Variability in the chemical defense of the sponge Chondrilla nucula against predatory reef fishes

Chondrilla nucula is a common Caribbean demosponge that grows in a range of habitats, from coral reefs to mangrove swamps. On reefs, C. nucula grows as a thinly encrusting sheet, while in mangrove habitats it surrounds submerged mangrove roots as fleshy, lobate clumps. Previous feeding experiments using predatory reef fish revealed a high degree of variability in the chemical defenses of C. nucula. The present study was undertaken to determine whether a relationship exists between habitat, growth form, and chemical defense of C. nucula. Both laboratory and field feeding-assays of crude extracts confirmed that C. nucula possesses a chemical defense with high intercolony variability, but there was no significant variation in feeding deterrency between reef and mangrove habitats at either geographic location (Bahamas and Florida). Extracts of C. nucula collected during September and October 1994 from the Bahamas were significantly more deterrent than those collected during August 1993, May 1994, and May 1995 from Florida, and extracts of these spring and summer Florida collections were more deterrent than extracts of C. nucula collected in December 1994 and February 1995 in the same locations. There was no evidence that deterrent compounds were concentrated in the surface tissues of the sponge, or that chemical defense could be induced by simulated predation. Laboratory and field assays of the fractionated crude extract revealed that feeding deterrency was confined to the most polar metabolites in the extract. Field transplants were used to determine whether predation influenced the growth form of C. nucula. Uncaged sponges transplanted from the mangrove to the reef were readily consumed by spongivorous reef fishes. Lobate mangrove sponges became thinner after being caged on the reef for 3 mo, but encrusting reef sponges did not become thicker after being caged in the mangroves for the same period of time. Reef sponges that were caged for 3 to 15 mo thickened by only a small amount (<1 mm) compared to uncaged and open-caged (i.e. in cages lacking tops) sponges. Simulated bite marks on both reef and mangrove sponges were repaired at a rapid rate (0.8 to 1.6 mm d⁻¹). Fish predation has an important impact on the distribution and abundance of C. nucula, but the thin growth form common to reef environments may be more the result of hydrodynamics than of grazing by spongivorous fishes. Received: 6 October 1997 / Accepted: 19 March 1998     

A brominated secondary metabolite synthesized by the cyanobacterial symbiont of a marine sponge and accumulation of the crystalline metabolite in the sponge tissue

The dictyoceratid marine sponge Dysidea herbacea (Keller, 1889) is common in shallow waters of the tropical Pacific Ocean. Polybrominated biphenyl ethers such as 2-(2,4-dibromophenyl)-4,6-dibromophenol (1) are characteristic secondary metabolites of some specimens of this sponge and may represent as much as 12% of the dry weight. We have found 1 to be deposited as conspicuous crystals throughout the sponge tissue. The dominant prokaryotic endosymbiont in the mesohyl of the sponge is a filamentous cyanobacterium (Oscillatoria spongeliae), although a vacuole-containing, heterotrophic bacterium is also present. The cyanobacteria were separated from the sponge cells and heterotrophic bacteria by flow cytometry. Coupled gas chromatography—mass spectrometry and proton nuclear magnetic-resonance spectroscopy revealed that the major brominated Compound 1 isolated from the intact symbiotic association is found in the cyanobacteria and not in the sponge cells or heterotrophic bacteria. This suggests that the production of the compound is due to the cyanobacterium, and not to the sponge or symbiotic heterotrophic bacteria, as had been suggested earlier.     

Photoadaptation of zooxanthellae in the sponge Cliona vastifica from the Red Sea, as measured in situ

In the Red Sea, the zooxanthellate sponge Cliona vastifica (Hancock) is mainly present at >15 m depth or in shaded areas. To test whether its scarcity in unshaded areas of shallower waters is linked to the functional inefficiency of its photosymbionts at high irradiances, sponges were transferred from 30 m to a six times higher light regime at 12 m depth, and then returned to their original location. During this time, photosynthetic responses to irradiance were measured as rapid light curves (RLCs) in situ by pulse amplitude modulated (PAM) fluorometry using a portable underwater device, and samples were taken for microscopic determinations of zooxanthellar abundance. The zooxanthellae harboured by this sponge adapted to the higher irradiance at 12 m by increasing both their light saturation points and relative photosynthetic electron transport rates (ETRs). The ETRs at light saturation increased almost fourfold within 15–20 days of transfer to the shallower water, and decreased back to almost their original values after the sponges were returned to 30 m depth. This, as well as the fact that the photosynthetic light responses within an individual sponge were in accordance with the irradiance incident to specific surfaces, shows that these photosymbionts are highly adaptable to various irradiances. There was no significant change in the number of zooxanthellae per sponge area throughout these experiments, and the different photosynthetic responses were likely due to adaptations of the photosynthetic apparatus within each zooxanthella. In conclusion, it seems that parameters other than the hypothesised inability of the photosymbionts to adapt adequately to high light conditions are the cause of C. vastifica's rareness in unshaded shallow areas of the Red Sea. Received: 25 April 2000 / Accepted: 13 October 2000     

The fractal branching of an arborescent sponge

The fractal properties of specimens of a planar branching sponge Raspailia inaequalis (Porifera, Demospongiae) were determined by analysing digitised photographs. The specimens, collected from a single site in northeastern New Zealand, had a wide range of morphology. Three different fractal methods were used: box counting; a method that gives the scaling of branch length with distance from the base of the fan; and an allometric analysis of the dependence of frontal area on specimen size. All three approaches gave a similar value for the fractal dimension. The conjecture that the specimens have a fractal branching structure is consistent with the results of a Horton analysis of their branching pattern. There is a significant relationship between fractal dimension and number of fingers, which implies that a simple count of the number of fingers is as useful for discriminating between individuals as the more complex fractal analysis. Using this relation, sponges from a site with less water movement are inferred to have a lower fractal dimension. This result is in agreement with the predictions of the Kaandorp model of sponge growth. Received: 13 March 2000 / Accepted: 11 October 2000     

Deep-sea caging of the mussel Mytilus galloprovincialis: Potential application in ecotoxicological studies

We experimented with caging the Mediterranean mussel (Mytilus galloprovincialis) at various depths for 69 d to measure basic physiological parameters, histological response and bio-accumulation of contaminants in a deep-sea contaminated area. In preliminary experiments, we demonstrated, under artificial pressure conditions, the ability of mussels Mytilus galloprovincialis to tolerate rapid immersion (at a speed of up to 120 m min^?1). In situ experiments were performed using submerged lines enabling mussels to be maintained at depths ranging of 40–1550 m with survival rates ranging from 80 to 38%, respectively. No significant differences in condition indexes were observed between treated and control specimens. However, histological observations demonstrated a clear reduction in thickness of the digestive epithelium with increasing depth exposure. By determining the contaminants in caged mussels, we found the following values for chromium accumulation: 27.4 μg g^?1 dry weight at 580 m depth and 9.8 μg g^?1 dry weight at 1550 m. Selected stations were located downstream of an industrial effluent at 420 m. The biological and environmental consequences of deep-sea contamination demonstrate the suitability of caged mussels for monitoring contaminant accumulation.     

Ecological roles of natural products from the marine sponge Geodia corticostylifera

In the Brazilian coast, high numbers of the small brittle star Ophiactis savignyi usually live associated with the sponge Geodia corticostylifera (Demospongiae, Geodidae), but not with other sympatric sponge species. In order to check whether this association was related only with the physical shelter provided by the sponge body or was chemically mediated, the crude organic extract of G. corticostylifera was added to sponge mimics made of phytagel and spongin skeleton. Control and treated mimics were simultaneously offered to previously sponge-associated O. savignyi in both static seawater and flow-through laboratory experiments. Ophiuroids were allowed to move towards the preferred mimic. The defensive properties of the sponge extract against fish predation and fouling were also evaluated. Chemotaxis assays showed that symbiotic ophiuroids were able to chemically recognize its host sponge, moving significantly more towards mimics containing G. corticostylifera extract. Chemical deterrence assays showed that the natural concentration of the extract of this sponge was also able to inhibit generalist fish predation on field experiments and the attachment of the common mussel Perna perna in laboratory assays. These results indicate that the crude extract of G. corticostylifera plays multiple functions in the marine environment, presumably being responsible for a closer association of this sponge with O. savignyi, providing protection for this ophiuroid and inhibition of epibionts on itself.     

Evidence for morphology-induced sediment settlement prevention on the tubular sponge<Emphasis Type="Italic"> Haliclona urceolus</Emphasis>

Several mechanisms are known to assist the survival of sponges in highly sedimented environments. This study considers the potential of sponge morphology and the positioning of exhalant water jets (through the osculum) in the adaptation of Haliclona urceolus to highly sedimented habitats. This sponge is cylindrical with an apical osculum, which is common in sedimented subtidal habitats at Lough Hyne Marine Nature Reserve, Cork, Ireland. Fifteen sponges were collected, preserved (killed with the structure and morphology maintained) and then replaced in a high sediment environment next to a living specimen (at 24 m). After 5 days, the sediment settled on both living and preserved sponges was collected and dried. No sediment was collected from living sponges, while preserved specimens had considerable amounts of settled sediment on their surfaces. The amount of sediment collected on these preserved specimens was significantly linearly correlated with sponge dry weight, maximum diameter and oscula width (R²>0.70, P<0.001, df=14). Observations of flow direction (using coloured dye) through H. urceolus showed that water is drawn into the sponge on its underside and exits via a large vertically pointing osculum. Sponge morphologies (shape) have often been considered as a means of passive adaptation to a number of different environmental parameters with oscula position enabling entrained flow through the sponge in high flow conditions. However, this study shows how the combination of sponge morphology (tubular shape) and positioning of the osculum may enable H. urceolus to survive in highly sedimented environments. Similar mechanisms may also aid the survival of some deep-water sponge species with similar morphologies.Communicated by J.P. Thorpe, Port Erin     

Rapid tissue reduction and recovery in the sponge Aplysinella sp.

We observed a pronounced, yet reversible tissue reduction in the tropical sponge Aplysinella sp. under non-experimental conditions in its natural habitat, after transfer into seawater tanks, as well as after transplantation from deep to shallow water in the field. Tissue reduction resulted in the formation of small “reduction bodies” tightly attached to the sponge skeleton. Although volume loss and gain were substantial, both tissue reduction and regeneration were often remarkably rapid, occurring within few hours. Microscopic analysis of the reduction bodies revealed morphological similarities to previously described sponge primmorphs, with densely packed archaeocytes and spherulous cells enclosed by a thin layer of epithelial-like cells. Denaturing gradient gel electrophoresis (DGGE) revealed pronounced changes in the sponge-associated microbial community upon tissue reduction during laboratory and field experiments and following changes in ambient conditions after transplantation in the field. Generally, the microbial community associated with this sponge proved less stable, less abundant, and less diverse than those of other, previously investigated Verongid sponges. However, one single phylotype was consistently present in DGGE profiles of Aplysinella sp. This phylotype clustered with γ-proteobacterial sequences found previously in other sponge species of different taxonomic affiliations and geographic provenances, as well as in sponge larvae. No apparent changes in the total secondary metabolite content (per dry weight) occurred in Aplysinella sp. upon tissue reduction; however, comparative analysis of intact and reduced tissue suggested changes in the concentrations of two minor compounds. Besides being ecologically interesting, the tissue reduction phenomenon in Aplysinella sp. provides an experimentally manipulable system for studies on sponge/microbe symbioses. Moreover, it may prove useful as a model system to investigate molecular mechanisms of basic Metazoan traits in vivo, complementing the in vitro sponge primmorph system currently used in this context.     

Morphology of<Emphasis Type="Italic"> Ecklonia radiata</Emphasis> (Phaeophyta: Laminarales) along its geographic distribution in south-western Australia and Australasia

Ecklonia radiata (C. Ag.) J. Agardh is a common macroalga on reefs in the warm-temperate parts of the southern hemisphere. It is a dominant habitat-former and as such has a strong structuring effect on associated algal assemblages. Morphological variation in E. radiata potentially affects its interactions with the surroundings and contributes to confusion about its taxonomy. We quantified the magnitude of morphological variation in fully developed E. radiata sporophytes across Australasia and tested the hypotheses that E. radiata has different morphology at different locations and that the degree of morphological difference depends on spatial distances among locations. A total of 11 morphological characters were sampled from 11 locations along the Australian coastline from Kalbarri in Western Australia to Sydney in New South Wales as well as from Doubtful Sound, New Zealand. Most morphological characters varied considerably from one location to another. For example, the average (±SE) thallus length was 135.2±12.5 cm in Kalbarri and only 69.7±5.5 cm in Sydney. There were no consistent spatial patterns of variation among individual morphological characters, and, generally, variations among individual characters were poorly correlated (-0.5< R<0.5). This suggests that individual morphological characters develop independently of each other in response to processes operating at different spatial scales. Multivariate measures of morphology were found to be different among some locations and similar among others (-0.37=Clarke's  R=1), but there was no correlation (Spearman's R=0.08) between morphological similarity and distance between locations. Consequently, our results do not support clinal variation in E. radiata morphology. Rather, they suggest the presence of discrete morphologically different populations, in which the morphology at any one location reflects multiple forcing factors operating on different morphological characters at different spatial scales.     

Isolation of Oscillatoria spongeliae,the filamentous cyanobacterial symbiont of the marine sponge Dysidea herbacea

The tropical marine sponge Dysidea herbacea (Keller) (Dictyoceratidae: Dysideidae) is always found associated with the filamentous cyanobacterium (blue-green alga) Oscillatoria spongeliae (Schulze) Hauck (Cyanophyceae: Oscillatoriaceae), which occurs abundantly throughout the sponge mesohyl. Intact, metabolically active, trichomes of O. spongeliae were isolated from the sponge by chopping the sponge tissue with a razor blade and squeezing the trichomes into a seawater-based medium containing polyvinylpyrrolidone, bovine serum albumin, dithiothreitol, glycerol, KCl and Na₂CO₃. The isolated cyanobacteria were concentrated by centrifugation and then washed several times in fresh medium. The isolated O. spongeliae have photosynthetic rates which are similar to the intact sponge-alga association for periods of at least 6 h after isolation. Addition of sponge homogenate to the isolated cyanobacteria causes rapid cell lysis.     

Oxygen dynamics and transport in the Mediterranean sponge Aplysina aerophoba

The Mediterranean sponge Aplysina aerophoba kept in aquaria or cultivation tanks can stop pumping for several hours or even days. To investigate changes in the chemical microenvironments, we measured oxygen profiles over the surface and into the tissue of pumping and non-pumping A. aerophoba specimens with Clark-type oxygen microelectrodes (tip diameters 18–30 μm). Total oxygen consumption rates of whole sponges were measured in closed chambers. These rates were used to back-calculate the oxygen distribution in a finite-element model. Combining direct measurements with calculations of diffusive flux and modeling revealed that the tissue of non-pumping sponges turns anoxic within 15 min, with the exception of a 1 mm surface layer where oxygen intrudes due to molecular diffusion over the sponge surface. Molecular diffusion is the only transport mechanism for oxygen into non-pumping sponges, which allows total oxygen consumption rates of 6–12 μmol cm⁻³ sponge day⁻¹. Sponges of different sizes had similar diffusional uptake rates, which is explained by their similar surface/volume ratios. In pumping sponges, oxygen consumption rates were between 22 and 37 μmol cm⁻³ sponge day⁻¹, and the entire tissue was oxygenated. Combining different approaches of direct oxygen measurement in living sponges with a dynamic model, we can show that tissue anoxia is a direct function of the pumping behavior. The sponge-microbe system of A. aerophoba thus has the possibility to switch actively between aerobic and anaerobic metabolism by stopping the water flow for more than 15 min. These periods of anoxia will greatly influence physiological variety and activity of the sponge microbes. Detailed knowledge about the varying chemical microenvironments in sponges will help to develop protocols to cultivate sponge-associated microbial lineages and improve our understanding of the sponge-microbe-system.     

Redwood of the reef: growth and age of the giant barrel sponge Xestospongia muta in the Florida Keys

The growth of animals in most taxa has long been well described, but the phylum Porifera has remained a notable exception. The giant barrel sponge Xestospongia muta dominates Caribbean coral reef communities, where it is an important spatial competitor, increases habitat complexity, and filters seawater. It has been called the ‘redwood of the reef’ because of its size (often >1 m height and diameter) and presumed long life, but very little is known about its demography. Since 1997, we have established and monitored 12 permanent 16 m diameter circular transects on the reef slope off Key Largo, Florida, to study this important species. Over a 4.5-year interval, we measured the volume of 104 tagged sponges using digital images to determine growth rates of X. muta. Five models were fit to the cubed root of initial and final volume estimates to determine which best described growth. Additional measurements of 33 sponges were taken over 6-month intervals to examine the relationship between the spongocoel, or inner-osculum space, and sponge size, and to examine short-term growth dynamics. Sponge volumes ranged from 24.05 to 80,281.67 cm³. Growth was variable, and specific growth rates decreased with increasing sponge size. The mean specific growth rate was 0.52 ± 0.65 year⁻¹, but sponges grew as fast or slow as 404 or 2% year⁻¹. Negative growth rates occurred over short temporal scales and growth varied seasonally, significantly faster during the summer. No differences in specific growth rate were found between transects at three different depths (15, 20, 30 m) or at two different reef sites. Spongocoel volume was positively allometric with increasing sponge size and scaling between the vertical and horizontal dimensions of the sponge indicated that morphology changes from a frustum of a cone to cylindrical as volume increases. Growth of X. muta was best described by the general von Bertalanffy and Tanaka growth curves. The largest sponge within our transects (1.23 × 0.98 m height × diameter) was estimated to be 127 years old. Although age extrapolations for very large sponges are subject to more error, the largest sponges on Caribbean reefs may be in excess of 2,300 years, placing X. muta among the longest-lived animals on earth.     

Symbiotic zooxanthellae enhance boring and growth rates of the tropical sponge Anthosigmella varians forma varians

Several species of boring sponges harbor symbiotic zooxanthellae, and it is believed that the symbiont enhances boring activity of host sponges. This hypothesis was tested using manipulative field experiments to assess the effect of intracellular zooxanthella populations on boring rates of the tropical sponge Anthosigmella varians forma varians. Portions of sponge were attached to 60 calcium carbonate blocks of known weight. Three sets of 10 blocks were grown at high light levels and three sets of 10 blocks were grown at low light levels for 105 d in the Florida Keys, Florida, USA. Boring rates, growth rates (lateral growth and within-substratum tissue penetration), and zooxanthella populations were measured at the end of the experiment. Absolute rates of boring and growth of A. varians forma varians were significantly greater when zooxanthella densities were higher. Boring rate and tissue penetration related to final surface area of sponge attachment was also enhanced when zooxanthella densities were higher, suggesting that the symbiont plays a physiological role in the decalcification process. This is in contrast to the role that zooxanthellae play in coral hosts. Based on the results of this study, it appears that the presence of zooxanthellar symbionts has important ecological and life-history consequences for host sponges. Ability to laterally overgrow competitors will be correlated with the size and activity of zooxanthella populations. In addition, the fitness of host sponges will be enhanced by algal symbionts, since greater penetration within substrata will result in an increase in production of tissue that can be converted into storage, feeding and reproductive functions.     

Ultrastructural aspects of the symbiosis between two species of the genus Aphanocapsa (Cyanophyceae) and Ircinia variabilis (Demospongiae)

The association between two species of the genus Aphanocapsa (Cyanophyceae) and the sponge Ircinia variabilis has been studied by electron microscopy. A. feldmanni is localized in the mesohyl or inside the cells of the sponge, while the larger A. raspaigellae is located only in an extracellular position inside cavities of the mesohyl. Both algae differ from other symbiotic Cyanophyceae in having a normal cell wall. They are able to reproduce in symbiotic condition, but also undergo, in their various extracellular and intracellular positions, a massive process of disintegration. A large amount of algal material is dispersed in the sponge tissues, which is a confirmation, at the ultrastructural level, of trophic relationships in the symbiosis Aphanocapsa-Ircinia.     

Changes in morphology and physiology of an East Mediterranean sponge in different habitats

The sponge Tetilla sp. (Tetractinomorpha: Tetillidae) is a common species in the eastern Mediterranean. This sponge inhabits four different habitat types differing in wave impact and irradiance levels. Two of these habitats (a shallow cave and deep water) are characterized by relatively calm water, whereas the other two (shallow exposed site and tide pools) are in turbulent water with high energy flow. The present study examined the influence of physical (depth, illumination and water motion) and biotic factors on morphology, skeletal plasticity and reproductive traits among the four spatially separated populations. Sponges from tidal pools had significantly larger body volume than sponges from deep water and from shallow caves (ANOVA: tidal-deep P<0.0001; tidal-shallow caves P<0.05). Sponges from exposed habitats were significantly larger than deep-water sponges (ANOVA: P=0.01). In addition, individuals from tide pools and from the exposed habitat had a significantly higher proportion of structural silica than sponges from the calmer deep water and from the cave sites. Oxea spicules in sponges from the calm habitats were significantly shorter than in those from the tidal pools and the exposed habitats. The percentage of spicules out of a sponges dry weight in individuals transplanted from deep (calm) to shallow (turbulent) water significantly increased by 21.9±12.9%. The new spicule percentage did not differ significantly from that of sponges originally from shallow water. Oocyte diameter differed significantly between habitats. The maximal size of mature eggs was found in deep-water sponges in June (97±5 m). In the shallow habitats, a smaller maximal oocyte diameter was found in the cave, in May (56.5±3 m). Furthermore, oocyte density in shallow-water sponges was highest in May and decreased in June (with 88.2±9 and 19.3±9 oocytes mm^–2, respectively). At the same time (June), oocyte density of deep-water sponges had just reached its maximum (155±33.7 oocytes mm^–2). The difference in oocyte size and density between deep- and shallow-water individuals indicates an earlier gamete release in the shallow sponge population. The results suggest that plasticity in skeletal design of this sponge indicates a trade off between spicule production and investment in reproduction.Communicated by O. Kinne, Oldendorf/Luhe     

Genetic evidence for cryptic speciation in allopatric populations of two cosmopolitan species of the calcareous sponge genusClathrina

Many sponge species are considered to be cosmopolitan. However, the systematics of marine sponges are very difficult because of the paucity of taxonomically useful characters, and hence the apparently cosmopolitan nature of many species may be simply a consequence of this. In this paper, geographically distant populations of two pairs of cosmopolitan calcareous sponges of the genusClathrina were compared genetically.C. clathrus andC. cerebrum were collected by SCUBA diving between January and March 1989 from two localities: the Mediterranean Sea at La Vesse, near Marseille, Frances, at 9 to 12 m depth, and from the South West Atlantic at Arraial do Cabo, about 200 km east of Rio de Janeiro, at 2 to 10 m depth. Very high levels of gene divergence were found between the allegedly conspecific populations. The levels of genetic identity,I, observed are so low (I=0.128 and 0.287) that the populations clearly cannot be considered conspecific. New species names ofC. aurea sp. nov. andC. brasiliensis sp. nov. are therefore assigned to the southwest Atlantic counterparts ofC. clathrus andC. cerebrum, respectively. It is concluded that, at least for the species studied, and probably for many other species in taxonomically difficult groups, the actual distributions of single species may be far more geographically restricted than is generally assumed.     

Verification of fractal growth models of the sponge Haliclona oculata (Porifera) with transplantation experiments

The growth form of the sponge Haliclona oculata is to a significant extent determined by the environmental conditions in which the form emerges. One of the main environmental parameters affecting the growth form is exposure to water movement. In this study, a morphological growth model is used to simulate the effect on the growth process of a change in exposure to water movement. Predictions based on the model are verified by experiments in which sponges from a sheltered growth site are transplanted to an exposed site, and vice versa. The effect of the transplantation on growth forms is determined by morphological comparisons. By combining the morphological simulation model with interpretation of growth forms, it becomes possible to use the growth form of H. oculata for bio-monitoring purposes. This form reflects the environmental conditions governing the growth process.     

A comparison of two cartographic exposure methods using Fucus vesiculosus as an indicator

Morphological variation and vertical distribution of Fucus vesiculosus were quantified at several sites in the Finnish archipelago (Baltic Sea). F. vesiculosus samples were obtained from skerries at geographical distances of 1 km or more (large scale) and at intervals of ca 100 m around a single island (small scale). The results were examined in relation to wave exposure, calculated by Baardseth and effective fetch cartographic methods. Despite the fact that the exposure indices were calculated differently they correlated strongly. Vegetative morphological characteristics of F. vesiculosus illustrate the morphological differences both within and between exposure gradients. The tallest and widest F. vesiculosus plants were found at the sheltered end of the large-scale exposure gradient. Those from equally sheltered sites of the island were smaller in all respects. Thus, the trend from small narrow plants to large wide sheltered plants was expressed differently over the different geographical scales. Consequently localities with similar exposure indices may have morphologically different F. vesiculosus populations. Shores with similar cartographic exposure indices can be different in nature. Underwater topography and shore locations, either close to the mainland or at the outermost sites of the archipelago, affect the exposure. Although a sheltered shore is indicated, the sublittoral zone may be quite exposed to the movements of water. In contrast, in an open shore environment underwater rocks, boulders and shallow water areas can provide sheltered habitats. The depth range of the F. vesiculosus belt exhibited two distinctive patterns. At sheltered sites, around islands in the outermost reaches of the archipelago F. vesiculosus can grow to a maximum depth of 5 m. In exposed habitats the belt becomes narrower, reaching a maximum depth of 3 m. Closer to the mainland F. vesiculosus is found at exposed sites to a maximum depth of 5 m; the depth range at sheltered sites is narrower, only reaching depths of 2 m or less. In conclusion, the changes in plant morphology and in the vertical belt distribution are similar to each other along both gradients at the exposed ends of the wave action spectrum; however, the two gradients diverge at the sheltered ends of the spectrum. Received: 10 August 1998 / Accepted: 11 January 1999     

Transmission,plasticity and the molecular identification of cyanobacterial symbionts in the Red Sea sponge <Emphasis Type="Italic">Diacarnus erythraenus</Emphasis>

Cyanobacterial symbionts in the sponge Diacarnus erythraenus from the Red Sea were identified in both adult sponges and their larvae by 16S rDNA sequencing. A single cyanobacterial type was found in all samples. This cyanobacterial type is closely related to other sponge cyanobacterial symbionts. The cyanobacterial rDNA, together with the morphological analysis by electron and fluorescence microscopy, provided evidence for vertical transmission of the symbionts in this sponge. In addition, we show phenotypic plasticity of the symbionts inside the sponge, probably as a result of variability in light availability inside the sponge tissue. Finally, the reproduction of Diacarnus erythraenus is also described.Matan Oren, Laura Steindler have contributed equally to the work.     

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.