Photosensitivity of the calanoid copepod Acartia tonsa

The light intensity and spectral sensitivities of the calanoid copepod Acartia tonsa Dana were determined by measuring phototactic responses. Adult females displayed only positive phototaxis. The dark-adapted copepod, which possesses a single naupliar eye, perceived light at intensities as low as 2.8x10¹¹ photons m^-2 s^-1. The action spectrum for positive phototaxis had no clear maxima but rather showed a broad range of greatest sensitivity from 453 to 620 nm. This sensitivity encompassed those wavelengths that are maximally available at the depth where the copepod is found during the day. This spectral overlap, coupled with the finding that the copepod requires light cues for nocturnal vertical miration, suggests that broad spectral sensitivity is an adaptive mechanism to maximize light intensity sensitivity during migration.     

Responses of cirripede larvae to light. I. Experiments with white light

The phototactic response of the nauplius larva of Balanus balanoides, B. crenatus and Elminius modestus shows darkadaptation; the response of the cyprid of B. balanoides shows both phototaxis and low photokinesis. The phototactic responses and the orientation of the cyprid to white light at settlement require an intensity of illumination slightly above 10^-5 lux. The ability to select a shaded position by cyprids of R. balanoides requires a higher intensity of 10^-2 to 10^-4 lux; hence a different mechanism may be involved. Barnacle larvae are sufficiently sensitive to be able to respond to light beneath the sea surface, even on cloudy, moonless nights.     

Comparative study of crustacean larval photoresponses

Ovigerous females of four brachyuran (Cancer gracilis, Lophopanopeus bellus bellus, Hemigrapsus oregonensis and Scyra acutiforns) and two anomuran (Pagurus beringanus and P. granosimanus) species, which live as adults in coastal areas, were collected near Friday Harbor, Washington, USA, in 1985, and spectral sensitivity and phototactic pattern of their larvae were measured. Responses were compared with previous measurements on estuarine species to determine whether responsiveness varies with adult habitat. Estuarine and coastal species have similar photoresponses. Spectral sensitivity of the test brachyran species had two maxima, one near 400 to 420 nm and another around 500 nm. The anomuran species were similar, but had an additional peak in the region of 580 to 620 nm. This sensitivity is adapted to daytime light conditions in the adult environment, and available spectra at the time of larval movement during diel vertical migration. Upon light adaptation and stimulation with a narrow light field, all six species showed positive phototaxis to high light intensities and a pronounced negative response to low intensities. Only the negative response will occur in natural underwater light conditions, and it is part of a predator-avoidance shadow-response which operates in areas of higher light intensity. The same phototactic pattern is observed in all species when darkadapted except H. oregonensis. If nocturnal vertical migration occurs, this negative response may be responsible for the descent at sunrise and depth maintenance during the day.     

The ontogeny of phototaxis by larvae of the crab Rhithropanopeus harrisii

Phototaxis by each zoeal stage of the crab Rhithropanopeus harrisii (Gould) was quantitatively measured by means of a microscope closed-circuit television system. The megalopa stage is indifferent to light stimulation and was thus not tested. The action spectrum for positive phototaxis is similar at each zoeal stage, having the most pronounced maximum at about 500 nm and smaller maxima at 400 and 280 nm. Responsiveness to various intensities of 500 nm light is also similar at each stage. After dark-adaptation strongest positive phototaxis occurs at intensities between 1.0 and 3.0x10^-4 W/m², with no negative phototaxis to lower intensities. After light-adaptation, the positive response occurs to higher intensities between 20 and 0.01 W/m², with a pronounced negative response to lower intensities. Generally, mean swimming speeds during positive phototaxis do not change with stimulation intensity, although during negative phototaxis mean speeds do vary with intensity. At the highest intensities which initiate a negative response, mean values are significantly greater than values for positive phototaxis. Based on the pattern of phototaxis at each developmental stage, a prediction of events during diurnal vertical migration is possible. The negative response after light-adaptation might also function as a shadow reflex.     

Phototaxis and cannibalism in gammaridean amphipods

Development changes in phototaxis were investigated in three species of estuarine amphipods: Gammarus lawrencianus. G. tigrinus, and G. mucronatus. In water of salinity 16 S, juveniles of G. tigrinus and G. mucronatus were initially photoneutral and became photonegative within two days after hatching. Juveniles of G. lawrencianus were photopositive on hatching and became photoneutral approximately 11 d later. In all three species, the change in phototactic behaviour coincided with the age at which juveniles became less vulnerable to cannibalism by adults. Evidence is presented showing genetic variation for phototactic behaviour in G. lawrencianus. Our data are consistent with the hypothesis that juvenile phototaxis is an evolutionary adaptation to minimise cannibalism by adults.     

Effects on growth and reproduction of the coral Stylophora pistillata by the mutualistic damselfish Dascyllus marginatus

Although coral dwelling fishes are common on coral reefs, the nature of their effect on the host corals is poorly understood. The present study, conducted in the Gulf of Eilat (Red Sea) between July 1989 and August 1990, demonstrated that the branching coral Stylophora pistillata (Esper) benefits, in two components of coral fitness, from the presence of the damselfish Dascyllus marginatus (Rüppell), an obligate coral dweller. The growth rate of damselfish-inhabited corals was significantly higher than that of corals without damselfish. This was observed, using two growth assessment methods, in long-term (>7 mo) comparisons between: (1) corals where the damselfish were experimentally removed versus corals with unaltered fish groups; and (2) naturally inhabited versus non-inhabited corals. The presence of damselfish did not affect the coral's specific (per surface area) reproductive output, whether it was assessed by the number of female gonads per polyp or by the number of planulae released cm^-2 surface area d^-1. However, the more rapid increase in branch size in damselfish-inhabited corals resulted in an apparent increase in the total reproductive output, with age, in growing corals. These findings demonstrate that the association between the damselfish D. marginatus and its host coral, S. pistillata, is mutualistic.     

Do corals select zooxanthellae by alternative discharge?

Loss of zooxanthellae (dinoflagellate Symbiodinium) from corals will sometimes lead to mass mortality of corals. To detect and quantify Symbiodinium released from corals, we developed a zooxanthellae “trap” and a quantitative PCR (qPCR) system with Symbiodinium clades A–F-specific primer sets. The trap was attached to a branch or the surface of several wild stony corals, and the water samples within the traps, including released Symbiodinium, were subjected to qPCR. All tested corals released clade C Symbiodinium at estimates of ~5,900 cells h⁻¹ cm⁻² of coral surface. Although all tested Pocillopora eydouxi harboured both clades C and D, some of these colonies released only clade C or released a lesser amount of clade D than that in the tissues. Our Symbiodinium quantification system revealed that wild hermatypic corals constantly release Symbiodinium to the environment. Our result suggests that some corals may discharge certain clades of Symbiodinium alternatively.     

Rafting of reef corals and other organisms at Kwajalein Atoll

Studies conducted at Kwajalein Atoll (9°N; 168°E) in early 1988 reveal that marine organisms are commonly rafted into the area on drift pumice, drift wood and other flotsam. Coralskeletons on pumice provide the most useful quantitative data because they persist after rafted motile organisms have departed and rafted sessile forms have decomposed or been scavenged. The estimated minimum number of pumice fragments carrying corals into Kwajalein Atoll during its geological history is on the order of 10⁹, with a more realistic estimate in excess of 10¹¹. The estimated number of coral colonies rafted into the atoll would probably have to be increased several-fold if rafting on floating organic materials such as wood, charcoal, nuts and seeds could be determined. In the present study, a typical sample of beached pumice from Kwajalein Atoll contained 10³ coral colonies per m³ of bulk pumice fragments. Major pumice rafting episodes frequently result from volcanic eruptions. Estimates based on the observed K wajalein coral fouling rate and literature values for drift pumice production suggest that 10⁵ colonies could be rafted through the tropics by the smallest reported pumice-producing events and up to 10¹² colonies for large events. In the present study, most of the rafted corals that were recovered belong to species of the genus Pocillopora, but speccics of Porites and Millepora were also collected. Analysis of surface-current data and reported drift patterns for pumice, logs, drift bottles and wrecks suggests that movement of rafted corals and drifting larvae is predominantly from peripheral areas of low coral-species diversity into centers of high coral diversity. Coral diversity centers might be viewed as areas of coral species accumulation rather than centers of coral species origin.     

Occurrence of a shadow response among brachyuran larvae

A closed-circuit television system was used to study phototaxis and behavioral responses upon sudden decreases in light intensity in light-adapted Stage I zoeae from 7 species of Brachyura (Panopeus herbstii, Menippe mercenaria, Uca pugilator, Callinectes sapidus, Sesarma cinereum, Pinnotheres maculatum, and Libinia emarginata). All species except one show positive phototaxis to high intensities of 500 nm light and negative phototaxis to low intensities. Upon a sudden light intensity decrease, larvae show a shadow response, which depends upon the initial and final intensities. If the initial intensity is sufficient to induce positive phototaxis, and the light is extinguished, the larvae of 6 species stop swimming and passively sink (sinking response). Reducing the intensity to an absolute level that causes negative phototaxis, induces a sinking response followed by negative phototaxis. For intensity reductions that occur in the intensity range above that for negative phototaxis, a sinking responseis observed. The intensity decrease that initiates the sinking response is independent of initial stimulus intensity and duration, is consistent between species, and is equivalent to a decrease by 0.5 OD (optical density) units. The sinking response is greatest at intensity decreases of at least 1.0 to 1.1 OD units. It is concluded that the shadow response is common in brachyuran larvae inhabiting coastal-estuarine areas.     

The effects of antifoulant-paint-contaminated sediments on coral recruits and branchlets

The 184-m cargo ship "Bunga Teratai Satu" ran aground on Sudbury Reef, within the Great Barrier Reef Marine Park, on 2 November 2000. Although no cargo or fuel was lost, the ship remained aground for 12 days and a large quantity of antifoulant paint containing tributyltin (TBT), zinc, and copper was scraped from the hull during the grounding and subsequent refloating operation. This resulted in extensive contamination of the reef sediments for up to 250 m surrounding the grounding site. Two laboratory-based experiments assessed the impact of contaminated sediments on the survival of both newly settled corals of Acropora microphthalma and branchlets of A. formosa. Newly settled corals exposed to sediments containing 8.0 mg kg^–1 TBT, 72 mg kg^–1 Cu, and 92 mg kg^–1 Zn or greater suffered significantly higher mortality after 72 h, compared to control or low-concentration treatments. Coral recruits exposed to 40 mg kg^–1 TBT (Sn), 306 mg kg^–1 Cu, and 403 mg kg^–1 Zn were all killed within 38 h. Branchlets from adult corals exposed to sediments with a high concentration of contaminants (TBT 160 mg kg^–1, Cu 1,180 mg kg^–1, and Zn 1,570 mg kg^–1) suffered significant mortality (38%), whereas branchlets placed in treatments with lower levels of contaminants suffered no mortality. Visual bleaching of the branchlets was observed at high contaminant levels, but an overall reduction in the symbiotic zooxanthellae populations was not observed in surviving corals. The photosynthetic yields of light-adapted zooxanthellae remained constant in live branchlets, indicating that the TBT-contaminated sediment may be more toxic to the host than the symbiont. Our results show that antifoulant contamination at ship-grounding sites has the potential to cause major mortality of resident coral communities and can have a negative impact on the recovery of adult populations.Communicated by P.W. Sammarco, Chauvin     

Larval behavior of predacious sister-species: orientation,molting site,and survival in Chrysopa

Field and laboratory studies compared two features of larval behavior in a pair of predacious sisterspecies of green lacewings: one (Chrysopa slossonae) a specialist on a single species of colonial aphids (the woolly alder aphid) that occur on branches and trunks of alder trees, the other (C. quadripunctata) a general aphid feeder whose primary prey is dispersed on foliage of diverse types of trees. First, a few hours after hatching, larvae of the two species develop significantly different phototactic responses; the differences correspond well with the spatial distributions of their prey. Most C. slossonae exhibited negative phototaxis, a response that helps move hatchlings inward on alder trees toward the woolly alder aphid colonies, whereas most C. quadripunctata hatchlings showed positive orientation to light, a response that tends to keep them in tree canopies with their prey. Second, in greenhouse experiments, a significantly greater proportion of C. slossonae larvae (second instars) molted within woolly alder aphid colonies and remained with the aphids than did C. quadripunctata larvae. These differences indicate that the specialist larvae have evolved a high degree of behavioral fidelity to their prey. However, larvae (second instars) of the two species that were released near ant-tended woolly alder aphid colonies in the field had similar recovery (= survival) rates. Consequently, natural selection may not act on behavioral traits that influence larval fidelity to prey during the late second and early third instars.     

Distribution of deep-water gorgonian corals in relation to benthic habitat features in the Northeast Channel (Atlantic Canada)

The distribution and abundance of deep-water gorgonian corals were investigated along 52 transects at 183–498 m depth in the Northeast Channel, between Georges Bank and Browns Bank in the northwest Atlantic, using a remotely operated vehicle and a towed video-camera system. Three species (Paragorgia arborea, Primnoa resedaeformis, and Acanthogorgia armata) were observed. Primnoa occurred on 35 transects below 196 m depth, with highest local abundance in stands of 104 colonies per 100 m². Paragorgia was present on 21 transects deeper than 235 m, with highest local abundance of 49 colonies per 100 m². Acanthogorgia was observed at only four transects between 231 m and 364 m, with a local maximum abundance of 199 colonies per 100 m². The maximum abundance averaged for whole transects was 19.2 and 6.2 colonies per 100 m² for Primnoa and Paragorgia, respectively. The corals were more common in the outer part of the channel along the shelf break and slope than on the shelf in the inner part. All three species showed a patchy distribution with no signs of competitive exclusion at any spatial scale. Transects with high abundance of corals were characterised by depths greater than 400 m, maximum temperatures less than 9.2°C, and a relatively high percentage coverage of cobble and boulder (more than 19% and 6%, respectively). High temperatures probably control the upper depth limit of the corals, and Primnoa seems to tolerate slightly higher temperatures than Paragorgia. Abundance of both species was negatively correlated with average temperature and positively with cobbles. Together, temperature, percentage cobble and salinity accounted for 38% of the variance of Primnoa. The comparable figure for Paragorgia was 15%. The observed distribution indicated that the abundance of coral is controlled by additional factors such as larger-scaled topographic features governing the current regimes and thus also the supply of food and larvae.Communicated by R. J. Thompson, St. Johns     

Allometric variability of heavy metals in corals of Lakshadweep islands

Heavy metal contents of the corals collected from the Lakshadweep archipelago are discussed in the light of allometric variations. Plots of log flesh weight versus log flesh metal concentration exhibited negative correlations for all the nine metals studied. Metals with rapid exchange rate are likely to show such negative correlation and in such situations the elimination rate far exceeds the metal accumulation rate. Similar plots in the skeletal component showed positive correlations. The only linear relationships were observed in log flesh density versus log partition coefficient plots. Comparative flesh enrichment of metals were found to decrease against increasing flesh weights (for both essential and non-essential metals), indicating a regulation of flesh metal burden beyond certain limits. In the skeletal component, anthropogenic input as well as competitive interactions between metal ions having radii similar to that of Ca²⁺ paves way for metal adsorption in calcite/aragonite skeletal lattice of corals.     

Initial observations of the zooplankton microdistribution on the fringing coral reef at Aqaba (Jordan)

Diurnal and nocturnal zooplankton was sampled simultaneously at the surface and at 35 m depth on the outer slope of the fringing reef off the Jordanian coast in the Guff of Aqaba, Red Sea, over a period of 8 d during July 1982. Complementary surface sampling was carried out above the reef and in the open sea in the morning and afternoon for a period of 5 d. Five groups of organisms were classified according to changes in their vertical distribution in the water column which is analyzed and discussed. Zooplankters of the first group appear to be positively phototactic (Calanus robustior, Mecynocera clausi, Oithona helgolandica, Corycella rostrata). Those of the second group also seem to be positively phototactic, but only as regards average and low light (moonlight) intensities (Nannocalanus minor, Acartia negligens, Sagitta enflata, S. hexaptera, Thalia democratica, Oikopleura fusiformis, O. longicauda). The zooplankton of the third group displays a negative phototaxis (Oithona nana, calyptopis and furcilia euphausiid stages, pelecypods and other mollusc veligers). Species exhibiting uncertain behaviour or caught in low numbers and exhibiting no particular distribution are relegated to the fourth and the fifth groups. The horizontal microdistribution of the zooplankton community is not easily discernible: certain neritic or oceanic affinities can be identified, however, by comparing the WP2 standard net surface samples collected above the reef with those from the open sea. The importance of zooplankton communities is emphasized, and the necessity of elucidating the temporal and spatial migrations and exchanges between reef and planktonic organisms in order to gain more insight into their trophic relationship with the reef system.     

Behavioral responses of larvae of the crab Rhithropanopeus harrisii (Brachyura: Xanthidae) during diel vertical migration

Larval Rhithropanopeus harrisii (Gould) show nocturnal vertical migration. Larval behavioral responses to different rates of increase and decrease in light intensity were measured in an apparatus with a natural angular light distribution. A central objective was to establish whether phototaxis actually participates in vertical migration. At sunset the level of light adaptation controlled the readiness of the larvae to migrate, while an ascent was initiated by a preductable relative decrease in intensity (e.g. 4.0x10^-3s^-1). Rates of relative decrease around sunset would evoke continuous upward swimming. Gravity was the orienting cue and there was no change in swimming speed during the ascent. At sunrise, the larval descent was initiated by exposure to an absolute light intensity of about 0.23 log unit above the lower visual threshold. Light served as an orienting cue, as larvae descended by a negative phototaxis. Thus, phototaxis is not a laboratory artifact and does participate in vertical migration. A consideration of behavioral responses of other crustacean zooplankton indicates there is considerable variation in the initiating and directing cues for vertical movements. The variety of behavioral responses of R. harrisii suggests that a synthesis of hypotheses about migration may provide the proper basis for explaining the mechanisms underlying diel vertical migration.     

Coral bleaching, bleaching-induced mortality, and the adaptive significance of the bleaching response

Coral bleaching events are often associated with higher levels of coral mortality but when this occurs in the chronology of individual bleaching events is poorly documented. Knowing when mortality occurs is important for understanding molecular mechanisms and the putative adaptive significance of the response (the Adaptive Bleaching Hypothesis). In a detailed study of a coral bleaching event on the Great Barrier Reef, involving weekly and twice weekly repetitive observations of >200 individually marked corals over an 18 month period (∼16,000 observations), it is shown that bleaching in Acropora latistella, A. subulata and Turbinaria mesenterina was an acute, rapid response, occurring within days of a peak in seawater temperatures exceeding previously described thresholds. Subsurface light levels, measured over the duration of the event, were not anomalous. Full bleaching (i.e. whole colonies turning bone-white) and partial bleaching (white patches) was observed in the Acropora spp. whilst the T. mesenterina colonies typically paled to a light brown colour. Algal densities in bleached corals were 10–30% of those of normally pigmented corals (∼2.5 × 10⁶ algae per cm²), and in this instance bleaching was clearly a sudden, isolated, stress event and not an extreme low-point in the seasonal fluctuation of the density of symbiotic algae. Bleached corals were associated with high levels of partial and whole-colony mortality, but mortality was exclusively limited to the two Acropora spp. Importantly, most of this mortality was recorded in surveys conducted 1 and 2 weeks after bleaching was first observed, and for A. latistella as little as 1 week after bleaching was first observed. This suggests that in this particular bleaching event, for the Acropora species, that bleaching and mortality were intimately linked: this in turn suggests it was a pathological phenomenon. The study highlights a problem in the adaptive bleaching hypothesis, whereby significant levels of mortality can occur in a bleaching event before any chance for subsequent recombination of the host-symbiont unit. It is argued that in order to further evaluate the significance of bleaching as a potentially adaptive mechanism, bleaching-induced and bleaching-related mortality have to be fully considered. It is necessary to incorporate the cost (in terms of mortality) of a bleaching event, the recurring cost of reverting to the original, mortal, stress–prone combination after the event, and the higher cost associated with forming a maladaptive combination.     

Experimental ecology of the temperate scleractinian coral Astrangia danae I. Partition of respiration,photosynthesis and calcification between host and symbionts

Calcification, photosynthesis and respiration of the scleractinian coral Astrangia danae were calculated from the changes in total alkalinity, pH, calculated total CO₂, and oxygen concentration produced by colonies incubated in glass jars. A correction for changes in ammonia, nitrate and nitrite was taken into account and the method evaluated. The fluxes of oxygen and CO₂ were highly correlated (r=0.99). The statistical error of alkalinity determinations was less than 10% of the changes observed in the slowest calcifying samples. Metabolism of polyparium alone was estimated by difference after removal of tissue and reincubation of bare corallum. Zooxanthellae concentration in the polyps was obtained from cell counts made on homogenates of polyp tissue. The calculated photosynthetic rate of the zooxanthellae in vivo was 25 mol O₂ (10⁸ cell)^-1 h^-1 at a light intensity of 120 Ein m^-2 s^-1. In corals having 0.5x10⁹ zooxanthellae/dm² of colony area up to 8% of the total photosynthesis was attributed to the corallum microcosm. Polyp respiration, photosynthesis, and CaCO₃ uptake rates were all much higher than rates previously reported from A. danae, apparently because in these experiments the organisms were better fed. This increased photosynthesis in turn enhanced calcification still further. The symbiosis therefore appears to provide a growth advantage even to fed corals, under the conditions of these experiments.     

Laboratory observations on spawning,fecundity and early development of a mesopelagic ostracod,Conchoecia pseudodiscophora,from the Japan Sea

The fate of the polycyclic aromatic hydrocarbon [³H]benzo[a]pyrene (BaP) was examined in two species of scleractinian corals, Favia fragum (Esper) and Montastrea annularis (Ellis and Solander), which were collected in the patch reefs surrounding Alina's Reef (25°23.25N; 80°09.8W) in Biscayne National Park, Florida, USA, in July, 1990. Corals were exposed to initial concentrations of 5 g/l in a simple static system for 25 h. BaP uptake was estimated from the disappearance of BaP from the water. Uptake rates were 6.5±0.7 and 10.8±0.2 g BaP cm^-2h^-1 for F. fragum and M. annularis, respectively, at initial BaP concentrations and were directly proportional to the concentration of BaP in the water. The separation of zooxanthellae from coral tissue revealed that zooxanthellae can accumulate up to 53 and 64% of the total BaP-derived radioactivity present in F. fragum and M. annularis, respectively. Both corals metabolized BaP slowly, as most of the accumulated radioactivity was present as the unmetabolized chemical. However, aqueous and organic-soluble metabolites were found in both the animal and zooxanthellae fractions. Analysis by high-performance liquid chromatography (HPLC) revealed that both species of corals metabolized BaP to various tetrols, triols, dihydrodiols, quinones and phenols, although the pattern of metabolites differed between species. Zooxanthellae contained some of the same Phase I metabolites found in the animal tissue; however, tetrols and triols were absent in extracts from the zooxanthellae. The elimination of BAP from corals was also slow; approximately 38 and 65% of the accumulated radioactivity was still present in F. fragum and M. annularis, respectively, 144 h following the transfer of exposed corals to an uncontaminated flow-through seawater system.     

Water flow and prey capture by three scleractinian corals, Madracis mirabilis, Montastrea cavernosa and Porites porites, in a field enclosure

Scleractinian corals experience a wide range of flow regimes which, coupled with colony morphology, can affect the ability of corals to capture zooplankton and other particulate materials. We used a field enclosure oriented parallel to prevailing oscillatory flow on the forereef at Discovery Bay, Jamaica, to investigate rates of zooplankton capture by corals of varying morphology and polyp size under realistic flow speeds. Experiments were carried out from 1989 to 1992. Particles (Artemia salina cysts) and naturally occurring zooplankton attracted into the enclosures were used as prey for the corals Madracis mirabilis (Duchassaing and Michelotti) (narrow branches, small polyps), Montastrea cavernosa (Linnaeus) (mounding, large polyps), and Porites porites (Pallas) (wide branches, small polyps). This design allowed corals to be used without removing them or their prey from the reef environment, and avoided contact of zooplankton with net surfaces. Flow speed had significant effects on capture rate for cysts (M. mirabilis), total zooplankton (M. mirabilis, M. cavernosa), and non-copepod zooplankton (M. mirabilis). Zooplankton prey capture increased with prey concentration for M. mirabilis and M. cavernosa, over a broad range of concentrations, indicating that saturation of the feeding response had not occurred until prey density was over 10⁴ items m⁻³, a concentration at least an order of magnitude greater than the normal range of reef zooplankton concentrations. Location of cyst capture on coral surfaces was not uniform; for M. cavernosa, sides and tops of mounds captured most particles, and for P. porites, capture was greatest near branch tops, but was close to uniform for M. mirabilis branches in all flow conditions. The present study confirms laboratory flume results, and field results for other species, suggesting that many coral species experience particle flux and encounter rate limitations at low flow speeds, decreasing potential zooplankton capture rates. Received: 17 September 1996 / Accepted: 22 November 1997     

Coral mucus stable isotope composition and labeling: experimental evidence for mucus uptake by epizoic acoelomorph worms

Mucus released by scleractinian corals can act as an important energy and nutrient carrier in coral reef ecosystems, and a distinct isotopic signature would allow following the fate of this material. This study investigates the natural C and N stable isotopic signatures of mucus released by four scleractinian coral genera (Acropora, Fungia, Pocillopora and Stylophora) in comparison with those of suspended particulate organic matter (POM) in seawater of a Northern Red Sea fringing coral reef near Aqaba, Jordan. The natural δ¹³C and δ¹⁵N signatures of coral mucus differed significantly from seawater POM for the majority of seasonal comparisons, but were inappropriate for explicit tracing of mucus in the coral reef food web. Thus, a labeling technique using stable isotope tracers (¹³C and ¹⁵N) was developed that produced δ¹³C values of up to 122 ± 5‰ (mean ± SE) and δ¹⁵N of up to 2,100 ± 151‰ in mucus exuded by Fungia corals. ¹³C and ¹⁵N-enriched compounds were rapidly (within 3 h) and light-dependently transferred from the endosymbiotic zooxanthellae to the mucus-producing coral host. The traceability of ¹⁵N-labeled mucus was examined by evaluating its uptake and potential utilization by epizoic acoelomorph Waminoa worms naturally occurring on a range of scleractinian coral taxa. This tracer experiment resulted in uptake of coral mucus by the coral-associated acoelomorphs and further demonstrated the possibility to trace stable isotope-labeled coral mucus by revealing a new trophic pathway in coral reef ecosystems.