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     

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.     

Bioerosion of coral reefs by two Hawaiian parrotfishes: species, size differences and fishery implications

Parrotfishes can be significant bioeroders and sediment producers on coral reefs. We quantified the bioerosion rates of two similarly sized Hawaiian parrotfishes with two different feeding modes (Scarus rubroviolaceus—a scraper and Chlorurus perspicillatus—an excavator). The results showed that feeding modes did not affect bioerosion rates but that bioerosion rates were size dependent, with largest individuals (S. rubroviolaceus 45–54 cm FL) bioeroding up to 380 ± 67 kg ind⁻¹ year⁻¹. The size for onset of bioerosion capabilities for both species was 15 cm. Grazing by the two species consumed 60% of the carbonate production of the fore reef area, suggesting that large parrotfishes in Hawaii are ecologically important bioeroders. As individual large S. rubroviolaceus contributed disproportionately more to bioerosion and sediment production than the equivalent biomass of smaller conspecifics, management strategies designed to retain normal reef bioerosion rates should seek to preserve the historical size structure of S. rubroviolaceus populations and to especially protect the larger size classes.     

Microbial associations in sponges. I. Ecology,physiology and microbial populations of coral reef sponges

Illumination, current strength and physical turbulence influence the distribution of 4 tropical sponges. Three sponges with cyanobacteria in exposed tissues grow only in poen shallow habitats: Pericharax heteroraphis in moderate-current, lowturbulence regions on the reef slope; Jaspis stellifera in low-current, moderate-turbulence regions of the outer reef flat; and Neofibularia irata in moderate-current, high-turbulence areas below the reef crest. Ircinia wistarii contains no cyanobacteria and occurs in deeper, strong-current, high-turbulence regions. N. irata agressively overgrows neighbouring corals and its growth form is influenced by the current strength. The sponges efficiently filter bacteria from the water. The efficiency is related to the aquiferous structure, particularly the size of choanocyte chambers, and is unrelated to the existing bacterial populations in sponge tissue. The numbers of bacteria associated with the sponges are proportional to the sponge mesohyl density, with the dense sponges J. stellifera and I. wistarii containing many bacteria whereas P. heteroraphis is not dense and has few bacteria.     

Trends in space occupation by the encrusting sponge Crambe crambe: variation in shape as a function of size and environment

The relationship between sponge size, habitat and shape was studied in the encrusting sponge Crambe crambe (Schmidt, 1862), which is distributed widely throughout the shallow Mediterranean littoral. Examination of sponge patches in shaded and well-illuminated habitats showed that the degree of peripheral irregularity of the edges of a patch is directly related to patch size. This relationship is valid only for sponges of >100 mm² in area. Photophilic and sciaphilous sponges display different growth forms. The pattern of growth is interpreted in terms of competition for space. The directional growth of sciaphilous sponges may be due to the presence of dominant neighbours that are good space competitors, and the irregular growth of photophilic sponges to the absence of such neighbours.     

Variability in reproductive output across a water quality gradient for a tropical marine sponge

To establish a complete understanding of reproductive variability, larval supply and ultimately population demographics of a species it is important to determine reproduction across a broad spectrum of environmental conditions. This study quantified sexual reproduction of the brooding, gonochoristic sponge Rhopaloeides odorabile from populations across the shelf reefs of the central Great Barrier Reef (GBR). Histological sections of reproductive sponges collected at increasing distances from the coast were used to determine if numbers of reproductive sponges, reproductive output (using a reproductive output index), size at sexual maturity, and sex ratios varied according to their location (distance) from the coastline and therefore from influences of terrigenous/riverine discharge. Significantly higher proportions of reproductive sponges occurred with increasing distance from the coast. The proportion of all reproductive sponges (both male and female) on offshore reefs ranged from 77 to 90%, during November and December, the peak reproductive months of this sponge, compared to 47 to 50% for sponges occurring on coastal reefs. Levels of female reproduction increased with increasing distance from the coastline on two levels. First, oocytes from offshore sponges were significantly larger than oocytes from coastal sponges. Second, sponges from offshore reefs showed a reproductive index (proportions of oocytes, embryos and larvae mm⁻²) approximately 15 times higher than coastal reef sponges. Therefore, both numbers of oocytes, embryos and larvae in conjunction with larger oocytes contribute to a higher reproductive output index for offshore sponges. The production of spermatic cysts in males was consistent across the GBR. Sex ratios for coastal reef sponges showed a male bias while offshore sponges showed approximate equal sex ratios. The effect of terrigenous riverine input from coastal fluvial plains to the inner GBR is well established and is likely to contribute to the lower levels of reproduction associated with female sponges inhabiting coastal reefs of the central GBR.     

Life history and production of the mysid <Emphasis Type="Italic">Orientomysis robusta</Emphasis>: high <Emphasis Type="Italic">P/B</Emphasis> ratio in a shallow warm-temperate habitat of the Sea of Japan

Although mysids play important roles in marine food chains, studies on their production are scarce, especially for warm-water species. We investigated life history and production of Orientomysis robusta in a shallow warm-temperate habitat of the Sea of Japan. Its spawning and recruitment occurred throughout the year; 19 overlapping cohorts were recognizable over an annual cycle. The summer cohorts recruited in July–September exhibited rapid growth, early maturity, small brood size, and small body size. A converse set of life history traits characterized the autumn–winter cohorts recruited in October–March. The spring cohorts recruited in April–June had intermediate characteristics of both cohorts. Life spans were 19–33, 21–48, and 69–138 days for summer, spring, and autumn–winter cohorts, respectively, and mortality rates were high for spring and summer cohorts, especially during June–August but were low for autumn–winter cohorts. Production calculated from the summation of growth increments was 488.8 mg DW m⁻² year⁻¹ with an annual P/B ratio of 21.26. The short life span seems to be responsible for such an extremely high P/B ratio. A method not requiring recognition and tracking cohorts gave similar values (534.0 mg DW m⁻² year⁻¹ and 20.49). The close agreement in production values between the two methods indicates our estimates are valid.     

Sponge-feeding by the Caribbean starfish Oreaster reticulatus

The common Caribbean starfish Oreaster reticulatus (Linnaeus) feeds on sponges by everting its stomach onto a sponge and digesting the tissue, leaving behind the sponge skeleton. In the San Blas Islands, Republic of Panama, 54.2% of the 1549 starfish examined from February 1987 to June 1990 at eight sites were feeding, and 61.4% of these were feeding on sponges, representing 51 species. Sponges were fed on disproportionately heavily in comparison to their abundance, which was only 9.7% of available prey. In feeding choice experiments, 736 pieces of 34 species of common sponges from a variety of shallow-water habitats, and also 9 ind of a coral, were offered to starfish in individual underwater cages. Acceptance or rejection of sponge species was unambiguous for 31 of the 34 species, and there was a clear relationship between sponge acceptability and sponge habitat. Starfish ate 16 of 20 species that normally grow only on the reefs, but only 1 of 14 species that live in the seagrass meadows and rubble flats surrounding the reefs. The starfish live in the seagrass meadows and rubble flats, and avoid the reefs, and so the acceptable reef sponges are generally inaccessible until a storm fragments and transports them into starfish habitat. After Huricane Joan washed fragments of reef sponges into a seagrass meadow in October 1988, starfish consumed the edible species. When the seagrass meadow was experimentally seeded with tagged reef sponge fragments in June 1994, O. reticulatus consumed edible species and accumulated in the area seeded. Reef sponges that were living in a seagrass meadow, from which O. reticulatus had been absent for at least 4 yr (from 1978 to 1982), were eliminated when the starfish migrated into the area, and the sponges have been unable to recolonize up to June 1994. O. reticulatus feeding and habitat preferences appear to restrict distributions of many Caribbean reef sponge species to habitats without O. reticulatus and may have exerted significant selective pressure on defences of those sponges that live in O. reticulatus habitats.     

Carbon conversion and metabolic rate in two marine sponges

The carbon metabolism of two marine sponges, Haliclona oculata and Dysidea avara, has been studied using a ¹³C isotope pulse-chase approach. The sponges were fed ¹³C-labeled diatoms (Skeletonema costatum) for 8 h and they took up between 75 and 85%. At different times, sponges were sampled for total ¹³C enrichment, and fatty acid (FA) composition and ¹³C enrichment. Algal biomarkers present in the sponges were highly labeled after feeding but their labeling levels decreased until none was left 10 days after enrichment. The sponge-specific FAs incorporated ¹³C label already during the first day and the amount of ¹³C label inside these FAs kept increasing until 3 weeks after labeling. The algal-derived carbon captured by the sponges during the 8-h feeding period was thus partly respired and partly metabolized during the weeks following. Apparently, sponges are able to capture enough food during short periods to sustain longer-term metabolism. The change of carbon metabolic rate of fatty acid synthesis due to mechanical damage of sponge tissue was studied by feeding sponges with ¹³C isotope–labeled diatom (Pheaodactylum tricornutum) either after or before damaging and tracing back the ¹³C content in the damaged and healthy tissue. The filtration and respiration in both sponges responded quickly to damage. The rate of respiration in H. oculata reduced immediately after damage, but returned to its initial level after 6 h. The ¹³C data revealed that H. oculata has a higher metabolic rate in the tips where growth occurs compared to the rest of the tissue and that the metabolic rate is increased after damage of the tissue. For D. avara, no differences were found between damaged and non-damaged tissue. However, the filtration rate decreased directly after damage.     

Sexual reproduction and settlement of the coral reef spongeChalinula sp. from the Red Sea

Characteristics of the sexual reproduction and larval settlement of the haplosclerid spongeChalinula sp., which inhabits the shallow waters (1 to 6 m) of Eilat, Red Sea, were investigated from September 1985 through to November 1987. This species was found to be a simultaneous hermaphroditic brooder, hence gonochorism is not the rule in the order Haplosclerida. Brooding always takes place in special brooding chambers. While the oocytes in the brooding chambers are among the largest known in sponges (355±37µm), the spermatic cysts distributed in the choanosome are among the smallest known for this phylum (average 26±7µm).Chalinula sp. breeds throughout the year and in experiments most larvae (74%) settled within 1 to 8 h post-release, generally within 4.5 h. Metamorphosis from larval shape to a sessile sponge lasts 1 to 6 h. Thus, larvae had a short swimming period, settled fast, and metamorphosed rapidly (within 1 to 6 h). The large size of the larvae may contribute to their ability to rapidly reorganize their body shape into that of a sessile sponge. In addition, the existence of already differentiated choanocyte chambers in the larvae, facilitates fast construction of the water filtration system in the newly settled sponges. The reproductive and larval characteristics ofChalinula sp. enable the larvae to settle on any vacant space in the reef, which may explain its abundance in the Red Sea.     

Linking abundance and diversity of sponge-associated microbial communities to metabolic differences in host sponges

Many sponge species contain large and diverse communities of microorganisms. Some of these microbes are suggested to be in a mutualistic interaction with their host sponges, but there is little evidence to support these hypotheses. Stable nitrogen isotope ratios of sponges in the Key Largo, Florida (USA) area grouped sponges into species with relatively low δ¹⁵N ratios and species with relatively high δ¹⁵N ratios. Using samples collected in June 2002 from Three Sisters Reef and Conch Reef in the Key Largo, Florida area, transmission electron microscopy (TEM) and denaturing gradient gel electrophoresis were performed on tissues of the sponges Ircinia felix and Aplysina cauliformis, which are in the low δ¹⁵N group, and on tissue of the sponge Niphates erecta, which is in the high δ¹⁵N group. Results showed that I. felix and A. cauliformis have large and diverse microbial communities, while N. erecta has a low biomass of one bacterial strain. As the low δ¹⁵N ratios indicated a microbial input of nitrogen, these results suggested that I. felix and A. cauliformis were receiving nitrogen from their associated microbial community, while N. erecta was obtaining nitrogen solely from external sources. Sequence analysis of the microbial communities showed a diversity of metabolic capabilities among the microbes of the low δ¹⁵N group, which are lacking in the high δ¹⁵N group, further supporting metabolic differences between the two groups. This research provides support for hypotheses of mutualisms between sponges and their associated microbial communities.     

Silica uptake kinetics of Halichondria panicea in Kiel Bight

Besides diatoms Demospongiae are the most important consumers of dissolved silica in the sea. They can play an important role for the silica budget especially in the shallow water areas of the Baltic Sea. The dependence of the silica uptake rate on the silica concentration of the seawater was measured for the sponge Halichondria panicea (Pallas, 1766). The sponges were collected in Kiel Bight. The uptake conformed to Michaelis–Menten kinetics with a half-saturation constant of 46.41 μM and a saturated uptake rate of 19.33 μmol h⁻¹ g⁻¹ ( p < 0.01). In the red algae zone of Kiel Bight the sponges depend on silica supply from the surrounding waters and may be silica-limited rather than food-limited in growth. Because of the much faster uptake of silica by diatoms and their lower saturation point, as well as the difference in spatial distribution of the two main silica consumers, a competition for silica between sponges and diatoms seems unlikely. Received: 21 June 1997 / Accepted: 15 July 1997     

Parrotfish predation on cryptic sponges of Caribbean coral reefs

Some sponge species that live in crevices in the reef frame appear to be restricted to their cryptic habitat by predation. When cryptic sponges were excavated, on Guigalotupo reef, San Blas, Panama, exposing them to potential predators, they were eaten by fishes that are generally considered to be herbivores, primarily parrotfishes of the genus Sparisoma: S.aurofrenatum (Cuvier & Valenciennes), S.viride (Bonnaterre), and S.chrysopterum (Bloch & Schneider). Of the 9150 bites observed to be taken by these species during paired (i.e., with sponges versus without sponges) trials conducted in defined feeding areas during 1986, 1987, and 1988, 72% (i.e. 6581 bites) were on cryptic sponges, even though these were only offered during half of the total observation time and never constituted more than 7% of the cover of the feeding observation areas. Individual parrotfish returned over and over to take bites of the exposed cryptic sponges until they were entirely consumed. They vigorously chased each other away from the sponges, but exhibited no such defense of their usual algal foods. A total of 18 sponge species were tested. Of the cryptic and semi-cryptic sponge species tested, only one of six was rejected by the parrotfish. Two of these six sponge species were consistently consumed entirely, and two were consumed entirely whenever their surfaces were sliced off with a razor blade, demonstrating that these sponges concentrate defenses against predators in their surfaces. One semi-cryptic species and one semi-exposed species were fed upon, but not entirely consumed. By contrast, 11 of 12 of the exposed and semi-exposed species were rejected. Cryptic sponges grew out of their cavities in the reef only when protected by seasonally thick mats of macroalgae or by cages that excluded fish. Received: 10 January 1997 / Accepted: 4 February 1997     

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     

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     

Biodiversity and rockfish recruitment in sponge gardens and bioherms of southern British Columbia,Canada

In the Strait of Georgia and Howe Sound, British Columbia, colonies of individual cloud sponges, growing on rock (known as sponge gardens) receive resource subsidies from the high biodiversity of epifauna on adjacent rock habitats. Bioherms are reefs of glass sponges living on layers of dead sponges. In the same area as the sponge gardens, newly discovered bioherms in Howe Sound, BC (49.34.67 N, 123.16.26 W) at depths of 28- to 35-m are constructed exclusively by Aphrocallistes vastus, the cloud sponge. The sponge gardens had much higher taxon richness than the bioherms. The sponge garden had 106 species from 10 phyla, whereas the bioherm had only 15 species from 5 phyla. For recruiting juvenile rockfish (quillback, Sebastes maliger), the food subsidy of sponge gardens appears to be missing on bioherms of cloud sponge, where biodiversity is relatively low. While adult and subadult rockfishes (S. maliger, S. ruberrimus, S. proriger, and S. elongatus) were present on bioherms, no evidence for nursery recruitment of inshore rockfishes to bioherms was observed, whereas the sponge gardens supported high densities of newly recruited S. maliger, perhaps owing to the combination of both refuge and feeding opportunities. These results indicate that sponge gardens form a habitat for early stages of inshore S. maliger, whereas A. vastus bioherms are associated only with older juvenile and adult rockfishes.     

Life history characteristics of a coral reef sponge

Study of the life history characteristics of the common Red Sea sponge Mycale fistulifera (Poecilosclerida: Demospongiae) reveals several traits which may categorize it as an opportunistic species: (1) at least part of the population is reproductively active throughout the year, providing a continuous supply of larvae for settlement; (2) sponges may produce and release a large number (152±39 larvae d^-1) of brooded larvae; (3) released larvae are capable of fast settlement and metamorphosis (minutes to 30 hours post-release); (4) the turnover of the population is high and population size varies with time. From 48 initial sponges plus 94 recruits, only 25 remained alive after 14 months of observation; (5) most members of the population (>70%) have a small body size (<30 cm²); (6) sexual maturity may occur at an early age. These traits facilitate continuous establishment of M. fistulifera in new spaces on the reef; (7) M. fistulifera, preferred substrate is another opportunistic species, the coral Stylophora pistillata which it overgrows.     

Sponge-inhabiting barnacles on Red Sea coral reefs

In this study eight different species of barnacles were found within nine species of sponges from the Red Sea. This brings to 11 the number of sponge-symbiotic barnacles reported from the Red Sea, two of these are new Acasta species (not described herein) and one (A. tzetlini Kolbasov) is a new record for this sea. This number is much higher than that of symbiotic barnacles found within sponges from either the N. Atlantic (2) or the Mediterranean (4). Two possible explanations for this are the presence of numerous predators in coral reefs and scarcity of available substrate for settlement. These factors can lead to high incidence of symbiotic relationships. Of the nine sponge species, only one (Suberites cf. clavatus) had previously been known to contain barnacles. Even at the family level, this is the first record of symbiotic barnacles in two out of the seven sponge families (Latrunculiidae, Theonellidae). Our present findings strengthen the apparent rule that the wider the openings in a barnacle shell, the fewer the host taxa with which it will associate, usually from one or two closely related families, and the more frequent it will associate with elastic sponges. Most Neoacasta laevigata found on Carteriospongia foliascens were located on the same side as the sponge's ostia, i.e. facing the incoming water. This adaptation allows the barnacles to catch more suspended particles from the water, provides them with more oxygen and prevents their exposure to discharged sponge waste. The highest density of barnacles observed on one face of a “leaf ” (with ostia) was 0.389 barnacles cm⁻² (one barnacle per 2.57 cm²) and on average 0.181 ± 0.68, while the average on the other side was only 0.068 ± 0.52 barnacles cm⁻². As indicated by the Morisita index, these barnacles most frequently (58%, n = 12) had a clumped spatial distribution (while the rest were randomly distributed), as is to be expected from such sessile organisms with internal fertilization via copulation. The presence of N. laevigata induced the growth of secondary perpendicular projections of its host C. foliascens. Of the N. laevigata examined, 17% brooded 324 ± 41 embryos each, of 286 ± 17 μm total length; only 5.7% (n = 123) were found to be dead. Size distribution analysis of skeletal elements from dead barnacles showed them to be significantly larger than the skeletal elements of the population of live barnacles ( p < 0.05). Received: 26 June 1998 / Accepted: 1 December 1998     

Photosynthetic performance of giant clams, Tridacna maxima and T. squamosa, Red Sea

Two species of giant clams, Tridacna maxima and T. squamosa, coexist in the Red Sea, but exhibit distinctly different depth distributions: T. maxima mostly occurs in shallow waters (reef flat and edge), while T. squamosa may occur down to the lower fore-reef slope. Giant clams have been described as mixotrophic, capable of both filter-feeding and photosynthesis due to algal symbionts (zooxanthellae), therefore, observed depth preferences were investigated in relation to possible differences in autotrophy vs. heterotrophy. This study was conducted from April to June 2004, at the reef near the Marine Science Station, Aqaba, Gulf of Aqaba, Red Sea, and in May 2007, at a reef near Dahab, Sinai Peninsula, Egypt. In situ measurements using a submersible pulse amplitude modulated fluorometer (Diving PAM), revealed no significant differences in effective PSII quantum yield (ΔF/Fm′) and relative electron transport rates (ETR) between the two species; but rapid light curves (ETR vs. light, photosynthetically active irradiance, PAR) showed significant differences in maximum photosynthetic rates (ETR_max), with 20% higher values in T. maxima. Chamber incubations displayed higher net and gross oxygen production by T. maxima (88.0 and 120.3 μmol O₂ cm⁻² mantle area day⁻¹) than T. squamosa (56.7 and 84.8 μmol O₂ cm⁻² mantle area day⁻¹); even under shading conditions (simulated depth of 20 m) T. maxima still achieved 93% of the surface gross O₂ production, whereas T. squamosa reached only 44%. A correlation was found between ETR and net photosynthesis measured as oxygen production (T. maxima: R ² = 0.53; T. squamosa: R ² = 0.61). Calculated compensation depth (CD) (gross photosynthesis equals respiration) in T. maxima (16 m) matches the maximum depth of occurrence in this study (17 m). By contrast, the CD of T. squamosa (9 m) was much shallower than the maximum vertical range (42 m). Findings suggest T. maxima is a strict functional photoautotroph limited by light, whereas T. squamosa is a mixotroph whose photoautotrophic range is extended by heterotrophy. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Among-habitat variation in herbivory on Sargassum spp. on a mid-shelf reef in the northern Great Barrier Reef

Herbivory is widely acknowledged as a key process determining the benthic community structure and resilience of coral reefs. Despite numerous studies that have examined herbivory across reef gradients in the Caribbean, few studies have directly quantified this process on Pacific reefs. Bioassays of two species of erect macroalgae (Sargassum swartzii and S. cristaefolium) were used to quantify variation in grazing intensity across seven habitats of varying depth and wave exposure on a mid-shelf reef in the northern Great Barrier Reef. Removal rates of Sargassum varied significantly among habitats, with both species displaying broadly similar patterns. The shallow habitats on the exposed aspect of the reef (i.e. reef crest, flat and back reef) experienced the highest reductions in mass (81.4–91.6% day⁻¹) for both S. swartzii and S. cristaefolium, while the deeper exposed habitats (reef slope and base) displayed the lowest reductions (3.8–13.4% day⁻¹) over a 24 h period. In contrast, the grazing intensity varied between the two species in the three habitats on the leeward aspect of the reef. Reductions in mass remained relatively high for S. swartzii on the patch reef and sheltered reef base and flat (62.7–76.5% day⁻¹) but were considerably lower for S. cristaefolium (37.9–63.5% day⁻¹) across the same habitats. Surprisingly, the rates of removal of Sargassum displayed no relationship with the density or biomass of roving herbivorous fishes or those species known to consume erect macroalgae, either collectively or independently. These results suggest that the relationship between browsing rates and herbivorous fish biomass is complex and may be driven by species that are underestimated in visual surveys. Direct quantification of browsing intensity using assays revealed a different pattern to inferences based on herbivore densities and highlights the potential difficulties of evaluating ecosystem processes based on visual census data alone.