Reproduction and development of the blue coralHeliopora coerulea (Alcyonaria: Coenothecalia)

Up to the present time, the reproductive cycle and larval development ofHeliopora coerulea, living fossil and sole member of the alcyonarian order Coenothecalia, have been undescribed. Populations ofH. coerulea studied on the Great Barrier Reef between 1984 and 1989 were found to have separate sexes and an annual gametogenic cycle. Mature eggs were large (>800µm diam), and their development was initiated internally. There appeared to be a lunar or semi-lunar periodicity in reproductive behaviour, with brooding of larvae commencing around either full or new moon. Development proceeded via superficial cleavage and a solid stereoblastula. Subsequent development took place on the surface of the colony, within a temporary brood-space created by protracted distension of the polyps. Larvae lacked zooxanthellae and were brooded for between 6 and 14 d prior to leaving the colony. Reproduction and development reflected the taxonomic affinities ofH. coerulea, but brooding and embryological development also appeared to be constrained by its morphology. Reproductive and larval characteristics ofH. coerulea may offer a partial explanation of the current biogeographic range ofH. coerulea.     

Bioluminescence of deep-sea coronate medusae (Cnidaria: Scyphozoa)

Bioluminescence is the production of visible light by a living organism. The light commonly appears as flashes from point sources (involving one or more cells, usually described as photocytes) or as a glandular secretion. A visible flash usually involves synchronous light emission from a group of cells or, if from a single-celled organism such as a dinoflagellate, from a group of organelles. The number of cells (or organelles) responding synchronously is the main determinant of the flash intensity. Bioluminescence is a common phenomenon in many deep-sea animals and is widespread among the Cnidaria. In this paper, we compare and contrast in situ and laboratory recordings of the bioluminescent responses of specimens of the deep-sea scyphozoans Atolla wyvillei, Atolla vanhoffeni, Atolla parva, Nausithoe rubra, Paraphyllina intermedia, Periphyllopsis braueri and Periphylla periphylla. Displays in all seven species consist of localised flashes and propagated waves of light in the surface epithelium. The first few single waves propagate at rates of up to 60 cm s^-1 but subsequent ones in any sequence of stimuli gradually decrease in speed. After several single wave responses, a subsequent stimulus may elicit multiple waves that persist for several seconds. Following such a frenzy, the specimen becomes temporarily refractory to further stimuli, but if rested will recover its normal responses and may produce further frenzies. The dome area, situated above the coronal groove, of the genera Paraphyllina, Periphylla, and Nausithoe is covered with luminescent point sources. Such point sources are generally absent from the dome of species of Atolla. Captured specimens of A. parva also produce secretory bioluminescence, corroborating prior in situ observations of this ability. Secretory bioluminescence in P. periphylla takes the form of scintillating particles released from the lappet margins. We did not observe secretory displays in specimens of any other species in the laboratory, but one instance of apparent secretory luminescence was recorded in situ in a specimen of A. wyvillei.Communicated by J. P. Thorpe, Port Erin     

Scanning and transmission electron microscope study of specialized mantle structures in dorid nudibranchs (Gastropoda: Opisthobranchia: Anthobranchia)

Mantle surfaces of the dorid nudibranchs Rostanga arbutus and Jorunna sp. were examined using scanning and transmission electron microscopy, and two specialized structures are described. These are caryophyllidia and mantle rim organs; the latter being described for the first time. Caryophyllidia occur in large numbers (several thousand) uniformly distributed over the entire upper surface, while the mantle rim organs, numbering only 40 to 60, are restricted to the upper mantle margin. Caryophyllidia are minute (40 to 50 m diam), erect tubercles supported internally by 4 to 7, vertical, calcareous spicules which emerge in a crown surrounding an apical knob. Caryophyllidia display a high level of spicular organization and incorporate a complex muscle system at their base. The apical knob is formed from specialized epidermis capping a sub-epithelial ganglion. The highly organized structure of the caryophyllidium indicates its potential importance as a new character in dorid taxonomy and phylogeny. Mantle rim organs (80 to 300 m diam) contain large numbers of vacuolated cells and cells containing pellet-shaped nodules.     

Density-dependent mortality of recruits of the abaloneHaliotis rubra (Mollusca: Gastropoda)

In three wild populations ofHaliotis rubra surveyed off southeastern Australia during the three years 1987–1989, 0 to 10% post-settlementH. rubra survived for 5 mo. Sequential censuses (February, May and December in each year) provided evidence that mortality of post-settlementH. rubra increased with the density of settlement. The occupation of cryptic habitat byH. rubra soon after settlement may well be an adaptation for avoiding predators or grazers and dislodgement by violent water movement.     

Light availability and diurnal growth of a green macroalga (Caulerpa cupressoides) and a seagrass (Halophila decipiens)

The effects of daily light period on diurnal growth patterns of a green macroalga [Caulerpa cupressoides v.lycopodium f.elegans (J. Agardh) Weber-van Bosse] and a seagrass (Halophila decipiens Ostenfeld) were investigated in Salt River submarine canyon in the US Virgin Islands in summer 1984. The daily light period, in which quantum irradiance exceeded the light saturation point for photosynthesis of the macroalga and seagrass, was manipulated in situ using lamps and shades. Plant growth was measured every 6 h for 7 d under natural and experimental daily light periods.C. cupressoides grew at the same rate day and night.H. decipiens grew more during the day than at night, a pattern that persisted under continuous light and dark treatments, indicating endogenous control of diurnal growth. Growth vs daily light period curves indicate thatC. cupressoides grew faster thanH. decipiens in short daily light periods, consistent with the observation that the macroalga penetrates to deeper water than the seagrass in Salt River canyon. Overall growth (day + night) ofH. decipiens was unaffected in lengthened light periods and reduced in shortened light periods. Chlorophyll content ofC. cupressoides was not correlated with light availability, while that ofH. decipiens was positively correlated. The alga and seagrass had different diurnal growth patterns but similar overall growth responses to daily light periods. This study shows that diurnal growth patterns are probably under endogenous control, while overall growth is a response to in situ light conditions.Contribution^#193 from West Indies Laboratory and the National Undersea Research Program     

Defensive secretions from the larvae ofApatania fimbriata (Pictet) (Trichoptera: Limnephilidae)

Summary When the larvae of the caddis flyApatania fimbriata (Pictet) are threatened, drops of fluid appear on their head capsules. The secretions are produced in a gland in the dorsal part of the prothorax. The neck region contains an eversible sac with numerous single setae, groups of 3 setae, or rows of setae on the surface. The secretion is released through two paris of orifices on the lateral sides of the sac. For the most part the secretion is composed of some 30 fatty acids, with the major components having 12–14 carbon atoms and up to 4 double bonds (approx. 1–2 µg secretion per specimen). Biotests with synthetic saturated acids (C₆–C₁₂) in a stream and in the laboratory demonstrated a paralysing effect on small invertebrate predators (Rhyacophila sp.,Plectrocnemia conspersa, Hydropsyche sp., larvae, all Trichoptera). In choice experiments,Rhyacophila sp. larvae preferred larvae ofAgapetus fuscipes andDrusus annulatus (Trichoptera) as food as compared withApatania fimbriata larvae. Larger predators, such asDinocras cephalotes (Insecta, Plecoptera) and the fishCottus gobio, did not discriminate betweenApatania fimbriata and other prey species. The use of fatty acids in defensive secretions is interpreted as an adaptation to the running water environment. They are effective repellents againstRhyacophila sp. larvae, the most important predator in the natural environment ofApatania larvae.     

An evaluation of the ABC-method (abundance/biomass comparison) as applied to macrozoobenthic communities living on tidal flats in the Dutch Wadden Sea

The ABC-method proposed by Warwick (1986) for detecting pollution effects on marine zoobenthic communities, was tested on distribution patterns of numbers and biomass among species in macrozoobenthos samples taken annually for 13 yr in a uniform way at 15 tidal-flat stations in the western part of the Dutch Wadden Sea. Along the margins of the tidal-flat area studied, where exposition to either drainage or water movements is extreme, the k-dominance plots for numbers were generally situated above those for biomass. In these areas values for biomass and species richness were relatively low and a few small-sized species such asCorophium volutator andHydrobia ulvae were very numerous. In the central part of the area, where environmental conditions are less severe, values for biomass and species richness were higher. Biomass was dominated by large adults ofMya arenaria, Mytilus edulis and/orArenicola marina. However, in only a restricted part of this area were plots for numbers consistently below biomass plots. In most of the central area high numbers ofH. ulvae occasionally occurred, causing plots for numbers to be situated above those for biomass during such periods. There appears to be no reason to interprete a high abundance ofH. ulvae as a sign of pollution or any other kind of disturbance or stress. It is concluded that the ABC method cannot be applied to tidal flat communities without reference to long-term and spatial series of control samples. In areas where such smallsized and sometimes numerous species asH. ulvae occur in strongly fluctuating numbers, the method appears useless for assessing the pollution status of a benthic community.     

Genotype-dependent fecundity and temporal variation of spawning in hybrid mussel (Mytilus) populations

Mussel samples were collected at 4 to 6 wk intervals throughout 1987 from two hybridMytilus edulis/M. galloprovincialis populations, at Croyde Bay and Whitsand Bay, in southwest England. These were analyzed at two polymorphic loci which are diagnostic for allozyme differences which typifyM. edulis andM. galloprovincialis. Dry mantle weight as a function of shell length was determined for all individuals of each sample. Size-frequency data for the two populations was obtained in September 1987 and March 1988. For all genotypes at both sites, fecundity was a function of shell length, and in both populations the frequency ofM. galloprovincialis alleles was positively correlated with shell length. At both sites, allozyme genotype explained a significant amount of variation in mantle weight either when assessed as a main effect or when assessed as an interaction with shell length or time of collection. At Croyde,M. galloprovincialis mussels had greater estimated fecundity per unit length than theM. edulis mussels. Differences in the timing of spawning activity between theM. edulis and theM. galloprovincialis mussels were inferred, and these differences might act to reduce the amount of interbreeding at Croyde. At Whitsand, a reduced level of variability in the timing of spawning activity and fecundity between the genotypes was observed and explained by a higher degree of genetic mixing. Because theM. galloprovincialis mussels had (1) a greater estimated fecundity at any length, and (2) a greater mean length than theM. edulis mussels, the mean genotypic annual fecundity perM. galloprovincialis mussel was 2.8 times greater than an individualM. edulis mussel at Croyde, and 2.2 times greater than an individualM. edulis mussel at Whitsand. This evidence thatM. galloprovincialis mussels have an advantage in fecundity, and thus perhaps in fertility, taken together with the evidence thatM. galloprovincialis also has a higher viability, indicates directional selection in favour of theM. galloprovincialis phenotype. Because of the observed temporal stability of the population it seems likely that this selection is counterbalanced by a massive imigration ofM. edulis spat from neighbouring populations.     

Reproduction of sympatric populations ofHeliocidaris erythrogramma andH. tuberculata (Echinoidea) in New South Wales

Reproduction ofHeliocidaris erythrogramma (Valenciennes) andH. tuberculata (Lamarck) was compared through examination of oogenesis, spermatogenesis and monthly measurement of the gonad index. These species occur sympatrically in the Sydney region. Their reproduction was examined at two sites near Botany Bay, New South Wales, Australia, from February 1989 through January 1990.H. erythrogramma produces buoyant, 450µm-diam eggs and the sperm have a head region 10µm in length. By contrast,H. tuberculata produces negatively-buoyant, 95µm-diam eggs and the sperm have a head region 4µm in length. Histochemical examination of the gonads revealed that periodic acid Schiff-positive (PAS +) material stored in the nutritive phagocytes appears to support vitellogenesis in both species. InH. tuberculata this material is utilized in the formation of PAS + yolk oligolecithal eggs, whereas inH. erythrogramma the PAS + material appears to be converted to lipid yolk in macrolecithal eggs.H. erythrogramma had a seasonal reproductive pattern with a 3 mo summer spawning period, whereas both populations ofH. tuberculata had a 9 mo breeding period characterized by the continual presence of nutrient reserves and vitellogenic oocytes which rapidly replaced spawned ova. Spawning ceased only for 3 mo over the summer. Due to the 9 mo spawning ofH. tuberculata it is not clear what factors serve to cue reproduction in this species.     

Bud formation and metamorphosis inCassiopea andromeda (Cnidaria: Scyphozoa): A developmental and ultrastructural study

Asexual reproduction by formation of swimming buds which metamorphose directly into polyps plays a most important role in the propagation ofCassiopea andromeda (Cnidaria: Scyphozoa). (C. andromeda polyps, originally supplied by the Löbbecke Museum and Aquarium Düsseldorf, FRG, were cultured in our laboratories.) We have defined five budding stages and investigated epithelial recruitment and dynamics during bud formation using intracellular vital stains. The region of cell recruitment was found to encircle the budding site asymmetrically. The aboral side contributing considerably less to the developing bud than the oral and lateral sides. Furthermore, it was found that the epithelial flow involved in bud formation is part of a permanent apico-basal displacement of ectodermal cells. Light and electronmicroscopic investigations revealed that no drastic changes occur at the cellular level, neither in the ectoderm nor in the endoderm which both participate in bud formation. Scanning and transmissionelectron microscopic investigations of the swimming bud revealed that the ectoderm is composed of three, and the endoderm of two, cell types. Nerve elements have been detected near the mesoglea between both ecto- and endodermal cells. Amoebocytes are regularly found either at the basis of epidermal cells or within the mesoglea, whereas symbionts are located in the endoderm. Buds induced to metamorphose by a bacterial-inducing factor were used to investigate morphological and ultrastructural changes occurring during metamorphosis and scyphistoma morphogenesis. Metamorphosis starts with the settling of a bud, followed by the formation of the pedal disk in which desmocytes, as typical cnidarian adhesive structures, are differentiated. Metamorphosis is completed with the formation of the mouth and tentacles. Whereas metamorphosis of cnidarian planulae implies considerable changes at the cellular level, tissue remodeling processes prevail in bud metamorphosis ofC. andromeda.     

Dorippids are Heterotremata: Evidence from ultrastructure of the spermatozoa ofNeodorippe astuta (Dorippidae) andPortunus pelagicus (Portunidae) Brachyura: Decapoda

Ultrastructural comparison between the sperm of the dorippid crabNeodorippe astuta (Fabricius, 1793) and the portunidPortunus pelagicus (Linnaeus, 1766) from Queensland, Australia, supports placement of dorippids with portunids and their relatives in the heterotreme section of the Eubrachyura (the Heterotremata - Thoracotremata or the Oxyrhyncha - Cancridea - Brachygnatha assemblage) and not withRanina ranina (in the Archaeobrachyura or the Oxystomata). Similarities between spermatozoa ofN. astuta and ofP. pelagicus (and other Eubrachyura) andR. ranina include: the large spherical, multi-layered, capsule-bound acrosome vesicle; the electron-dense operculum capping the vesicle; an invaginated core, or perforatorium; concentric zonation of the contents of the vesicle; a layer of cytoplasm, between the acrosome vesicle and the nucleus, which contains mitochondria (mostly degenerating) and lattice-like lamellar complexes or membrane remnants; a diffuse nucleus which is bounded externally by a combined nuclear and plasma membrane and cups the scanty cytoplasm and the large acrosome vesicle; and lateral arms into which the chromatin extends. Characteristic eubrachyuran features of theN. astuta sperm absent fromR. ranina are the long perforatorium (short and conical with a unique subacrosomal chamber inR. ranina) extending almost to the operculum; presence in the perforatorium of longitudinally arranged convoluted tubules; a zone of acrosomal rays forming the outer part of an inner dense zone; the presence of a thickened ring surrounding the basal part of the perforatorium; and, basally, two centrioles (absent fromR. ranina but also from some eubrachyurans). The sperm ofN. astuta is more similar to that ofP. pelagicus than to that of other investigated Brachyura. A heterotreme status ofN. astuta is thus unequivocally supported. Both species lack the posterior median process seen in the nucleus of majids andR. ranina.     

Pheromones inBlattella germanica andBlaberus craniifer (Blaberoidea): Glandular source,morphology and analyses of pheromonally released behaviours

Females ofBlattella germanica andBlaberus craniifer produce a volatile sex pheromone attractive at a distance for conspecific males. During the emission of the pheromone, females ofB. craniifer adopt a typical calling posture; we never observe such a stance inB. germanica females. For both species, the glandular structures responsible for pheromonal production are located on the female pygidium. InB. germanica, the thickness of the glandular epithelium is clearly correlated with the attractiveness of females, being maximum for 13 day old females. The functions of the pygidial glands are discussed in both species.     

Seasonal cycle of lysosomal enzyme activities in the mantle tissue and isolated cells from the musselMytilus edulis

InMytilus edulis L., gametogenesis takes place in the mantle at the expense of the connective storage tissue. There are two main types of storage cells: vesicular (VC) cells storing large amounts of glycogen and adipogranular (ADG) cells containing large numbers of protein granules, lipid droplets and lesser amounts of glycogen. One of the ways in which stored reserves can be mobilized for gamete formation is by controlled autophagy, in which the cellular constituents are degraded by lysosomes. Mussels were collected from the Menai Strait, North Wales, and monthly measurements made, over two years (1984–1986), of the activities of lysosomal acid hydrolases (acid phosphatase,-N-acetyl-D-glucosaminidase and-glucuronidase) and Cathepsins B and L in the mantle tissue, isolated ADG cells, low-density cells and, during spawning, in the mature oocytes of female mussels. The lysosomal proteinases, Cathepsins B, L and H, were further characterised by activation with thiol compounds and inhibition with thiol blockers and by leupeptin. Because of the low activity in the mantle tissue ofM. edulis, Cathepsin H was not assayed on a seasonal basis. There was a general increase in lysosomal enzyme activity during the winter, which can be related to increased autolysis in the storage cells and to the process of maturation in the developing oocytes. The activity of Cathepsin B was highest in the ADG and low-density cells, implying an important role in proteolysis within the ADG cells. By contrast, Cathepsin L displayed the highest activity in the mature oocytes, suggesting a major function of Cathepsin L in the development and maturation of the oocytes. Two different-glucosidase activities were measured in the monthly assays, one with a pH optimum of 4.5 (acid) and the other at pH 7.5 (neutral). Highest activities of the acid-glycosidase were found in the low-density cells, but there were no significant seasonal changes in the mantle tissue as a whole. Activities of the neutral-glucosidase were low in the ADG cells and mature oocytes, but showed high activities in the mantle tissue, with marked seasonal changes that corresponded to the mobilization of glycogen reserves in the VC cells.     

Karyotypes in two species of the genusHippocampus (Pisces: Syngnatiformes)

Diploid modal numbers of chromosomes 2n=48 forHippocampus ramulosus and 2n=44 forH. hippocampus, collected in the Gulf of Palermo in May 1985, are established. No sex-linked heteromorphic pairs are observed in the two species analyzed. The karyotype ofH. hippocampus seems to be a derived condition.     

“Glandular pockets” of the integument and feeding mechanism inPagurus bernhardus (Crustacea: Anomura)

The benthic anomuran crabPagurus bernhardus (L) were collected from the North Sea near Helgoland, West Germany, during the years 1986–1987. The histological study of the integument ofPagurus bernhardus reveals a double structure with dorsal and ventral cuticular layers and with two epithelial cell layers. Additional structures have been developed within the integument to trap micro food organisms. The structure and function of the cupshaped glandular pockets of the integument is described. The cells which secrete mucus are also described. Histological and histochemical methods have been employed to study the chemical nature of various cells of the integument and the secretions of the glandular pockets. The ecological significance of the sensory hairs, glandular pockets, cuticular layers and the supra-branchial groove is discussed with reference to the crab's feeding mechanism.     

Cytochrome P-450 monooxygenase system and benzo(a)pyrene metabolism in echinoderms

A comparative study on mixed-function oxygenase (MFO) systems was carried out on four echinoderm species: the asteroidsAsterias rubens andMarthasterias glacialis, a holothurianHolothuria forskali and an echinoidEchinus esculentus. Cytochromes P-450 andb ₅ and the MFO system-associated NADH-ferricyanide reductases NADH-cytochromec reductase, NADPH-cytochromec reductase and benzo(a)pyrene (BP) hydroxylase activities were present in microsomal fractions of pyloric caeca ofA rubens andM. glacialis and in the haemal plexus ofH. forskali. In contrast, cytochrome P-450 and BP hydroxylase activity were not detectable in the gonads ofE. esculentus. The tissue and subcellular distribution of the MFO system was studied inA. rubens. MFO system components were found in the stomachs and gonads, although detection of cytochrome P-450 in the latter tissue was difficult. Sex-related differences were not significant. The contents of the MFO system components and BP hydroxylase activities were highest in the microsomal (100 000 ×g) fractions, but MFO system components were also found in the mitochondrial (12 000 ×g) and cytosolic fractions. The BP hydroxylase activity in pyloric caeca microsomes ofA. rubens was NADPH-dependent and was inhibited by several agents known to be inhibitors of vertebrate cytochromes P-450. In the former respect, the characteristics of the MFO system were more like those of vertebrates and crustaceans than that of molluscs.     

Morphology and field behavior of pteropod molluscs: feeding methods in the families Cavoliniidae,Limacinidae and Peraclididae (Gastropoda: Thecosomata)

From 1983 to 1985 we studied the in-situ feeding behavior of fourteen species belonging to nine genera of thecosomatous pteropods, one in the family Limacinidae, seven in the family Cavoliniidae, and one in the family Peraclididae. All the euthecosomes we examined fed with a large spherical mucous web. Peraclis reticulata (D'Orbigny), however, uses a funnel-shaped mucous sheet, resembling that of other pseudothecosomes. Although the feeding mechanisms are similar in all the animals we examined closely, we found great differences in the size and morphology of the external part of the mantle. In species of Diacria and Cavolinia the external mantle attains its greatest complexity, and appears to be used for the sorting or rejection of food and for flotation. Species of the other five cavoliniid genera we studied have greatly reduced external mantles and sink slowly while feeding. Species of Limacina have no external mantle but appear to arrest sinking by setting their mucous webs. P. reticulata, which also appears to be neutrally buoyant, has an extensive external mantle that completely covers the shell. We conclude that feeding with a large mucous web explains the presence of fragments of large, fast moving prey in the stomach contents of both pseudothecosomes and euthecosomes. At present, it is impossible to decide whether thecosomes should be regarded as carnivorous trappers of prey or as suspension feeders. The use of an external mucous web for feeding is probably common to all thecosome pteropods and is the fundamental andaptation that enables them to live holoplanktonically.     

Relationship between mouthpart structures and in situ feeding habits of species of the family Pontellidae (Copepoda: Calanoida)

Pontellid copepods were collected from the surface waters of a tidal front region in the Bungo Channel (the Inland Sea of Japan) in June 1986 to examine the relationship between the morphology of cephalic appendages and gut contents. In particular, two dominant species,Labidocera japonica Mori andPontellopsis yamadae Mori, were compared in detail. Large setae on the second maxillae ofL. japonica possessed two rows of setules at right angles to each seta along its inner margin except a terminal part which was serrated, whereas the inner margin of those setae ofP. yamadae was entirely serrated. Judging from the structure of the mouthparts, especially the second maxillae, the former species seems to employ both suspension and raptorial feeding modes, in contrast to the latter, which may use only the raptorial mode of feeding. InP. yamadae, the first maxilla and the maxilliped are also modified for carnivory. Gut content analysis supported the morphological evidence for feeding differences, and revealed thatP. yamadae is a carnivore preying mainly on copepodids whileL. japonica feeds omnivorously on copepod nauplii and phytoplankton particles. Since the mouthpart structures of congeners are quite similar to each other, the feeding behavior and habits might also be similar. Within the family Pontellidae, the generaAnomalocera, Calanopia, Epilabidocera, andPontella have mouthpart structures similar to those ofLabidocera, whereas the genusPontellina resemblesPontellopsis. Morphological similarities would suggest that the first group of genera employs both suspension and raptorial feeding modes, and thatPontellina is a carnivore likePontellopsis.     

Spermatocyte chromosome banding studies inBuccinulum corneum (Prosobranchia: Neogastropoda): Variation in silver-NOR banding pattern

Diploid number (2n=72), and haploid number (n=36) forBuccinulum corneum (L. 1758) collected from the Gulf of Palermo in December 1987 were determined. A simple method to obtain nucleolar organizer regions (NOR), and constitutive heterochromatin regions (C-bands) of chromosomes ofB. corneum is described. Analyses of silver-stained chromosome preparations ofB. corneum suggest that a within-individual variability in NOR-banding pattern is present in each of the five specimens analysed.     

Functional morphology of Petricola (Rupellaria) typica (Bivalvia: Petricolidae)

The bivalve Petricola typica occurs from North Carolina (USA) to Brazil where, according to Rios (1970), its range extends to Rio Grande do Sul. The species can be found in coral blocks of Mussismilia hispida, inside colonies of Schizoporella unicornis (a calcarean bryozoan) and, more frequently, inside the sandy reefs built by the polychaete Phragmatopoma lapidosa (intertidal zone). Initial settlement probably occurs in a pre-existing cavity, which is enlarged to form a hole into which the bivalve fits exactly when its valves open. The shell, mantle, and siphons are described in detail. Special attention was paid to the ctenidia, labial palps, and stomach.