MARINE FAUNA and FLORA of BERMUDA a Systematic Guide to the Identification of Marine Organisms

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-lnterscience Publication JOHN WILEY & SONS New York * Chichester • Brisbane • Toronto * Singapore PORIFERA 111 stomulida {p. 211), Gastrotricha (p. 213), and brown are commonly caused by sym- Nematoda (p. 216), Rotifera (p. 219), biotic algae. Sponges are crustose or mas- Kinorhyncha (p. 220), Priapulida {p. 222), sive, cushion-, fan-, tube-, tree- or cup- Acanthocephala (p. 223), Sipuncula (p. shaped. Attached to the substrate, they 224), Echiura (p. 228), Pogonophora (p. show little movement except contraction of 230), Annelida (p. 232), Tardigrada (p. the entire body or of the openings upon 265), Arthropoda (p. 268), Mollusca (p. disturbance. They are most commonly 392), Bryozoa (p. 500), Phoronida (p. 516), confused with compound ascidians. Brachiopoda (p. 518), Chaetognatha (p. 519), Echinodermata (p. 522), Hemichor- Of the 4 recent sponge classes, the reef- data{p. 541), and Chordata (p. 545). The 5 dwelling Sclerospongea have hitherto not phyla not represented are the microscopic, been found in Bermuda, and Hexactinel- primitive, marine Placozoa, Nematomor- lida occur in the deep sea. Owing to many pha (nema lode-like worms with few new data on embryology, histology and marine representatives), the recently de- chemistry, the positions of higher taxa re- scribed marine interstitial Loricifera, the main in a state of llux. Of about 5,000 terrestrial Onychophora and the parasitic species known, 70 (approximately, because Pentastomida. Recent phylogenetic anal- of many uncertain identifications) are yses support the subdivision of Metazoa found in Bermuda; 49 are reported here. into Piacozoa, Parazoa ( = Porifera) and Eumetazoa (all other phyla), and the latter OCCURRENCE: Predominantly marine (3 into Coelenterata (= Radiata, i.e., Cnidaria Demospongea families in fresh water, but and Ctenophora) and Bilateria. Only the none reported from Bermuda), on stable latter term is used for classification pur- substrates, Calcarea are most common in poses here (p. 197). very shallow water; Demospongea (95% of all recent species) occur in all depths and climatic zones but, except for small, crus- Phylum Porifera (Sponges) tose and endolithic forms, avoid high- energy environments. Substrates include CHARACTERISTICS: Sedentary, aquatic, rock (particularly in caves), dead coral, filter-feeding METAZOA bounded by a 1-cell subtidal mangrove roots, sea grass layer of flat pinacocytes (pinaeoderm) and con- rhizomes, algae, other sponges, shells of taining flagellated choanocytes (choanoderm) mollusks and crabs, and artificial struc- that create a unidirectional water current, tures (buoys, pilings). Soft bottoms in calm through the body. Water enters numerous water are colonized by initial settlement on small ostia (pores) and leaves through larger rubble fragments. oscula. Mesohyle (between pinaeoderm and Collect by wading, snorkeling or choanoderm) contains various mobile cells, col- SCUBA diving (turn over rocks, look in- lagen and, usually, a skeleton of spongin, min- side caves); on deep level bottoms also by eral (silica or calcium carbonate) or both. The dredging. Cut with knife or chisel; include size ranges from millimeters to more than substrate where possible (with encrusting 1 m diameter, commonly 0.1-10 1 in or excavating forms in particular). Wear volume. Consistency varies with nature gloves for some forms are irritating to the and density of the skeleton from soft skin (by s pic tiles, toxins or epizoic cnida- crumbly to stiff elastic and stony hard. Col- rians). Commercially usable species have ors are often vivid (yellow, red, blue), par- been reported from Bermuda but are now ticularly in dark locations; shades of green very rare or absent. Other forms are under 112 KINGDOM ANIMALIA investigation for their antimicrobial prop- structures include exhalant openings (oscula), protru- erties. sions (conuli, papillae), embedded sand grains and special spiculous reinforcements (cortex). The sur- IDENTIFICATION: Some species can be face layer (ectosome) covers an internal layer identified from color, shape, consistency (choanosome) that contains skeletal material and is traversed by canal systems. Three types of canal sys- and surface structures. These characters tems are recognized in sponges; all occur in Calcarea. should be noted from fresh specimens, to- A scon is a simple lube (spongocoel) lined with gether with possible presence of pigment choanocytes (choanoderm), e.g., in Ctatkrina. Sycon, exudate or color changes shortly after named after the sponge where it occurs, has a folded collecting. Color photographs for doc- choanoderm; the choanocytes line short tubes that radiate from a common atrium, l.eucon occurs in umentation are useful if immediate many Calcarea (e.g., Leutandra) and in all Demo- identification is not possible. Presence and spongea; the choanocytes are restricted to small type of skeleton should be determined chambers that are dispersed through the thick from microscope preparations. Make 0.2- mesohyl; the choanocyte chambers are connected to 0.5 mm sections (razor blade) of dehy- each other and to the outside by a system of canals without flagellated cells. drated (alcohol or air dried) material; cut Note reticulation and structure of spongin fibers, perpendicular and, subsequently, parallel and mineral composition, size classes, position and to the surface and note whether a surface type (shape) of spicules. Collagenous spongin can oc- layer (ectosome) is detachable from the in- cur as patches connecting spicules or it can build up a ternal layer (choanosome); clear in xylene substantial elastic framework. Skeleton structure can be reticulate (net-like), with ascending primary and and mount in balsam under cover glass. connecting secondary fibers, dendritic (tree-like Alcohol-hardened material can also be branching) or intermediate such as dend roreticulate stained to make soft tissue components or pi u mo reticulate (feather-like). Spongin fibers can (e.g., spun gin, choanocyle chambers) bet- be clear or cored by a pith, by sediments, or by ter visible. A convenient stain for this pur- spicules. Spongin fibrils (In.iniu) and spongin spicules (DartiiineUa) are structures not connected to the pose is a saturated solution of basic fuchsin framework. Most sponge spicules are siliceous, except in 95% ethanol; rinse well with alcohol. in Calcarea where they are of calcium carbonate (test Spicules are isolated by digesting small but with acid!). One distinguishes megascleres (com- representative tissue fragments in cold monly >5(l p.m) that are structurally important, and sodium hypochlorite (C lor ox) or (siliceous microsclcrcs that occur tin oriented in certain parts of the tissue, Megascleres can be arranged in radiate, spicules only) boiling nitric acid; examine reticulate or felted fashion, in strands, coring (fully under high-power optics. Permanent embedded), or echinating (partly embedded in) spon- mounts of dry spicules in balsam require gin fibers. Megascleres can have I (monaxon), 3 thorough rinsing with water and absolute (triaxon) or 4 (letraxon) axes. Monaxon forms can be alcohol. Let spicules settle in test tube after pointed at both ends (oxea); pointed at one end, rounded at the the other (style); rounded at both each change (minimum 1 hr in water, 0.5 (strongyle); pointed at one, knobbed at the other (ty- hr in alcohol) to prevent loss of mi- lostyle); or knobbed at both ends (tylote). Cladolylotes croseleres. Use standard histological tech- are tylote at one end, anchor-like with recurved dads niques to determine shape, size and ar- (rays) at the other. Spitted spicules have the prefix rangement of choanocytes and choanocyte acantho- (e.g., acanthostyle). Monactines, diactincs, triactines and tetractines chambers for classification of some Kera- are radiate spicules with 1-4 rays, respectively. tosa and Calcarea. Triaenes are tetractines with 1 ray (rhabd) commonly Common sponge shapes are crustose, chambered much larger than the other 3, which can point for- (excavating), irregularly massive, spherical, tubular ward of (protriaeu) or toward (anatriacn) the rhabd. or cylindrical. Most colors are represented, but fade Desmas are irregular complex branching and inter- or change quickly after collecting (especially when ex- locking megascleres. Microscleres can be various posed to air), even before preservation. Consistency forms of asters where rays originate from 1 point can be mucous son, compressible, elastic, stiff, car- (euaster) or from an axis (strcptaster). Euasters in- tilaginous or hard and brittle. Conspicuous surface clude those with long free tapered rays (oxyasler). PORIFERA 113 with short ray and thick centrum (spheraster) and formed by some species (e.g., Utosa ruetz- with coalescenl rays and special surface ornamenta- leri, Cliona lampa). Life-span ranges from a tion (sterraster). The most common kind of strep tas- few months to about 10 yr; some very large ter has a s pi r;iled axis and is termed s pi raster. A related type, the amphiaster, has spines radiating from (0.8-2.5 m diameter) specimens of Demo both ends of the shaft. Microsderes can also be hair- spongea observed in deep reef zones, out- like (i aphid), C- or S-shaped (sigma; sigmaspire, if side the range of wave action, are es- contorted) or bow-shaped (tuxa). Another common timated to be 50-100 years old. Sponges group of microsderes arc anchors (chelae) with either can be classified as unselective suspension equal-sized (isochela) or unequal-sized (anisochcla) recurved ends. Arcuate chelae have a bow-shaped feeders. They create a unidirectional water shaft. Hexactinal spintles (e.g., hexaster) characteris- current by uncoordinated beating of tic of the class I lexactinellida have 6 rays arranged in choanocyte fiagella, and filter bacterial and $ perpendicular axes (triaxon). other cells and detrital organic particles Sizes given in plates refer to the largest dimension under 50 u,m in diameter, the maximum (height or width) of the specimen shown. size of ineurrent openings (ostia). Thirty to Fix in 10% formalin-seawater, neutral- over 100 1 water can be micro-filtered in 1 ized and buffered by me then a mine (20 g/l hr by a sponge of 1 I volume. Digestion final solution), store in 70-80% ethyl al- takes place intracellularly (phagocytosis). cohol (change twiee). Dry large specimens Many tropical and subtropical shallow- for more convenient storage but fix representative portions in liquid. Use formalin water species harbor symbiotic bacteria or Bouin's fixation and Mallory's triple (e.g., Aplysina spp., Pseudoceratina crassa) Stain for routine histological examination. and unicellular algae (zoocyanellae, e.g., Ircinia felix, Ckondrilla nucula; zooxanthel- If necessary decalcify in 5% nitric acid lae, e.g., Cliona caribbaea) that are partly (rinse well, neutralize in 5% aqueous phagocytized, partly used as a source of sodium suifate solution). For sectioning dissolved nutrients. Sponges provide sub- embed in polyester wax or epoxy resin, or strate or hiding places for many epibionts in 12% gelatin for freezing and cryostat microtomy. such as algae (jania), Hydrozoa (Sphaero- coryne), Zoanthidea (Parazoanthus), Ento- procta (Loxasomdla), Amphipoda {Caprella) BIOLOGY: Most sponges are successive and endobionts (many polychaetes and hermaphrodites, producing male and fe- crustaceans). Some of these are parasites male gametes anywhere in the endosome. or predators because they regularly feed Male specimens emit sperm through their on the tissue of the host {e.g., Alpheidae). oscula into the water column. In vivipa- Predators not living in or on sponges are rous species, which constitute the majority, many fishes and some sea turtles. Some females receive sperm through the inhal- crabs (Dromiidae, Majidae) hold sponges ant water current and incubate fertilized on their backs for camouflage without eggs until they are expelled as free- harming them except for an occasional swimming larvae. The few known ovipa- trimming. Several species are remarkable rous species release numerous eggs en- for their capacity of exacavating limestone veloped in mucous sheets for outside (Clionidae, in particular). fertilization. Several methods of asexual reproduction are common. Most species are able to regenerate from fragments. Plate 29 External budding is commonly observed (e.g., Tethya spp.), and gemmules, cell DEVELOPMENT: TWO principal types of aggregates comparable to but less complex larvae occur in Porifera; both have a than those of freshwater sponges, are ciliated anterior portion. The parenchy- 114 KINGDOM ANIMALIA

otynthus(Sycon) 1 mm

1.5 mm

2 mm parenchymula(Lissodendoryx) 0.3 mm

parenchymula(lrcinia) amphiblastula (Sycon)

29 PORIFERA: Development

mula proper to most Demospongea is oval Bermuda sponges have been treated by DE or pear-shaped, solid, 0.5-2 mm long; it is LAUB1NFELS (1950} and RUTZLER (1974). A re- comparable to the cnidarian planula. The cent monograph on a related fauna from the Bahamas, including chapters on systematic proce- amphiblastula of Calcarea is spherical, dures and an illustrated glossary, is presented by smaller {50-300 u.m), and has a central cav- WIEDF.NMAYER (1977). Modern revisions of 3 or- ity. Hexactineltid larvae, as far as they are ders of Caribbean demosponges are by SQEST (1978, known, are of the parenchymula type but 1980, in press). may not be able to swim actively. Parenchy- mula larvae, in contrast to amphiblastulae, CL. DEMOSPONGEA: Porifera with can have a high degree of differentiation siliceous spicules or spongin fibers, when released from the parent. Most commonly both, rarely neither sponge larvae swim in characteristic rotat- present. Megascleres monaxon or ing or spiraling fashion close to the bot- tetraxon. Great variety of color, tom. Settlement is preceded by a short shape and size. creeping phase. Attachment by the anterior pole occurs usually after a few hours or a day. Then the larva flattens and devel- Plate 30 ops a canal system and 1 osculum within 2 days. Higher Calcarea (e.g., Sycettida) go O. KERATOSA: Demospongea lack- through a tubular asconoid development ing proper mineral spicules. Com- stage (olynthus) shortly after settlement. monly elastic in life owing to spongin Sponge larvae are best caught by sta- skeleton. Generally in shallow water. tionary plankton nets installed near the (About 20 spp. from Bda.) bottom and facing a unidirectional current. Nearly mature larvae are often re- F. SPONGIIDAE: Keratosa with re leased by the parent sponge in the lab ticulum of primary (ascending) and under adverse conditions (e.g., heating, secondary (interconnecting, thinner) oxygen depletion). spongin fibers. Fibers without pith, some cored by moderate quantities REFERENCES: Recent comprehensive presentations of foreign inclusions (sediment of the phylum are found in BRIEN et al. (1973) and BERGQUIST(1978). Techniques for systematic and grains). Small, spherical choanocyte ecological study are outlined in RUTZLER (1978). chambers (<50 u,m diameter). PORIFERA 115

Irciniafelix (Duch. & Mich.) { — I.fas- Pseudoceratina crassa (Hyatt) ciculata sensu de Laub.): Genus ( — lanthella ardis de Laub.) (Blue with filamentous spongin threads bleeder); Dendritic knotty fibers, fiiling the choanosome, which make dark amber, to 0.5 mm thick. Sur- the sponges extremely tough and face with rounded conuli, otherwise difficult to tear; reticulated sand pat- smooth. Rubbery, firm consistency. tern on surface.—Typical form {f. Cushions or clusters of massive con- felix) of species crustose, with raised ical chimneys. Dull green, brown yel- oscula, or massive, with 1-2 mm low or golden, aerophobic (turns conules on surface. Forma fistularis bluish-purple-black when killed); (Verrill) with simple erect hollow stains fingers with purple exudate branches and terminal oscula; f. when handled. Size to 25 cm. Very acuia (Duch. & Mich.) (non /. common in shallow caves, on open strobUina) conical or massive, with rocks and reefs, 0.5-5 m. (Color Plate elevated oscula, with large (2-4 mm) 3.1.) conuli. All forms grayish to chestnut-brown, up to 25 cm. With a very F. DYSIDEIDAE: Keratosa with lam- distinctive odor when exposed to the inated primary (ascending) and sec- air. On protected sediment and rock ondary (interconnecting) spongin bottoms (f. felix, f. fistularis) and on fibers, commonly packed with sand the reefs (f. felix, f. acutd), 0.5-5 m; grains. Choanocyte chambers ovoid very common. {Color Plate 3.3.) or sack-shaped and fairly large (>50 u,m diameter). APLYSINIDAE: Keratosa with lam- Dysidea etheria de Laub.: Genus inated fibers cored by a granular with primary fibers packed, secon- pith, without foreign inclusions. daries cored by sand grains and bro- Small {<50 (xm) ovoid choanocyte ken (foreign) spicule fragments; cho- chambers. anosome also charged with foreign material. Unelastic, easily torn.— Aplysina fistularis (Pallas)( = Verongia Species occurs as crusts or lumpy- fistularis): Genus with regular hexa- cushions, to 12 cm, with small (1 gonal reticulum of amber fibers.—■ mm) conules. Two color varieties: Species fleshy spongy, with small low brilliant clear blue and grayish blue; conuli. Color deep yellow with rarely transitions. Mainly on vertical brown or greenish {shallow water) hard substrates in inshore waters, tinge; turns dark purple to black in 0.5-2 m. (Color Plate 3.4, 5.) air (aerophobic). Typical form (f. fistularis) consists of clusters of D. janiae (Duch. & Mich.) {=Des- smooth tubes; f. ansa has more irreg- macella janiae Verr.; D.fragilis forma ular tubes with short or fairly long algaft.ra de Laub.): Genus as (15 cm) and sometimes branching above.—Species as aggregates of cy- digital processes rising from the rim lindrical lobes, to 6 cm, with apical of the cylinder. Maximum size 30 oscula. Entire sponge permeated by cm. Common in all reef and open- the branching alga Jama sp. The alga lagoon environments, 1-6 m; f. replaces the spongin skeleton partly fistularis only in outer reefs, 30-40 m. or entirely; it is dead (white) in the cm

Halisarcadujardini Dictyodendrillanux Aplysilialongispina 30 KERATOSA (Sponges 1)

116 PORIFERA 117

deeper portions of the sponge, sand grains.—Species encrusting, mostly live (pinkish to greenish pur- covering up to 20 cm x 20 cm, with ple) near the surface. Sponge color small oscular chimneys. Consistency accordingly whitish, pinkish, light soft, easy to tear. Deep purplish greenish purple. Common but in- black color. Common in inshore wa- conspicuous among Jania turfs on ters, fouling on buoys, pilings and patch reefs of inshore waters, 0.5-2 similar structures, 1-5 m. m. (Color Plate 3.6.) Darwinella rosacea Hechtel: Genus DICTYODENDRILLIDAE: Kera- with spongin spicules in addition to tosa with reticulate, distinctly pithed, the fiber skeleton. Thin rose-red laminated, and dark puplish colored conulose crusts, to 5 cm. In inshore fibers and delicate, cavernous tissue. waters, 1 in; uncommon. (Color Choanocyte chambers ovoid and Plate 7.11.) large (>50 |a.m diameter). HALISARCIDAE: Keratosa in Dictyodendrilla nux (de Laub.) which the spongin skeleton is ab- { = Dendrilla mix): Primary and sent. Large (>50 p.m) sack-shaped secondary fibers with only a few choanocyte chambers. isolated foreign inclusions. Massive, to 15 cm, with finely conulose sur- Halisarca dujardini Johnston: Soft face, cavernous choanosome; very yellowish brown lumpy incrusta- soft and compressible. Dark grayish tions, 5 cm X 10 cm, on shells and blue, appearing black in the field; seagrass hlades. In protected bays, 1- bleeds clear blue pigment when han- 2 m. dled. Moderately common on inshore hard bottoms, 1-2 m,

APLYSILLIDAE: Keratosa with Plate 31 simple or branched (dendritic), pithed and laminated ascending O. HAPLOSCLERIDA: Demospon- fibers and large (>50 p,m) choano- gea with reticulate skeleton of simple cyte chambers. spicules (oxeas, strongyles). Spicules single (unispicular) or in bundles Aplysilla longispina George & Wil- (multispicular), connected by more son ( — A. sulfurea sensu de Laub.): or less spongin, or coring solid spon- Thickly in crusting, covering large gin fibers. Usually no microseleres, areas, to 20 cm x 15 cm. Surface occasionally sigmas. (About 12 spp. with numerous large (1-5 mm), slen- from Bda.) der, frequently compound conules. Soft cavernous tissue, stiff elastic F. HALICLONIDAE: I laplosclerida fibers. Bright sulfur yellow color with small (generally <150 u.m long) turns dark purple upon preserva- oxeas of uniform length, lacking mi- tion. Abundant in shallow inshore croscleres and specialized tangential caves, 1 m. (Color Plate 3.10.) ectosomal skeleton.

Chelonaplysitla erecta (Row): Ge- Reniera hogarthi (Hechtel) {-Hali- nus with neat surface reticulation of clona permollis sensu de Laub.): Ge- 118 KINGDOM ANIMALIA

nus with isodictyal {equal-sided mesh) fibers cored by robust oxeas. Den- reticulation, some multispicular dritic or frazzled fiber ends protrud- tracts (spongin restricted to spicule ing above the surface. nodes).—Species encrusting or forming clusters of vo lea no- shaped Niphates erecta Duch. & Mich. or tubular elevations to 15 cm. Very (= Haliclona variabilis sensu de soft, limp consistency. Color violet. Laub.): Cushions or single creeping Common on rubble pieces among or erect branches to 20 cm. Flush os- sea grass in protected inshore bays, cular rims ragged owing to protrud- 1-3 m. ing fiber ends. Stiff spongy, resilient consistency. Color bluish to pinkish Amphimedon viridis Duch. & Mich. lavender. Some specimens have sig- ( = Haliclona viridis): Genus with as- mas for microscleres. Common on cending multispicular tracts ce- shallow (0.5-3 m) patch reefs. Most mented by spongin, with unoriented frequently found with Parazoanthus spicules in between.—Species ap- parasiticus covering surface. (Color pears as massive cushions with raised Plates 3.12, 7.9.) oscula, as groups of conical tubes, occasionally as solid branches. Max- F. CALLYSPONGIIDAE: Haploscle- imum size 18 cm. Consistency fleshy, rida with spongin libers cored by doughy, mucous when rubbed. spicules, and with special tangential Color dull green, occasional speci- surface reticulation of primary and mens purplish brown. Very common secondary meshes. in shallow caves, on open rocks and on mud bottoms in inshore waters, Cattyspongia vaginalis (Lam.): 0.5-5 m. (Color Plate 3.9.) Clusters of large, thin-walled tubes, to 25 cm. Surface of most specimens Haliclona molitba de Laub.: Genus covered by pronounced cone-shaped characterized by reticulation of projections ("spines," conules). spongin fibers cored by spicules.— Spongy elastic, grayish green to Species amorphous, encrusting or lavender. Surface commonly col- digitate-ramose, to 10 cm, very soft, onized by Parazoanthus parasiticus. compressible and limp. Clear vivid Most abundant on outer reefs and violet color. Typically attached to inshore patch reefs, 1-5 m. seaweeds in inshore bays, 1-3 m. (Color Plate 3.8.) F. ADOCIIDAE: Haplosclerida with spicules in isodictyal reticulation and H. monticulosa (Verr.) { — Liosina ascending multispicular tracts. Spic- rnonticvlosa): Genus as above.— ules in a range of sizes. Tangential Species encrusting or digitate, to 12 ectosomal spicule skeleton present. cm. Soft spongy, bright scarlet. On rocks or, characteristically, encrust- Adocia amphioxa (de Laub.) ( = Stron- ing other sponges (e.g., Ircinia felix), gylopkora amphioxa): Genus with in shallow inshore caves, 0.5-1 m. oxeas, no microscleres.—Species as (Color Plate 3.2.) dull gray brittle cushions, to 10 cm, with elevated oscula. Moderately NIPHATIDAE: Haplosclerida with common on rocks in inshore waters, irregular reticulation of stiff spongin 1 m. ^*v3!wS Xytopsuesosburnensis Acanthacarnussouriei 31 HAPLOSCLERIDA, POECILOSCLERIDA (Sponges 2)

119 120 KINGDOM ANIMALIA

O. POECILOSCLERIDA: Demo- mangrove roots, other organisms spongea with skeleton composed of (e.g., other sponges, crabs), rock and spicules and spongin fibers. Mega- mud bottoms of protected inshore scleres (monactine and diactine, water, from low tide level to 4 m. frequently spiny) and microscleres (Color Plates 3.11, 11.6.) (varied; chelae, sigmas, microxeas and toxa; no asters). Great variety in MYXILLIDAE: Poecilosclerida shape, structure and color. (About with endosomal monactines (styles) 14 spp. from Bda.) in isodictyat reticulation, ectosomal diactines (tytotes), and isochelae and F. MYCALIDAE: Poecilosclerida with sigmas. dendroreticulate spicule strands (styles or subtylostyles), interstitial Lissodendoryx isodictyalis (Carter) anisocheles and sigmas. {Garlic sponge): Massive amorphous to lobate, with meandering Mycale microsigmatosa Arndt: surface convolutions. Size to 20 cm. Thin crusts of up to 8 cm x 15 cm Spongy compressible. Golden yel- area with meandering exhalant ca- low, frequently with tinges of bluish nals. Brick red over drab interior. green. Tendency to incorporate Coating subtidal mangrove roots; in- large quantities of foreign matter, shore waters. e.g. sand, rubble, algae, sea grass blades. Common on mangrove roots F. BIEMNIDAE: Poecitosclerida with and sediment bottoms of inshore wa- dendroreticulate spicule strands ters, 1-3 m. (Color Plate 3.7.) (styles), with sigmas and raphids. DESMACIDONIDAE: Poecilo- Biemna microstyla de Laub.: Thin sclerida with plumoreticulate ar- crusts (8 cm x 15 cm) with uneven, rangement of diactine megascleres tuherculate surface. Dull yellow to (strongyles), chelate and sigmoid yellow-orange. Common, coating microscleres. subtidal mangrove roots in inshore waters. Xytopsues osburnensis (George & Wilson): Genus packed with for- F. TEDANIIDAE: Poecitosclerida with eign materials.—Species rounded- dendroreticulate monactines (styles), massive, some oscular tubes, to 14 ectosomal diactines (ty totes) and cm. Lumpy surface with meandering roughened oxeote microscleres, subsurface canals. Endosome between spicule strands permeated Tedania ignis (Duch, & Mich.) (Fire densely with strands of the red alga sponge): Encrusting to massive, to jania. Soft, mucous, easily torn. 20 cm, with vents located on conical Color brownish purple, mottled, elevations; at some locations erect with tinges of pink and green. digitate. Cavernous, soft, easily torn. Inshore waters, moderately com- Bright red ectosome, brownish red mon, 1 m. inside. Occasionally specimens ap- pear blackish at the surface owing to CLATHRIIDAE: Poecilosclerida dense populations of epizoic Loxo- with ascending tracts of monac tines somella tedaniae. Very common on (styles, acanthostyles) that are PORIFERA 121

echinated by accessory megascleres. with spicules as strongyles. Massive, Microscleres are isochelae and toxa. with oscula on rounded lobes or low tubes, to 12 cm. Very spongy, limp, Acanthacarnus souriei Levi; Genus conn lose surface, chestnut to black- with echinating acanthostyles and ish brown. Fairly common, inshore characteristic cladotylotes {rose-stem waters, 1-3 m. spicules).—Species a thin, bright red-orange film, to 20 cm, coating and permeating substrate, possibly O. AXINELLIDA: Demospongea with boring. Common on coral rock, great variety of monaxon mega- patch reefs, 1-3 m. scleres, including acanthose forms and spongin. Microscleres absent, except in a few families. Typically with Plate 32 condensed axial, and plumose or plumoreticulate extra-axial skeleton. O. HALICHONDRIIDA: Demospon- (3 spp. from Bda.) gea with monaxonid megascleres (oxeas, styles, strongyles) and spongin, without microscleres. Endo- AXINELLIDAE: Axinellida with somal spicules in confusion (criss- styles, with or without oxeas, without cross) with ectosomal organization, microscleres. or arranged in ascending tracts. {About 5 spp. from Bda.) Homaxinella rudis (Verr.) (Red tree sponge): Genus ramose, with sim- F. HYMENIACIDONIDAE: Hali- ple spiculation of styles. Erect, tree- chondriida with ascending spicule like branching, to 12 cm, with strands. lumpy, hispid surface. Spongy, but firm elastic, slightly mucous. Clear Ulosa ruetzleri Wiedenmayer ( — Dy- red. Common on rock in shaded lo- sidea crawshayi sensu de Laub.}: cations, shallow caves, inshore wa- Genus with flat, ribbon-like dendro- ters, 0.5-2 m. (Color Plate 4.2.) reticulate spongin fibers cored by spicules; skin-like conulose ectosome Pseudaxinella explicata (Wieden- resembling keratose sponges; quan- mayer) { = P. rosacea sensu de Laub.); tities of foreign materials through- Genus massive, axial condensation out the choa no somes, including the replaced by ascending plumoreticu- fibers.—Species with spicules as long late spicule columns.—Species red styles. Encrusting to massive, to 20 orange crusts or cushions, to 10 cm, cm, very soft, compressible, limp; stiffly spongy; strong mucus produc- bright orange, with darker conspicu- tion when handled. Common on ous gemmules. On rocks, other rock in shallow caves, inshore waters, sponges {e.g., lrcinia felbc, Amphime- 0.5-2 m. (Color Plate 4.1.) don viridis), mangrove roots, in inshore waters and on patch reefs, 1-3 m. (Color Plate 4.12.) EURYPONIDAE: Encrusting Ax- inellida with principal megascleres U. bermuda (de Laub.) ( — Fibulia ber- perpendicular to the substrate, muda): Genus as above.—Species echinated by secondaries. 122 KINGDOM ANIMALIA

Homaxinella rudis

Eurypon clavatumWi Bern

32 HALICHONDRIIDA, AXINELLIDA (Sponges 3)

Eurypon ctavatum (Bowerbank): to 14 cm with meandering subsur- Genus with erect lylostyles echinated face canals; yellow-orange, reddish by acai it ho styles of 2 size classes.— orange to brownish red. Small speci- Species as very thin, hispid, orange mens very common under rocks of red encrustation of up to 18 cm. reef environments, inter tidal to 5 m. Moderately common, coaling rocks; (Color Plate 4.14.) inshore waters, 1 m. CLIONIDAE: Papillate, encrusting or massive Hadromerida, with lylostyles and spirasters or amphias- Plate 33 ters, some species with oxeas or raphids, Microscleres constitute only O. HADROMERIDA: Demospongea a small portion of the skeleton. All with monactinal megascleres (lylo- species excavate limestone, at least in the early stages of their life cycles. styles or subtylostyles, rarely styles) organized on a radial pattern. Spon- Spkeciospongia otkella de Laub.: gin present, but never as fibers. Mi- Large, massive, to 25 cm, boring croscleres, if present, astrose (asters, when young; with robust lylostyles spirasters) or oxeote (micro-oxeas). and minute spirasters and amphias- (About 12 spp. from Bda.) ters with compound spines. Grayish brown to black. Common in and on F. SPIRASTRELLIDAE: Encrusting rocks of inshore waters, intertidal to or massive Hadromerida with lylo- 2 m. Sometimes associated with styles and spirasters. The spi rasters snapping shrimps (Alpheus cylin- are mostly stout and form a substan- dricus) and barnacles (Membrano- tial part of the skeleton. balanus declivis),

Spirastrella mollis Verr. ( = S. coc- Cliona caribbaea Carter (non C. cinea sensu de Laub.): Encrusting, caribboea sensu de Laub.): Genus PORIFERA 123

Aaptos bergmanni Terpiosaurantiaca 33 HADROMERIDA (Sponges 4) mostly endolithic, with surface papil- zooxanthellae. Very common in lae; some stages encrust substrate. shells and rock, inner and outer Boring throughout life cycle.— reefs, 0.5-3 m. (Color Plate 4.11.) Species with tylostyles and 1 size class of thin wavy spirasters. Bores C. flavifodina Riitzler: Genus as small chambers, with tendency of above.—Species with tylostyles and merging inhalant and exhalant robust coarsely spined spirasters of 1 papillae to form small {3-20 mm) size class. Small (5 mm) discrete yel- crusts. Greenish, olive or brown, de- low-brown papillae, large (25 mm) pending on density of symbiotic ragged excavations filled with yellow 124 KINGDOM ANIMALIA

tissue. Very common in inshore wa- Terpios aurantiaca Duch. 8c Mich.: ters, open bays and outer reefs, 0.5- Genus characterized by tylostyles 10 m. with 3-Iobed heads. Irregularly massive, lobate, some specimens grape- C. vermifera Hancock: Genus as or finger-shaped with processes above.—Species with tylostyles and fused sideways. To 20 cm. Color yel- smooth, undulated rods as mi- low, orange or greenish blue. Green- croseleres. Vivid orange-red papillae ish-bluish tinges owing to symbiotic (2 mm), large but discrete chambers bacteria. Common on mangrove (to 8 mm). Very common in rock, in- roots and rocks in enclosed waters, shore waters and outer reefs, 1-5 m. even under low salinity conditions, 0.5-2 m. (Color Plate 4.13.) C. dioryssa (de Laub.) ( = Spirastrella dioryssa): Genus as above.—Species TETHYIDAE: Hadromerida with with tylostyles and 2 size classes of pronounced radiate structure, with spirasters. Orange to yellow-orange strongyloxeas or styles for mega- papillae or crusts (confluent papillae, scleres, asters for microscleres. 16 mm), chambered excavations {8 mm). Very common in rock, coral Tethya actinia de Laub. (Tangerine and shell, inshore waters and outer sponge): Spherical, to 5 cm diameter. With lumpy surface, buds and reefs, 0.5-5 m. (Color Plate 4.10.) attachment fibers protruding. Bright orange or green outside, dull orange C. lampa de Laub.: Genus as inside. Very common inshore and on above.—Species with tylostyles, reefs, inter tidal to 1 m. (Color Plate spined micro-oxeas and spiny mi- 4.3, 4.) crorhabds. Encrusting and boring large (to several square meters) areas of substrate, brilliant red. Less common yellow variety with same habit Plate 34 and distribution. Bears gemmules. Kills coral and clam substrates (e.g., O. ASTROPHORIDA: Demos pongea Siderastrea, Chamu), Very common at with tetractine and oxeote niega- shallow locations with strong water sclcres in some radial arrangement, currents, inshore waters, 0.5-1 m. and astrose microscleres. Either or (Color Plate 4.8, 9.) all spicule types can be lost. (5 spp. from Bda.) SUBERITIDAE: Hadromerida without microscteres. STELLETTIDAE: Astrophorida with long-shafted triaenes, and with Aaptos bergmanni de l,aub.: Genus euasters. with styles for spicules. Massive or subspherical, to 20 cm; firm consis- Myriastra crassispicula (Soil as): tency. Dark brown appearance, with With one category of oxy asters. rich yellow interior. Moderately Drab, spherical, to 5 cm; attached to common in shaded inshore habitats, pieces of rubble. Common on off- 0.5-2 m. shore secondary hard bottoms, 80 m. PORIFERA 125

Chondrilla nucula 34 ASTROPHORIDA, SPIROPHORIDA, LITHISTIDA {Sponges 5)

GEODIIDAE: Astrophorida with class only. Thickly encrusting, to long-shafted triaenes, and with ster- 15 cm, with slippery smooth sur- rasters forming surface armor. face, firm cartilaginous consistency. Brown, greenish brown (from cy- Geodia gibberosa Lam.: Encrusting anelles) to cream color, depending to amorphous massive, 12 cm. on light exposure. Very common on Smooth, partly pitted surface, leath- rocks, mangrove roots and fouling ery tough consistency. Color white to artificial structures, in caves as well dark gray, dependent on light expo- as illuminated habitats, inshore wa- sure. Moderately common but in- ters and reefs, 0.5-5 m. conspicuous under rocks or among Chondrosia collectrix (Schmidt): seaweeds, frequently overgrown by Genus without spicnles.—Species other sponges; inshore waters and reniform or lobate, cartilaginous outer reefs, intertidai to 10 m. cushions, 5 cm, off-white, gray to black. Moderately common in caves CHONDROSIIDAE: Astrophorida and under rocks, inshore waters, with reduced spicule complement. 0.5-2 m. Chondrilla nucula Schmidt (Chicken O. SPIROPHORIDA: Spherical De- liver sponge): Spherasters of 1 size mospongea with radial skeleton of 126 KINGDOM ANiMALIA

triaenes and oxeas. Microscleres are Clatkrina coriacea (Montagu) ( = Leu- contorted signias (sigmaspirae). (1 cosolenia canariemu sensu de Laub.): sp. from Bda.) Bright yellow cushions, to 8 cm, made up of trellis work of ascon TETILLIDAE: Spirophoridae with tubes with regular triactines packed characteristic inhalant and exhalant in the wall. Moderately common in depressions (porocalyces). caves, under rocks and on mangrove roots, inshore waters and outer Cinachyra alloclada Uliczka ( = C. reefs, 0.5-3 m. (Color Plate 4.6.) cavernosa setisu de Laub): Hemi- spherical or subspherical, to 8 cm, O. LEUCETTIDA: Calcarea with leu- with hispid surface caused by pro- conoid construction; choanocytes re- truding spicules. Firm consistency, stricted to chambers, with nucleus in ability to contract strongly. Yellow basal position (Only family: Leucet- color frequently obscured by tidae, with 1 sp. from Bda.) trapped sediments. Common in caves and under rocks; inshore wa- Leucetta microraphis (Haeckel) (=L, ters, 0.5-2 m. {Color Plate 4.5.) floridana) (Dead man's fingers): Irregular lobate to digitate, up to 50 O. LITHISTIDA: Demospongea with cm, fragile with rough surface. Color interlocked desmas forming a hard white to shades of pink. Two size skeleton. (2 spp. from Bda.) classes of triaxons. Very common in shallow caves inshore (1 m), also LEIODERMATIIDAE: Lilhistida from secondary hard bottoms in 80 with oxeas. m. (Color Plate 4.15.)

Leiodermatium pfeifferae (Carter) O. SYCETTIDA: Calcarea with sy- {—Azorica pfeifferae): Thin erect conoid or leuconoid canal system folded plate, 13 cm, stony hard, (choanocytes lining tubes or cham- ochreous white. Dredged twice, bers); choanocyle nucleus apical, from 800 and 1,900 m. connected to flagcllum. (About 5 spp. from Bda.)

SYCETTIDAE: Sycettida with tu- Plate 35 bular choanocyte chambers radiating from the atrium.

CL. CALCAREA: Porifera with min- Sycon ciliatum (Fabric!us) ( — Scypha eral skeleton composed of calcium ciliata): Vase-shaped, 12 mm, with carbonate. No distinction between spicule crown around osculum. megascleres and microscleres. Fragile, white. Varieties of monaxon, triaxon and tetraxonspicules O. CLATHRINIDA: Calcarea with present. Common in caves and simple tubular spongocoel lined by under rocks, inshore and near shore, choanocytes (asconoid type); cho- 0.5-1 m. (Color Plate 4.7.) anocyte nucleus in basal position (Only 1 family: Clathrinidae, with F. GRANTIIDAE: Sycettida with cho- about 3 spp. from Bda.) anocyte chambers and cortical wall. HYDROZOA 127

Leucetta microraphis 35 CALCAREA (Sponges 6)

Leucandra aspera (Schmidt) ( = Leu- explode when stimulated, discharging conia aspera): While, sack-shaped, thread-like tubes variously adapted to coil to 5 cm, tapering toward an oscular around, stick to or penetrate and poison a spicule crown. Brittle, with hispid prey organism or aggressor. There are 3 surface. Thin and thick oxeas, tri- basic and a number of subtypes of nemato- axons and tetraxons. Moderately cysts whose occurrence has significance for common in shallow inshore water, cnidarian classification (but is of only lim- 0.5-2 m. ited use for the purposes of identification within the framework of this book and K. RUTZLER therefore not considered in any detail). The presentation of the 3 classes Hy- dro zoa (p. 127), Scyphozoa (p. 155) and Anthozoa (p. 159) follows the conserva- Phylum Cnidaria tive order; more recent arguments variously propose either Scyphozoa or An- CHARACTERISTICS: Polyp- or medusa-shaped thozoa as the most primitive Cnidaria. In METAZOA of generally radial symmetry and some systems the Cu bo medusae, here with nematocysts (stinging capsules); body wall treated as an order of Scyphozoa, are of 2 cellular layers enclosing a (mostly) non- given the rank of a class, Cubozoa. celtutar membrane (mesoglea); the sole body cavity (gastrovascular cavity, or coelenteron) may be subdivided by radially arranged partitions Class Hydrozoa (mesenteries). The larva is typically a planula. CHARACTERISTICS: CNIDARIA occurring Distinguish between individuals and col- as either polypoid or medusoid stages or both, onies, the usually sedentary polypoid stage with tetramerous or polymerous radial symmetry and the usually free-swimming medu- (infrequently bilateral), non-cellular mesoglea, soid stage, and various skeletal materials and gonads that are usually epidermal. A and types. The mesoglea can be thin- stomodeum and nematocyst-bearing structures membranous, or massive-gelatinous. Ne- in the gastrovascular cavity, such as septa and matocysts, each produced by and con- gastric cirri, are lacking. Medusae are typically tained in an epidermal cell (cnidocyte), are craspedote (i.e., with a velum or shelf of tissue small egg- to spindle-shaped capsules that about the opening of the subumbrellar cavity). 128 KINGDOM ANIMALIA

Hydrozoan systematics has been com- face and are propelled by water currents plicated by separate classification systems and the wind. Several genera are known to for polyps (hydroids) and hydromedusae; have species that are luminescent. moreover, hydroid and medusa stages of a given species have often been known by About 2,000 species of hydroids are recog- different scientific names. Progress is be- nized worldwide. Of the 70 species ing made toward uniting different stages identified to date from Bermuda, 32 are of a species under a single name and in included here. eliminating the dual classification. The system used here recognizes 7 orders in the OCCURRENCE: Largely sessile and epi- class, all of which occur in Bermuda. The faunal, although representatives of the presentation as "Hydroida", "Hydro- group are found in the plankton, the neus- medusae" and Siphonophora is for conve- ton, and even the meiofauna. Bathymetri- nience only. A group of genera (Velella, cally, they occur from the intertidal zone to Porpita) often united as Chondrophora the deep sea. In shallow waters around (either as a separate order or within Bermuda, they are most diverse and abun- Siphonophora) are here considered as dant in areas swept by tidal currents or Athecata within the hydroid polyps. subjected to wave action. Some species dis- Nearly 3,000 species of hydrozoans are play a marked substrate preference, known worldwide, 160 of which have been whereas others occur on a large number of recorded from Bermuda. Of these, 70 are substrate types. Around Bermuda, hy- included here. droids commonly occur on rocks and rock rubble, algae, turtle grass, invertebrates including other hydroids, pilings, floats, HYDROIDA {Hydroid polyps) buoys and wrecks. Little is known about seasonal cycles and reproductive period- CHARACTERISTICS: (Polypoid HYDRO- icities of Bermuda hydroids. ZOA; usually colonial, berUJik and sessile; with Hydroids in shallow waters are best col- or wkftoui a medusa stage in the life cycle). From lected by snorkeling or SCUBA diving. about 1 mm to 2 m or more in height; most Collections from deeper waters may be reach a maximum size of a few centime- made using a dredge. Discovery of small ters. Although the living parts are typically species usually requires careful examina- soft and rather delicate, hydroids are com- tion of a variety of substrates in the labora- monly protected to some degree by a tory using a dissecting microscope or mag- chitinous envelope (perisarc), Hydrocorals nifying glass. Velella, Porpita, and species secrete a hard calcareous skeleton whose growing on pelagic Sargassum may be col- form superficially resembles that of the lected in neuston nets, by dipnetting or by true corals. Hydranths may be variously beachcombing. colored owing to pigment in the gastroder- Some hydroids, most notably Millepora mis, and the perisarc may be clear, golden alcicornis and Macrorhynchia philippina, are or even black. Several species are brown capable of stinging humans. owing to the presence of algal symbionts. Movements consist basically of hydranth IDENTIFICATION: Based principally upon and tentacle bending and contraction, and external characters, species determination mouth opening. Velella. and Porpita, re- requires the use of a stereoscope or micro- garded as colonies by some authors and as scope. Go nop ho res must be present or solitary polyps by others, float at the sur- medusae obtained for the identification of a

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COLOR PLATE 3: PORiFERA 1 (Sponges). 1 Pseudoceratina crassa: 2 Haliclona monticulosa; 3 Ircinia fetix; 4&5 Dysidea etherea:6Dysidea janiae; 7 Lissodendoryxisodictyalis.Q Haliclonamoiitba (with the red algae Laurencia papiitosa and Amphiroa fragilissima); 9 Amphimedon viridis: 10 Aplysitla longispina; 11 Tedania ignis; 12 Niphateserecta. (Sponsored by Tfte Trustees of the Bermuda Biological Station.) ff

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COLOR PLATE 4: PORIFEBA 2 (Sponges). 1 Pseudaxinetia explicata; 2 Homaxineila rudis; 3 & 4 Tethya actinia; 5 Cinachyra alloclada, 6 Ctathrina coriacea; 7 Sycon ciliatum; 8 & 9 Cliona iampa: 10 Ctiona dioryssa: 11 Cliona caribbaea; 12 Utosa ruetzieri; 13 Terpios aurantiaca (on /sognomon aiatus); 14 Spiras- trella mollis; 15 Leuceffa microraphis. (Sponsored by She// Company of Bermuda Ltd.)