intertidally on exposed F. ACTINIIDAE: Endomyaria shores, in pools, on and under with a distinct sphincter (rarely stones and in holes. A small form reduced) and with an adherent of this species occurs on floating pedal disc with well-developed Sargassum. Asexual reproduction basilar muscles. Column divided by longitudinal fission common. into scapus and a narrow ca­ (Color Plate 6.3.) pitulum. More than 6 pairs of mesenteries are perfect. (8 spp. Condylactis gigantea (Weinland) from Bda.) (=G. passifiora sensu Verrill, 1901 a; Ilyanthopsis longifilis sensu Actinia bermudensis (McMur­ Verrill, 1907) (B Purple-tipped rich) (B Red anemone): Col­ anemone): Column smooth, umn smooth and low, deep red short-cylindrical to trumpet­ to brownish. Fossa contains 0-24 shaped; color bluish gray, yellow­ blue, globular acrorhagi. Oral ish or brick-red. Acrorhagi ab­ disc with 96-140 rather short, sent. Up to 150 long, tapering, acute tentacles. Tentacles with­ greenish or brownish tentacles out pattern and often more with a conspicuous design of brightly colored than the rest of pale, densely arranged, ruffte­ the body. Two pairs of directive like striae. Tentacle tips blunt or mesenteries and 2 distinct si­ slightly swollen, often rose or phonoglyphs. Diameter of base purple (less frequently blue or to 4 cm. Intertidally in holes and bright green). Diameter of ex­ under stones, often in rather panded oral disc and tentacles to exposed localities. Viviparous; 30 cm; of base to 8 cm. Juveniles asexual reproduction by longi­ often with knobby tentacles. tudinal fission has occasionally Common in shallow water (less been reported. (Color Plate 6.2.) than 10m) usually on coral reefs, rubble flats and Thalassia fields. Pseudactinia melanaster (Verrill) With zooxanthellae. Often associ­ ( = Anemonia elegans Verrill, ated with shrimp (e.g., Pericli­ 1901a, 1907; Actinia melanaster menes anthophilus, Thor amboinen­ sensu Verrill, 1901 a, 1907; sis), crabs (e.g., Stenorhynchus Anemonia sargassensis; Anemonia seticornis, Mithrax spp.), and a va- Actinia bermudensis

Anthopleura carneola

51 ENDOMYARIA (Sea anemones 2)

177 178 KINGDOM ANI MALIA

riety of fishes (e.g., juvenile adhering to the warts. No asexual wrasses, Apogon spp.). (Color reproduction. Plate 9.12.) Bunodosoma granuliferum (Le­ Anthopleura carneola (Verrill) sueur): Column short and cy­ (= B unodactis stelloides var. car­ lindrical, completely covered with neola Verrill, 1907; A. varioar­ small vesicles and marked by a mata): Genus with acrorhagi pattern of alternating pale and and scapus covered with adhe­ dark longitudinal bands. Parapet sive warts. Sphincter often cir­ modified into small, marginal cumscript.-Species with dirty­ lobes that often bear cream or green, pink or wine-colored col­ pinkish, globular acrorhagi on umn, which is entirely covered their aboral face. Oral disc with with bright green, yellowish· or 96 (rarely more) brown, orange, reddish warts. Lower part of col­ purple to almost black tentacles umn often pale. Variable num­ with opaque, grayish or white ber of cream colored to faint red­ cross bars above. Diameter of dish acrorhagi in the fossa. Oral base to 5 em. Found intertidally disc with 30-60 short, acute tenta­ on exposed shores, but usually in cles, with or without a faint pat­ sheltered niches (under stones, in tern. Diameter of base to 2 em. holes); not common. (Color Plate Common in the upper parts of 6.12.) the intertidal zone under stones, in holes or crevices. Columnar Actinostella flosculifera (Lesueur) warts often encrusted with shell ( = Phyllactis ftosculifera; Asteractis fragments. Asexual reproduc­ expansa; Actinactis fiosculifera tion by longitudinal fission. sensu Verrill 1900a; Asteractis ftosculifera sensu Verrill 1899, A. catenulata (Verrill) (= Ac­ 1907) (Sand anemone): Col­ tinoides pallida sensu Verrill umn elongate when expanded, 1900a; Bunodactis stelloides var. with a broad, grayish to yellowish catenulata Verrill, 1907): Cenus green, disc-shaped collar, pro­ as above.-Species with elongate vided with numerous, radially ar­ milk-white to pale pink colored ranged, short, often branched column. Warts of the same colors protuberances. Scapus often with restricted to the upper part of a flamed pattern of cream and the column. Oral disc with 20-43 bluish gray or red; the upper tentacles, which have a conspicu­ scapus is darker and provided ous color pattern. Oral disc usu­ with longitudinal rows of distinct ally with a conspicuous green warts. Oral disc with 48 tentacles, ring around the mouth. Diame­ often green, yellow or red with ter of base to 1.5 em. Common cream colored crossbars above. under stones and in crevices in Expanded collar to 10 em across; tide pools; also found in sand, base to 4 em. With zooxanthellae. the base attached to buried On sandy flats, in bays and inlets, stones. Upper part of column of­ and on reefs; rarely seen. Often ten covered by shell fragments buried in the sand, the base at- ANTHOZOA 179

tached to a solid substrate, the itary or colonial Zoantharia very expanded collar resting on the similar to the Actiniaria but with sand. Viviparous. a calcareous skeleton. (Of ap­ proximately 2,500 living species, F. PHYMANTHIDAE: Endomy­ 34 are known from Bda; 25 are ana with or without a weak included here. All 5 suborders sphincter and with an adherent are represented in Bda.) pedal disc with well-developed basilar muscles. Column not di­ vided into regIOns. Oral disc wide; divided into a distinct, pe­ Plate 52 ripheral, tentaculate region and a central region with radial rows of wart-like protuberance's (discal S.O. ASTROCOENIINA: Scleracti­ tentacles). (l sp. from Bda.) ma with septa composed of a few simple trabeculae. Corallites Epicystis crucifer (Lesueur) small (less than 3 mm in diame­ (= Phymanthus crucifer) (B Cross­ ter); polyps rarely with more barred anemone): Column than 12 tentacles. Colonial. short and trumpet-shaped, with a flamed pattern of cream and red. F. ASTROCOENIIDAE: Astrocoe­ Numerous longitudinal rows of niina with beaded septal margins 3-6 conspicuous purple warts in and poorly developed coeno­ the marginal regIOn, each row steum. Phaceloid to cerioid col­ ending m a conical marginal onies formed by extratentacular wart. Tentacles short, up to 384 budding. (l sp. from Bda.) in number. Oral disc to 15 cm across; base to 8 cm in diameter. Two varieties occur in Bermuda: Stephanocoenia michelinii Milne a form with elevated crossbars on Edwards & Haime (= Plesiastrea the tentacles and with predomi­ goodei Verrill, 1900a, 1901 b, nantly gray (sometimes yellow, 1907) (B Small-eyed star coral): green or dark purple) oral disc Corallum massive, cerioid to and tentacles (Color Plate 6.9); plocoid, often encrusting. Calices and a form with smooth tenta­ 2-3 mm in diameter, containing cles, a brown oral disc, and ten­ 24 septa. Twelve paliform lobes tacles with yellow to faint or­ arranged before the 1st and 2nd ange, radial-longitudinal stripes. cycles of septa. Columella sty­ (Color Plate 6.10.) Intermediates liform. Brown. Rare on outer also occur. With zooxanthellae. reefs; more common in open, in­ In shallow water on reefs, rocky shore waters; 2-5 m. shores and stony-sandy flats. Of­ ten in sand, the base attached to F. POCILLOPORIDAE: Astrocoe­ stones; uncommon. niina having septa with smooth mner margms or reduced to o. SCLERACTINIA (= MADRE­ spmes. Well-developed, solid PORARIA) (Stony corals): Sol- coenosteum. Plocoid and usually 180 KINGDOM ANI MALIA

52 ASTROCOENIINA (Stony corals 1)

ramose colonies formed by ex­ Plate 53 tratentacular budding. (3 spp. from Bda.) S.O. FUNGIINA: Scleractinia with septa composed of numerous trabeculae in fenestrate arrange­ Madracis decactis (Lyman) (B ment. Synapticulae present. Ten-rayed star coral): Genus Corallites usually larger than 2 with well-developed septa ar­ mm in diameter; polyps usually ranged in groups of 6, 8 or 10. with more than 12 tentacles. Col­ Columella styliform and promi­ onial and solitary. nent.-Species with encrusting, massive, nodular or clavate coral­ F. AGARICIIDAE: Fungiina with lum. Calices about 2 mm in diam­ eter, with 10 septa each. Pali ab­ solid septothecal walls. Solitary or colonial, the latter condition sent. Green or brown. Common resulting from intratentacular on inner reefs and in inshore wa­ budding. (l sp. from Bda.) ters, especially on vertical reef edges; 1-4 m. Agaricia fragilis Dana (B Hat coral, Shade coral): Corallum M. mirabilis (Duch. & Mich.): pedicellated, with broad (rarely Genus as above.-Species with over 15 cm in diameter), thin, bushy colonies, to 2 m in diam­ saucer-shaped fronds. Calices eter. Branches 5-9 mm in diame­ only on upper side of frond. Up ter, with blunt tips. Calices 1-2 to 24 septa per calice. No col­ mm in diameter, with 10 septa umella. Chocolate- or purple­ each. Pali absent. Pale cream to brown. Very common in shaded bright yellow. Very common in areas such as shallow caves; inner 0.5-6.0 m, especially on the verti­ and outer reefs, and inshore wa­ cal edges of inshore reefs. ters; 1-15 m. Agaricia fragilis

S. siderea Siderastrea radians

53 FUNGIINA (Stony corals 2)

181 182 KINGDOM ANI MALIA

F. SIDERASTREIDAE: Fungiina irregular, stout branches. Calices with slightly porous, synap­ about 2 mm in diameter; septa ticulothecate walls. Usually colo­ poorly defined. Light brown to nial, formed by both extra- and purple. Common in open in­ intratentacular budding. (2 spp. shore waters; 0.5-3.0 m. from Bda.) P. astreoides Lamarck: Genus Siderastrea radians (Pallas) : as above.-Species with cerioid, Genus with extratentacular bud­ flat to hemispherical corallum, ding and cerioid corallum.­ often covered with small bumps. Species with spheroidal or hemi­ Calices about 1.5 mm in diame­ spherical corallum to 30 cm in ter. Septa poorly defined. Yel­ diameter. Calices 2.5-3.5 mm in lowish brown. Very common in diameter, with less than 48 septal­ inner and outer reefs and open calice. Inner edges of septa al­ inshore waters; 0.5-15 m. (Color most vertical. Greenish to brown. Plate 6.14.) Common on inner and outer reefs and in inshore waters, in- S.D. FAVIINA: Scleractinia with cluding mud flats where speci- septa composed of numerous mens may be partially buried in trabeculae in laminar arrange­ the mud and periodically ex- ment; septal edges dentate. Syn­ posed at low tide; 0-10 m. apticulae absent. Corallite diame­ ter usually greater than 2 mm. S. siderea (Ellis & Solander): Colonial and solitary. Genus as above.-Species with encrusting or hemispherical, cerioid coralla to 100 cm in di­ ameter. Calices 4-5 mm in diame­ ter, with 50-60 septa. Inner edges Plate 54 of septa slope gently (about 45°) toward the papillose columella. Light reddish brown. On inner F. FAVIIDAE: Faviina with ex­ and outer reefs and in inshore sert septa composed of 1 or 2 fan waters; fairly common; 0-10 m. systems producing a more or less regularly dentate inner margin. F. PORITIDAE: Fungiina with Colonies formed by both intra­ extremely porous walls and and extratentacular budding. septa. Septa composed of 3-8 (All 5 spp. from Bda included.) loosely united trabeculae. Colo­ nial, formed by extratentacular Favia fragum (Esper) (B Small budding. (2 spp. from Bda.) star coral, Golf ball coral): Small, plocoid, pebble-like coralla Porites porites (Pallas) (= P. poly­ or encrustations, usually less than morpha Verrill, 1901b): Genus 10 cm in diameter. Corallites with 2 cycles of septa (12) and mono-, di-, or tricentric. Calicu­ closely united corallites.-Spe­ lar diameter 5-6 mm. Yellowish cies with cerioid, ramose coral­ brown. Common on inner reefs, lum consisting of thick clumps or in open inshore waters, and in 15cm

D.labyrinthiformis

M. cavernosa

54 FAVIIDAE (Stony corals 3)

183 184 KINGDOM ANIMALIA

tide pools; 1-6 m. Planulation Vl­ M. cavernosa (L.) (= Orbicella VIII. cavernosa sensu Verrill, 1900a, 1901b, 1907) (Great star coral): Diploria strigosa (Dana) (=Mean­ Genus as above.-Corallum of dra cerebrum sensu Verrill, 1901b, species shaped as massive boul­ 1907) (Brain coral): Genus with ders, platy fronds, or encrusta­ meandroid, sinuous series of tions to 200 cm in diameter. corallites. Paliform lobes absent. Calices 5-11 mm in diameter; 48 Trabecular linkages between septa/calice. Columella well de­ corallite centers.-Species with veloped. Brown or green. Re­ hemispherical, spheroidal, or en­ stricted to the open clear waters crusting coralla to 200 cm in di­ of the inner and outer reefs; 3-30 ameter. Valleys about 6 mm m. (Color Plate 6.13.) wide; collines not grooved. 'With 15-20 septa/em. Yellow to green­ ish brown. Very common in in­ ner and outer reefs and even in Plate 55 muddy bays; 1-8 m.

D. labyrinthiformis (L.) F. RHIZANGIIDAE: Faviina with (=Meandra labyrinthiformis sensu septa composed of 1 fan system Verrill, 1901b, 1907) (Brain that produces an irregular septal coral): Genus as above.-Spe­ margin. Colony formation by cies with hemispherical coralla extratentacular, stoloniferous reaching 200 em in diameter. budding, with individual polyps Valleys about 5 mm wide; col­ often losing their original con­ lines distinctly grooved by am­ nections. (3 spp. from Bda.) bulacra. With 14-17 septa/em. Yellow or brown. Very abundant Astrangia solitaria (Lesueur): on the outer reefs and some in­ Solitary corallites or quasicol­ shore water; 1-30 m. onies formed by asexual budding from basal stolons. Stolons often Montastrea annularis (Ellis & subsequently disrupted, giving Solander) (= Orbicella hispidula; the appearance of exclusively sol­ Orbicella annularis sensu Verrill, itary corallites. Corallites cylin­ 1900a, 1901b, 1907) (Small star drical, 5-20 mm tall, 3-4 mm in coral): Genus with plocoid diameter. With 36 septa; no coralla and costate coenosteum. paliform lobes. Common but Columella spongy.-Corallum of cryptic; found on undersurfaces species shaped as massive boul­ of rocks or other corals; 1-3 m. ders, upright pillars, or encrusta­ Ahermatypic. (Co~or Plate 6.7.) tions to 200-300 em in diameter. Calices 2-4 mm in diameter; 24 Colangia immersa Pourtales: septa/calice. No paliform lobes; Solitary corallites or quasi­ columella well developed. Yel­ colonies formed by asexual bud­ 10wish to brown. Very common ding from a common basal sto­ on inner and outer reefs and in lon. Corallites cylindrical, to 10 open inshore waters; 1-30 m. mm tall, 6-7 mm in diameter. ".i'if., , O. valenciennesi

?~:'.".~..". 4'

55 RHIZANGIIDAE-MUSSIDAE (Stony corals 4)

185 186 KINGDOM ANIMALIA

With 48 septa; those of 1st cycle 40 cm tall. Branches long, 12-20 much more exsert than others. mm in diameter. Calices low, of­ Twelve paliform lobes. Green, ten sunken in the coenosteum, with clear tentacles. Rare; found surrounded by a low ridge. on undersides of platy corals; 3­ Calicular diameter 3-5 mm. Yel­ 95 m. Ahermatypic. low. Common in some inshore waters; 2-20 m. F. OCULINIDAE: Faviina with exsert septa composed of 1 fan F. MEANDRINIDAE: Faviina system that produces a smooth or with exsert septa composed of 1 minutely dentate septal margin. fan system that produces a Coenosteum dense. Colony for­ smooth or finely dentate septal mation by extratentacular bud­ margin. Well-developed en­ ding. (4 spp. from Bda.) dothecal dissepiments. Solitary, or colonial by intratentacular Oculina diffusa Lamarck (Ivory budding. (2 spp. from Bda.) coral, Bush coral): Genus with well-developed columella of M eandrina meandrites (L.): twisted processes. Pali arranged Massive, rounded or flat boul­ in an irregular crown before first ders reaching to 100 cm in di­ 1 or 2 cycles of septa. No axial ameter (rarely more than 30 cm corallites.-Species have ramose, in Bda); smaller coralla unat­ compact colonies rarely exceed­ tached. Meandroid. Septa 6-81 ing 30 cm in diameter. Branches cm, with smooth inner edges. to 10 mm in diameter. Calice 3-4 Columella lamellar. Yellow, mm in diameter; 24 septa/calice. brown, or white. Moderately Pale yellow. Very common on in­ common in open, clear waters shore reefs and some inshore wa­ such as outer patch reefs; 1-10 m. ters, especially in areas of high sedimentation; 1-3 m. Dichocoenia stokesi Milne Ed­ wards & Haime: Heavy, O. varicosa Lesueur (Large ivory rounded or flat coralla reaching coral, Tree coral): Genus as to 50 cm in diameter. Calices above.-Species with open, ir­ mono- or polycentric, arranged regularly branched, arborescent in a V-shape or in long (to 50 coralla to 40 cm tall. Branches mm) meandroid valleys. Septa long, crooked, to 50 mm in diam­ 101cm, with smooth inner edges. eter. Calices usually raised on Columella trabecular. Yellow or mounds. Calicular diameter 3-4 brown. Rare; on outer reefs; 7-10 mm. Rare; found in some In­ m. shore waters; 12-25 m. F. MUSSIDAE: Faviina with ex­ O. valenciennesi Milne Edwards sert septa composed of more & Haime (Ivory coral, Tree than 2 fan systems that produce coral): Genus as above.­ large, coarse, septal dentations. Species with open, irregularly Well-developed endothecal dis­ branched arborescent colonies to sepiments. Solitary, or colonial by ANTHOZOA 187

mm

, /-----' Coenocyathus goreaui Guynia annulata Rhizopsammia bermudensis 56 CARYOPHYLLIINA, DENDROPHYLLIINA (Stony corals 5)

intratentacular budding. (2 spp. Plate 56 from Bda.) S.D. CARYOPHYLLIINA: Sclerac­ Isophyllia sinuosa (Ellis & Solan­ tinia with septa composed of nu­ der) (=1. multiflora Verrill, merous trabeculae in laminar ar­ 1901b; Mussa anectens Verrill, rangement; septal edges smooth. 1901c, 1907; Mussa roseola Ver­ Synapticulae absent. Corallite di­ rill, 1907; Mussa dipsacea sensu ameter usually greater than 2 Verrill, 1907; Mussa fragilis sen­ mm. Mostly solitary. su Verrill, 1907; I. fragilis sensu Verrill, 1901b; I. dipsacea sensu F. CARYOPHYLLIIDAE: Caryo­ Verrill, 1901b) (Rose coral): phylliina with solid septothecal Small meandroid coralla usually walls. Solitary or colonial by in­ less than 20 em in diameter. tra- or extratentacular budding. Septa 7-9/cm, with inner edges (Of 7 spp. known from Bda the bearing coarse teeth. Columella only shallow-water species is in­ spongy. Valleys 20-25 mm wide. cluded here.) Form multiflora is smaller, with narrower valleys (12-15 mm Coenocyathus goreaui Wells: wide) and has 11-12 septa/em. Small bushy colonies of inter­ Both forms occur in a variety of twined cylindrical corallities. colors: green; white, lavender, Corallites to 20 mm long and 4-6 brown and variegated. Very com­ mm in diameter. Usually 32 septa mon on outer reefs and in in­ arranged octamerally. Eight pali. shore waters; 0.5-5 m. (Color Pale pink. Rare, known only Plate 6.8.) from cavities in reef rock. Aher­ matypic. Scolymia sp.: Solitary, subcylin­ drical, firmly attached coralla to 7 F. GUYNIIDAE: Caryophylliina em in diameter. Calice round, with epithecal wall penetrated by with 4 or 5 cycles of septa. Septa regular rows of pores. Exclu­ and costae coarsely dentate. Col­ sively solitary and ahermatypic. umella large and trabecular. (1 sp. from Bda.) (The specimens collected from Bda are too small to identify as to Guynia annulata Duncan: Cor­ species.) Rare, on outer reefs. allum cylindrical, attached ba- 188 KINGDOM ANI MALIA

sally or along its side. Extremely tractile tentacles, bearing con­ small; calicular diameter 1 mm, spicuous, globular acrospheres. lenght to 10 mm. Septa 12 or 16. Spirocysts always present. (l sp. Columella consists of one twisted from Bda.) lath. Inconspicuous, and ex­ tremely rare in shallow water. Corynactis parvula Duch. & Known depth range 3-653 m. Mich. Qewel anemone): With 50-100 tentacles arranged in ra­ S.O. DENDROPHYLLIINA: Scler­ dial rows of 3-7 tentacles each, actinia with irregularly perforate which increase in size toward the septa composed of numerous margin. Outer tentacles longest, trabeculae; septal edges smooth. often exceeding the diameter of Synapticulae present. Corallite the oral disc. Diameter of base diameter usually greater than 2 and oral disc about 5-8 mm, that mm. (Contains only I family, of the expanded crown of tenta­ Dendrophylliidae, and only 1 sp. cles to 25 mm. Color variable: is known from Bda.) body often orange to brown, ac­ rospheres bright orange to red. Rhizopsammia bennudensis Wells: On reefs, under stones and in Small colonies, formed by bud­ holes among dead coral; rare. ding of corallites from an en­ Often several specimens are clus­ crusting base. Corallites cylin­ tered as a result of asexual repro­ drical, 6-8 mm in diameter, 5-10 duction by pedal laceration. mm tall. Septa 48/calice. Septa (Color Plate 7.5.) and theca porous. Bright orange. Rare; known only from cavities F. DISCOSOMATIDAE: Cup- or in reef rock. Ahermatypic (Color disc-shaped, often firm Coral­ Plate 6.6.) limorpharia with simple or den­ dritic discal tentacles lacking acrospheres. Tropical shallow­ water forms, invariably with Plate 57 zooxanthellae. (2 spp. from Bda.)

Discosoma sanctithomae (Duch. & O. CORALLIMORPHARIA (= AS­ Mich.) (=Actinotryx sanctithomae CLEROCORALLIA) (Coral anem­ sensu Verrill, 1900, 1907; Rho­ ones, False corals): Solitary or dactis sanctithomae): Genus as colonial Zoantharia with hex­ the family.-Species with oral amerously arranged mesentery disc to 5 em in diameter, and with pairs. More than one tentacle a narrow, naked marginal zone, corresponds to each inter­ generally bordered by distinct mesenterial space. No skeleton. but tiny marginal tentacles which (Of approximately 35 spp. world­ often have white acrospheres. wide, 3 spp. are known from Discal tentacles usually well de­ Bda). veloped and distinctly dendritic. No more than 3-7 discal tentacles F. CORALLIMORPHIDAE: Cor­ in the principal radial rows, the allimorpharia with simple, re- total number not exceeding 300. 57 CORALLIMORPHARIA (False corals), ZOANTHIDAE (Sea mats)

189 190 KINGDOM ANIMALIA

Column greenish or brown; ten­ Palythoa variabilis (Duerden) tacles often iridescent; inside of ( = Protopalythoa grandis Ven-ill, stomodeum white. Very common 1900): Genus with encrusted on and around coral reefs, often mesoglea and single mesogleal forming extensive, brightly col­ sphincter muscle.-Species with ored mats. (Color Plate 7.12, 13.) polyps that may arise singly or be connected by a lamellar basal D. carlgreni (=Rhodactis coenenchyme in groups of4 or 5. (Watzl) 2 carlgreni; Paradiscosoma carlgreni): Large colonies cover to 0.5 m of Genus as above.-Species with a substrate. Length of column and smaller and more rigid body than width of oral disc of large polyps D. sanctithomae. No naked mar­ are 5 cm and 4 cm, respectively. ginal zone. Marginal te.ntacles Column light brown to white, de­ generally smaller and the margin pending on the degree of encrus­ often drawn out into small, blunt tation and concentration of zoo­ or trifid lobes. Discal tentacles xanthellae. Oral disc dark brown, wart-like to short and denaritic, light brown, slightly greenish or up to 20 in the principal radial variegated, with large white rows. Total number of discal ten­ areas. Tentacles light brown, tacles may reach over 1,000. numbering up to 82, but usually Color variable: variegated, green about 68. Common on rocks in with brown, orange or red; some shallow water; occasionally found discal tentacles may be conspicu­ in small colonies at depths to 30 ously white or yellow; inside of m. Often found with Zoanthus stomodeum usually yellow. On sociatus. (Color Plate 7.3,4.) and around coral reefs; uncom­ mon. (Color Plate 7.6, 7.) P. mammillosa (Ellis & Solander) (= P. grandiflora Verrill, 1900a, 1901a, 1907): Genus as above.­ O. ZOANTHIDEA (Colonial anem­ Species with variable colony form ones, Sea mats): Solitary and depending on habitat. On rocks colonial Zoantharia with paired exposed to wave action, the but no hexamerously arranged coenenchyme is about 3 cm thick; mesenteries. All mesentery pairs colonies are 8-12 cm across, and after the first 6 are formed in the polyps are separated by channels ventro-lateral exocoels. No skele­ about 1 cm wide. On sand, the ton, but sometimes calcareous coenenchyme is usually not thick­ encrustation in mesoglea. (Of ap­ er than 1 cm and the polyps are proximately 300 spp., 9 are spaced several centimeters apart, known from Bda; 7 are included usually with only the oral discs here.) visible. Expanded adult polyp 15 mm long with a 13 mm wide oral F. ZOANTHIDAE: Zoanthidea disc. Column and disc ocher, with 1 or 2 mesogleal sphincter never variegated. There are usu­ muscles and a brachycnemic ar­ ally 44-48 tentacles in fully grown rangement of mesenteries. (7 polyps. Moderately common on spp. from Bda.) rocks in shallow water. ANTHOZOA 191

P. caribaea Duch. & Mich. (= P. mid-column. Column light mammillosa sensu Verrill, 1901 a): green, often bluish towards the Genus as above.-Species usu­ distal end. Common in most of ally forming extensive colonies the shallow-water bays, some­ with numerous closely arranged times intertidally. Often found polyps. Proximal ends of polyps with Palythoa variabilis. (Color joined in a basal coenosteum Plate 7.1.) about 1 cm thick. When fully ex­ panded, the distal portions of the Isaurus duchassaingi (Andres): polyps rise about 4 mm from the Polyps occur singly or in small basal coenenchyme but when clusters. Column usually bent, contracted the surface of the col­ with tubercles on the convex side ony is almost flat. The colony is and a smooth concave side. Dur­ light ocher. Number of-tentacles ing the daytime the oral disc and increases with body size (from 20 tentacles are normally infolded, to 44). Common on most patch leaving a small aperture at the reefs and nearshore, often found center of the closed oral disc. growing on dead areas of coral Length and width of the column heads. (Color Plate 7.8.) of a large solitary specimen are 7 cm and 1 cm, respectively. Clus­ Zoanthus sociatus (Ellis & Solan­ tered polyps are smaller. Column der) (= Z. proteus Verrill, 1900) light brown, orange, white or (Green sea mat): Polyps squat, chartreuse, sometimes translu­ elongate, or trumpet-shaped. cent because of very thick non­ Basal coenenchyme laminar, cov­ encrusting mesoglea. Oral disc ering the surface of the substrate and tentacles light brown. Large to which the colony is attached, polyps generally have 44-46 ten­ or reticulate, composed of a net­ tacles. Filamentous green algae work of stolons that joins the and other debris often adhere to colony on several separate sub­ the upper warty surface of the strates. A typical colony covers column. On reefs and rocks; 0-20 2 about 10 cm . Mesoglea lacks en­ m; very rare. (Color Plate 7.2.) crustations. Length of column and width of oral disc of ex­ F. EPIZOANTHIDAE: Zoanthi­ panded polyp are 6 mm and 3 dea with a single mesogleal mm, respectively. Color variable, sphincter muscle, macrocnemic but most polyps have a bluish arrangement of mesenteries, and green or yellowish green oral encrusted body wall. (1 sp. from disc. The pigment of the disc is Bda.) often arranged in a series of con­ centric circles and there may be a Epizoanthus minutus Duerden: dark triangular patch at each cor­ Colonies vary in size, but usually ner of the slit-shaped mouth. 10-20 polyps joined by thin sto­ Tentacles light green, usually 46­ lons constitute 1 colony. Length 50. Column smooth, but often of column and width of oral disc polyps at edge of a colony have of a large, fully expanded polyp numerous protuberances on the are 10 mm and 4 mm, respec- 192 KINGDOM ANIMALIA

tively. Color of polyp dependent all subsequent mesenteries are on the type of mesogleal encrus­ formed in multiplication cham­ tation, but usually light brown. ber opposite siphonoglyph. Two The oral disc is darker brown tentacles correspond to each in­ with a few radiating white lines. termesenterial space. No skele­ Tentacles are brown with dark ton; tube dwellers. (Of approxi­ brown bands and white tips. mately 100 species, 1 adult sp. is Adult polyps have 32-40 tenta­ known from Bda.) cles. Sphincter muscle weak, located in several small cavities s.O. PENICILLARIA: Ceriantharia near the center of the mesoglea. with mesenteries arranged in Found attached to the undersur­ duplets. Craspedonemes absent. face of rubble in shallow wave­ Older, fertile mesenteries bear exposed locations; rare. EColor acontioids. (Only I family, Plate 7.11.) Arachnactidae, in the suborder.)

F. PARAZOANTHIDAE: Zoan­ thidea with a single endodermal Arachnanthus nocturnus den sphincter muscle, macrocnemic Hartog (= Cerianthus natans? arrangement of mesenteries, and Verrill, 1901a): Tube en­ encrusted body wall. (l sp. from crusted with sand or gravel. Body Bda.) to 30 cm long, yellowish brown (sometimes with dark brown Parazoanthus parasiticus (Duch. streaks). Upper part of body usu­ & Mich.): Polyps solitary or col­ ally white. Marginal tentacles onial, joined in groups of 3 or 4 brown with 3-6 pale crossbars on by thin stolons. Length of col­ the upper surface and fine green umn to 4 mm, width of oral disc streaks between their insertions. to 4.5 mm. Column encrusted Labial tentacles pale brown, with­ with fine calcareous and siliceous out crossbars. Siphonoglyph con­ materials that give it a greenish nected with about 1/3 of the total white color. Oral disc and tenta­ number of mesenteries. In shal­ cles light brown. Up to 28 tenta­ low, sandy bays (often just below cles per polyp. In shallow water tide level) where its tubes are on reefs and in enclosed bays; attached to stones. Locally com­ exclusively on sponges, such as mon, but easily overlooked be­ Niphates erecta and Cally.lpongia cause it is usually retracted into vaginalis. (Color Plate 7.9.) its tube during daytime. Juve­ niles, often without a tube, may be found under stones. (Color Plate 7.10.) Plate 58 O. ANTIPATHARIA (Black cor­ als): Colonial Zoantharia with O. CERIANTHARIA (Tube anem­ 6, 10 or 12 unpaired mesenteries ones): Exclusively solitary Zo­ and 6 tentacles. Internal axis antharia with unpaired mesen­ keratin-like. Colonies firmly at­ teries. After first 6 mesenteries, tached to substrate. (Of approx- ANTHOZOA 193

A. tanacetum

58 CERIANTHARIA (Tube anemones), ANTIPATHARIA (Black corals)

imately 175 spp., 5 are known pie or pinnulate with simple or from Bda; 4 are included here.) bifid spines. Polyps oval.-Spe­ cies with fan-shaped colonies to F. ANTIPATHIDAE: Antipatha­ 40 cm tall. Branches not pinnu­ ria with simple (non-pinnate), late. At 40-60 m, growing up­ non-retractile tentacles. right on a gently sloping bottom; not uncommon. Antipathes furcata Gray: Ge­ nus with sparsely to densely A. hirta Gray (=A. picea): Ge­ branched colonies. Branches sim- nus as above.-Species with 194 KINGDOM ANIMALIA

sparsely branched, monopodial have been chosen to illustrate 5 of the 6 colonies to 80 cm tall. Pinnules orders. arranged biserially in 4-6 longi­ tudinal rows. Tertiary pinnules OCCURRENCE: Marine; some species In rare. Not uncommon at 40-60 m brackish water. The great majority is on gently sloping bottom. pelagic, drifting in swarms with surface currents; some representatives of the or­ A. tanacetum Pourtales: Genus der Platyctenida are adapted for benthic as above.-Species with a mono­ and even parasitic life and found on en­ podial colony with pinnules ar­ crusting algae, mangrove roots, sedentary ranged in 4-6 rows along the animals and other substrates. length of the axis. Tertiary pin­ Collect small forms with a plankton net nules common. Not uncommon outfitted with a large-volume bucket; to below 50 m. obtain intact specimens of the more fragile species, individuals should be caught in Stichopathes lutkeni Brook: glass jars by SCUBA. Monopodial colony without pin­ nules (a naked rod). Polyps ar­ IDENTIFICATION: Only observation in the ranged on 1 side of the axis open water can fully reveal the delicate throughout its length. Colonies beauty of these organisms. Species are straight, but sometimes coiled identified under the dissecting micro­ near free end. To 4 m long but scope, preferably live, in a glass dish. not more than 1 cm in diameter. Note the 2 planes of symmetry (one through the tentacles, the other perpendicular). The generally 8 Most common between 30 and 60 rows of combs (plates made of fused cilia) converge, m in areas with great vertical re­ on the aboral pole, toward the statocyst to which they lief. are connected by delicate rows of cilia. The move­ ment of the combs always starts at, and is directed S. CAIRNS, J. C. DEN HARTOG toward, the aboral pole, propelling the animal ARNESON ROTZLER C. & K. through the water mouth first. Most species have I or [corrected after publication] more pairs of delicate tentacles, beset with adhesive bodies (colloblasts), with which they catch prey; in Phylum Ctenophora (Comb-jellies) Cydippida these retract into tentacular sheaths. The mouth opens into a stomach from which a system of canals extends into the tentacular sheaths and under CHARACTERISTICS: Extremely transparent, the comb rows; the latter canals also house the gelatinous, mostly pelagic, bisymmetric gonads. In Lobata, the laterally compressed body METAZOA with 8 rows of comb-like ciliary bears 4 short auricles and 2 often extensive lobes that plates; without nematocysts. Generally 1-10 are used in locomotion; in the ribbon-shaped Cestida, 4 comb rows are reduced. The benthic Platyctenida cm (rarely to 150 cm). Most species are so are extremely flattened, and have reduced comb transparent that only the iridescent flashes rows. emanating from the beating combs reveal Fix Cydippida and Beroida, preferably their presence. after anesthesia with magnesium chloride, in Flemming or a mixture of mercuric The phylum is thought to be an early chloride (100 g) and glacial acetic acid (3 offshoot from the ancestral medusoid ml) in seawater (300 ml). After 15-30 min, cnidarian. Of about 90 species known, carefully replace fixative with fresh water, many of the pelagic ones are cosmopoli­ and transfer to 70% alcohol by stages, tan; 5 species from the Bermuda region starting with 30%. Most Lobata and Ces-