Effects of monsoon-driven wave action on coral reefs of Guam and implications for coral recruitment

Benthic cyanobacteria can respond rapidly to favorable environmental conditions, overgrow a variety of reef organisms, and dominate benthic marine communities; however, little is known about the dynamics and consequences of such cyanobacterial blooms in coral reef ecosystems. In this study, the benthic community was quantified at the time of coral spawnings in Guam to assess the substrate that coral larvae would encounter when attempting settlement. Transects at 9, 18, and 25-m depths were surveyed at two reef sites before and after heavy wave action driven by westerly monsoon winds. Communities differed significantly between sites and depths, but major changes in benthic community structure were associated with wave action driven by monsoon winds. A shift from cyanobacteria to crustose coralline algae (CCA) accounted for 44% of this change. Coral recruitment on Guam may be limited by substrate availability if cyanobacteria cover large areas of the reef at the time of settlement, and consequently recruitment may in part depend upon wave action from annual monsoon winds and tropical storms which remove cyanobacteria, thereby exposing underlying CCA and other substrate suitable for coral settlement.     

Engineering of coral reef larval supply through transplantation of nursery-farmed gravid colonies

The continuous worldwide degradation of coral reefs raises an urgent need for novel active restoration techniques as traditional conservation practices have failed to impede the incessant reefs' decline. While applying the “gardening coral reefs” methodology in Eilat (Red Sea, Israel), we examined reproductive outputs of naturally-grown and outplanted, nursery-farmed Stylophora pistillata colonies from three coral-transplantation trials (November 2005, May 2007, and September 2008), along three reproductive seasons. Surprisingly, transplanted colonies showed better reproductive capacities than the natal Stylophora colonies during > 4 post-transplantation years. A higher percentage of nursery-farmed colonies released planula larvae as compared to naturally-grown colonies. Gravid transplants also shed more planulae per colony, yielding significantly augmented numbers of total planulae over naturally developed S. pistillata colonies. Our results indicate that nursery-grown corals may be used to enhance reef resilience by contributing to the larval pool, forming an engineered larval dispersal instrument for reef rehabilitation.     

The application of acoustic Doppler current profilers to measure the timing and patterns of coral larval dispersal

An experiment was conducted along the reefs off west Maui, Hawaii, during the summer of 2003 to monitor the spawning of the reef-building coral Montipora capitata and to determine the role of ocean currents in dispersing the larvae from the natal reef. Instruments documented the environmental forcing during the coral spawning season; drifters were deployed on three successive nights following direct observations of coral spawning. Both the timing and relative magnitude of the coral spawning were identifiable in acoustic backscatter data and correlated to plankton tow data. Each drifter track showed that the surface water containing coral eggs and planula larvae were transported rapidly offshore and not locally retained. Wind and current patterns during the previous year and during subsequent coral spawning events later in the summer were similar to those observed during the drifter releases. This suggests that the trajectories observed during the focused experiment are representative of the general pattern of larval dispersal off west Maui. These findings demonstrate the application of acoustic profilers for remotely imaging coral spawning and predicting their initial dispersal patterns.     

Effects of pressure on swimming behavior in planula larvae of the coral Porites astreoides (Cnidaria, Scleractinia)

Mechanisms governing the behavior of coral planulae are not well understood, particularly those manifesting themselves between the time when the larvae are released and when they settle. Larvae from the hermatypic coral Porites astreoides Lamarck were exposed to different levels of hydrostatic pressure—103.4, 206.9, 310.3, 413.8, and 517.1 kPa (including ambient pressure). Data were collected at stops of the above pressures for 15 min each, respectively. This was done in both an increasing sequence and a decreasing one. When exposed to increases in pressure from 103.4 kPa, larvae swam upward (negative barotaxis) in a spiraling motion. Upon exposure to decreasing pressure from 517.1 kPa, larvae moved downward (positive barotaxis), but the magnitude of the vertical movement was much less than in the case of increasing pressure. This suggests that these larvae are more sensitive to increased pressure than decreasing pressure. High variance was also observed in the responses of these larvae at both the intra- and inter-colony levels. Thus, this behavioral trait is variable within the population. The trait may be genetically based, and thus may be susceptible to alteration by natural selection, although this remains to be demonstrated. This study is the first to document these behavioral mechanisms in coral larvae.     

The biology of coral metamorphosis: molecular responses of larvae to inducers of settlement and metamorphosis

Like many other cnidarians, corals undergo metamorphosis from a motile planula larva to a sedentary polyp. In some sea anemones such as Nematostella this process is a smooth transition requiring no extrinsic stimuli, but in many corals it is more complex and is cue-driven. To better understand the molecular events underlying coral metamorphosis, competent larvae were treated with either a natural inducer of settlement (crustose coralline algae chips/extract) or LWamide, which bypasses the settlement phase and drives larvae directly into metamorphosis. Microarrays featuring > 8000 Acropora unigenes were used to follow gene expression changes during the 12 h period after these treatments, and the expression patterns of specific genes, selected on the basis of the array experiments, were investigated by in situ hybridization. Three patterns of expression were common—an aboral pattern restricted to the searching/settlement phase, a second phase of aboral expression corresponding to the beginning of the development of the calicoblastic ectoderm and continuing after metamorphosis, and a later orally-restricted pattern.     

The role of epibotic bacteria from the surface of the soft coral Dendronephthya sp. in the inhibition of larval settlement

It has been suggested that bacteria associated with soft-bodied organisms are suggested to produce bioactive compounds against the attachment of invertebrate larvae and bacteria onto the surface of these organisms. Our recent study has demonstrated that epibiotic bacteria from the surface of the soft coral Dendronephthya sp. (Coelenterata: Octocoralia, Alcyonacea) inhibit the growth of bacteria commonly found in marine natural biofilms. In the present study, the effect of 11 epibiotic bacteria isolated from the surface of Dendronephthya sp. on larval settlement of the tubeworms Hydroides elegans was examined using laboratory bioassay. Among 11 bacterial isolates, 2 strains (18%) inhibited the larval settlement of H. elegans (Haswell), 4 strains (36%) were “inductive” to larvae and the remaining 5 strains (46%) were “non-inductive”. There was no correlation between the antifouling activities of bacterial isolates and their phylogenetic origin, i.e. closely related bacterial strains showed different effects on larval settlement of H. elegans. When all “inductive”, “non-inductive” and “inhibitive” bacterial isolates were mixed in a 1:1:1 ratio, the effect of the resultant multispecies film on larval settlement became “inhibitive”. Waterborne compounds of Vibrio sp. and an unidentified α-Proteobacterium, which suppressed the settlement of H. elegans and Bugula neritina (L.) larvae, were further investigated using size fractionation and bioassay-guided enzymatic analysis. It was found that antilarval settlement compounds from these bacteria were heat-stable polysaccharides with a molecular weight >100 kDa. The results indicate that the bacteria associated with the soft coral Dendronephthya sp. may contribute to the antifouling mechanisms of the soft-bodied organisms by producing compounds that are against bacterial growth and settlement of macrofoulers on the surface of their host.     

Within-colony variation in inducibility of coral disease resistance

The abiotic environment influences a variety of ecological processes, including the emergence, transmission, and distribution of disease. In the oceans, increased temperatures associated with climate change are hypothesized to decrease host resistance and/or increase pathogen growth, virulence, or infectivity. Colonial organisms, such as corals, could face a unique challenge with respect to temperature and disease stress: heterogeneous within-colony distribution of constitutive and temperature-induced resistance to infection. This could facilitate disease if warming temperatures promote pathogen growth while decreasing resistance of some areas of the coral colony. Here, an experiment was used to test the hypothesis that temperature-induced disease resistance is heterogeneous within colonies of the sea fan coral, Gorgonia ventalina. Resistance, measured as activity of antifungal metabolites, increased (approx. 30%) with temperature only in young edge tissue, not in older center tissue, consistent with patterns of infection in older, larger sea fan colonies on Caribbean reefs.     

Temporal settlement patterns of larvae of the broadcast spawning reef coral Favites chinensis and the broadcast spawning and brooding reef coral Goniastrea aspera from Okinawa, Japan

The faviid corals, Favites chinensis and Goniastrea aspera are widely distributed in the Indo-Pacific region. Both corals are hermaphroditic broadcast spawners, but G. aspera is also known to brood planula larvae in Okinawa. This study investigated the temporal settlement patterns of planula larvae of the scleractinian corals F. chinensis and G. aspera that developed from spawned gametes, and planula release and settlement of brooded larvae of G. aspera from Okinawa, Japan. Some of the broadcast-spawned larvae of F. chinensis and G. aspera had very short pre-competency periods of 1–2 and 2–3 days after spawning, and relatively long maximum settlement-competency periods of 56–63 and 63–70 days after spawning, respectively. These pre-competency periods are among the shortest reported for larvae of broadcast spawning coral species, and appear to be negatively correlated with seawater temperature. F. chinensis larvae tended to settle rapidly with 34–39% of larvae settling in the first week after spawning, while broadcast-spawned G. aspera larvae had a slower settlement pattern with 11–15% of larvae settling in the first week after spawning. Brooded larvae of G. aspera settled more rapidly, with settlement rates of 27–31% within the first 24 h and 45–65% within the first week after the start of the experiment. The production of planula larvae with rapid settlement capabilities may enable F. chinensis and G. aspera to establish and maintain populations in shallow reef sites at Okinawa. The release of the brooded planulae for up to 2 months may explain why G. aspera is locally more dominant on shallow reefs in Okinawa than F. chinensis. On a broader scale, the longer settlement competency periods of some of the broadcast-spawned larvae of these species increase their potential for longer-distance dispersal and may partly explain the wide biogeographic distribution of these species in the Indo-Pacific region.     

Effects of competition and herbivory on interactions between a hard coral and a brown alga

Despite widespread acceptance of the negative effects of macroalgae on corals, very few studies have experimentally tested the competitive nature of the interaction, and most have ignored the potential effects of corals on algae. We report the effects of herbivory and competition on the growth of the branching scleractinian coral Porites cylindrica Dana and the creeping foliose brown alga Lobophora variegata (Lamouroux) Womersley, on an inshore fringing reef of the central Great Barrier Reef. L. variegata overgrows branches of P. cylindrica from the base up, forming a distinct boundary between the alga and the coral tissue. The experiment used exclusion cages to test for effects of herbivores, and removal of algae and coral tissue, at their interaction boundary, to test for inhibition of the competitors by each other. Comparisons of coral branches with the algae present or removed showed that the presence and overgrowth of the alga caused significant coral tissue mortality. Comparisons of branches with coral tissue unmanipulated or damaged showed that the coral inhibited the overgrowth by L. variegata, but that the algae were markedly superior competitors. Importantly, reduced herbivory resulted in faster algal growth and consequent overgrowth and mortality of coral tissue, demonstrating the critical importance of herbivory to the outcome of the competitive interaction.     

The expression of the let-7 small regulatory RNA is controlled by ecdysone during metamorphosis in Drosophila melanogaster

In Caenorhabditis elegans, the heterochronic pathway controls the timing of developmental events during the larval stages. A component of this pathway, the let-7 small regulatory RNA, is expressed at the late stages of development and promotes the transition from larval to adult (L/A) stages. The stage-specificity of let-7 expression, which is crucial for the proper timing of the worm L/A transition, is conserved in Drosophila melanogaster and other invertebrates. In Drosophila, pulses of the steroid hormone 20-hydroxyecdysone (ecdysone) control the timing of the transition from larval to pupal to adult stages. To test whether let-7 expression is regulated by ecdysone in Drosophila, we used Northern blot analysis to examine the effect of altered ecdysone levels on let-7 expression in mutant animals, organ cultures, and S2 cultured cells. Experiments were conducted to test the role of Broad-Complex (BR-C), an essential component in the ecdysone pathway, in let-7 expression. We show that ecdysone and BR-C are required for let-7 expression, indicating that the ecdysone pathway regulates the temporal expression of let-7 in Drosophila. These results demonstrate an interaction between steroid hormone signaling and the heterochronic pathway in insects.     

Keeping and breeding of the coral catshark (Atelomycterus marmoratus) at the Aquarium of the Cologne Zoo

Coral catsharks (Atelomycterus marmoratus) are kept at the Aquarium of Cologne Zoo in a public exhibit tank measuring 300 x 160 x 110 cm (length x width x water height) with a water volume of ca. 5000 l. The 1.3 adults have been producing fertilized eggs since early 2010. Deposited eggs are longish-oval in shape, with a length of about 10 cm. Juveniles hatched in our rearing tanks (measuring 100-210 x 45-80 x 30-50 cm, with 280-430 l water volume) after periods of 4-6 months. The freshly hatched coral catsharks measured from 10-13 cm in total length. After 3 months, each juvenile had grown about 4-5 cm in length and the distinct, bright color pattern of the juveniles had faded towards the more subtle adult colour pattern. The keeping and breeding of coral catsharks can be classified as uncomplicated, given that sufficient swimming space and hiding places are provided. So far, 14 juvenile coral catsharks have been successfully reared at the Cologne Zoo Aquarium and are now available for breeding projects.     

Larval identification of Lutjanus Bloch in Nansha coral reefs by AFLP molecular method

Species of Lutjanus Bloch are highly valued fish in some fisheries of the world and some species are cultured in the Southeast Asia especially in South China. Wild larvae are still the major source of mariculture in South China because artificial breeding techniques for most Lutjanus species have not yet been available. The Nansha coral reefs (also called Spratly Archipelago) water area, which is located in the South China Sea, is the main habitat and spawn area for Lutjanus in China. Larval identification of Lutjanus is important for relative ecological studies and mariculture, but larvae of many closely related species, such as those of the genus Lutjanus, are different to be distinguished morphologically. In the present study, a PCR-based fingerprinting technique called amplified fragment length polymorphism AFLP was used in the characterization and identification of 11 Lutjanus species captured in Nansha coral reefs. Optimal AFLP patterns were obtained with primer combination of E+AGC/M+CAA selective nucleotides. There were in total 132 AFLP loci in all specimens, and AFLP markers of each species varied from 44 to 69, but only 7 markers were fixed in all specimens. Meanwhile, high levels of intraspecific homogeneity were observed. All 11 species of Lutjanus were successfully identified by the comparative analyses of AFLP patterns. Moreover, neighbour-joining and UPGMA analyses of AFLP data were compared with current morphological taxonomic systems.     

Improvements to a passive trap for quantifying barnacle larval supply to semi-exposed rocky shores

Planktonic cyprid larvae of the intertidal barnacle, Semibalanus balanoides (L.) and the infralittoral/sublittoral Balanus crenatus Brug., are readily captured in small, chambered, passive traps deployed on rocky shores. Several structural developments of a previous trap design improved urea (killing solution) retention and capture of cyprids of both species. The design adopted for assessment of the larval supply/settlement relationship of the focal species, S. balanoides, included additional internal baffles and the replacement of the cylindrical trap opening with a coned aperture. Three sizes of cone aperture area were compared, with the intermediate size (1 cm²) being optimal for measuring larval supply over the full range of wave action on local shores. Most daily catches included larvae that were recognizable as having been dead before they were trapped, and had recently passed through the guts of grazing limpets (Patella vulgata L.). One site showed that ∼32% of trapped S. balanoides were so-called ‘faecal’ cyprids. An inability to distinguish these in trap samples would lead to a significant over-estimation of larval supply. Preliminary data indicated that S. balanoides larval supply to the intertidal is enhanced by onshore winds (or perhaps increased wave turbulence), whereas B. crenatus supply was positively correlated with offshore winds (or decreased turbulence). A clear ‘supply/settlement’ relationship for S. balanoides at six sites along ∼40 km of coastline was derived from 1 cm² coned trap catches and daily counts of settlement on ceramic tiles. There were no instances of disproportionately high settlement in relation to larval supply, but at five sites there were days when settlement apparently failed. The identification and explanation of settlement success and failure is fundamental to understanding the importance of larval supply in structuring marine benthic populations and communities.     

Effects of nutrient enhancement on the fecundity of a coral reef macroalga

On coral reefs, there is concern that increased nutrient supply (e.g. due to eutrophication) causes increased algal growth and hence increased algal abundance, in turn causing colonisation and invasions of coral populations, resulting in reef degradation, or a coral-algal phase shift. For example, species of Sargassum, a highly seasonal, large, brown seaweed, are suggested to be colonising corals on inshore coral reefs of the Great Barrier Reef, as a result of anthropogenic increases in terrestrial runoff of sediments and nutrients. However, implicit in this argument is the assumption that nutrient-related increases in growth will lead to increased fecundity (and/or propagule success), since without such changes, increased abundance can only occur by vegetative means. Whilst plausible, there is no experimental evidence for this assumption in coral reef algae.We here present an initial study in which experimental increases in nutrient supply apparently did not lead to increased fecundity of Sargassum siliquosum; rather, density and biomass of receptacles were reduced in nutrient-enhanced algae. There was little effect of nutrient treatments on the proportional allocation of biomass to reproductive and vegetative structures: nutrient enhancement led to similar decreases in biomass of reproductive and vegetative tissue. Tissue nutrient levels indicated effective enhancement of nitrogen supply, although phosphorus levels were not different across nutrient treatments.The reduced fecundity with increased nutrients may reflect either a genuine inhibition by higher nutrient levels, as found in previous studies, or accelerated maturation, causing increased tissue losses due to more advanced seasonal senescence. Either way, this exploratory experiment provides initial evidence that nutrient effects on tropical coral reef macroalgae may be complex, and does not support the assumption that increased nutrient supply will result in a numeric increase in populations of Sargassum spp. Our results should not be taken as an unequivocal demonstration that nutrients inhibit fecundity overall, but illustrate the need to distinguish between effects on different life-history processes (e.g. growth and reproduction). For increased growth of pre-existing individuals to contribute to algal invasions and phase shifts, that growth must result in either increased fecundity, or increased propagule success.     

Activities of natural methyl farnesoids on pupariation and metamorphosis of Drosophila melanogaster

Methyl farnesoate (MF) and juvenile hormone (JH III), which bind with high affinity to the receptors USP and MET, respectively, and bisepoxy JH III (bisJH III) were assessed for several activities during Drosophila larval development, and during prepupal development to eclosed adults. Dietary MF and JH III were similarly active, and more active than bisJH III, in lengthening larval development prior to pupariation. However, the order of activity was changed (JH III > bisJH III > MF) with respect to preventing prepupae from eclosing as normal adults, whether administered in the larval diet or as topically applied at the white puparium stage. If endogenous production of all three larval methyl farnesoids was suppressed by a strongly driven RNAi against HMGCR in the corpora allata cells, most larvae did not attain pupariation. Farnesol (which has no demonstrated life-necessary function in larval life except in corpora allata cells as a precursor to methyl farnesoid biosynthesis) when incorporated into the diet rescued attainment of pupariation in a dose-dependent manner, presumably by rescuing endogenous production of all three hormones. A more mild suppression of endogenous methyl farnesoid production enabled larval attainment of pupariation. However, in this background dietary MF had increased activity in preventing puparia from attaining normal adult eclosion. The physiological relevance of using exogenous methyl farnesoids to block prepupal development to normally eclosed adults was tested by, instead, protecting in prepupae the endogenous titer of methyl farnesoids. JH esterase normally increases during the mid-late prepupal stage, presumably to clear endogenous methyl farnesoids. When JH esterase was inhibited with an RNAi, it prevented attainment of adult eclosion. Cultured adult corpora allata from male and female Aedes aegypti released both MF and JH III, and the A. aegypti nuclear receptor USP bound MF with nanomolar affinity. These A. aegypti data support the use of Drosophila as a model for mosquitoes of the binding of secreted MF to USP.     

Effect of dietary polyunsaturated fatty acids on reproductive output and larval growth of bivalves

The pre-spawning condition of adult bivalves is influenced by quantity and quality of available food. For bivalves, the essential polyunsaturated fatty acids (PUFAs) eicosapentaenoic acid (EPA) 20:5(n-3) and docosahexaenoic acid (DHA) 22:6(n-3) are presumed to determine the nutritional value of algae. Macoma balthica kept on a broodstock diet supplemented with PUFAs spawned a larger number of eggs (average 22220) per female and larger sized eggs (106.7 μm) compared to adults kept on a diet without PUFA supplementation (962.5 eggs with average size 99.8 μm).Larvae of M. balthica from the same parental pool however did not profit from a diet where a part was replaced with PUFA spheres. Instead, larvae reared on Isochrysis sp. showed lower mortality and higher growth rates than larvae fed on the same algae supplemented with lipid spheres. Crassostrea gigas larvae showed no clear response to a PUFA supplemented diet.     

Effects of the fish anesthetic, clove oil (eugenol), on coral health and growth

Ecological research within coral reefs often requires the use of anesthetics to immobilize organisms. It is therefore important to consider the effect of these chemicals on the surrounding flora and fauna, particularly to the corals themselves. We quantified the effects of clove oil, a commonly used fish anesthetic, on the growth and occurrence of bleaching in three species of corals: Acropora striata, Pocillopora verrucosa, and Porites australiensis. We compared coral responses to five treatments: a gradient of four clove oil concentrations (0-28%) in seawater, and one concentration of clove oil (14%) in ethanol. Each week, we assessed the presence of bleaching, and then applied the treatment. We measured growth over the duration of the 6-week experiment using the buoyant weight technique. Growth and bleaching showed a dose response to clove oil exposure, and the use of ethanol as a solvent had an additional deleterious effect, as also suggested by observed changes in concentrations of eugenol following field application. Overall, growth was reduced by 37.6% at the highest concentration (28% clove oil in seawater) relative to the control (0% clove oil). The reduction in growth was nearly as great (35.3% of the control) at half the concentration of clove oil (14%) when dissolved in ethanol. These results suggest the repeated use of clove oil (even without a solvent) can deleteriously affect corals.     

A simple technique for measuring buoyant weight increment of entire, transplanted coral colonies in the field

Estimating the impacts of global and local threats on coral reefs requires monitoring reef health and measuring coral growth and calcification rates at different time scales. This has traditionally been mostly performed in short-term experimental studies in which coral fragments were grown in the laboratory or in the field but measured ex situ. Practical techniques in which growth and measurements are performed over the long term in situ are rare. Apart from photographic approaches, weight increment measurements have also been applied. Past buoyant weight measurements under water involved a complicated and little-used apparatus. We introduce a new method that combines previous field and laboratory techniques to measure the buoyant weight of entire, transplanted corals under water. This method uses an electronic balance fitted into an acrylic glass underwater housing and placed atop of an acrylic glass cube. Within this cube, corals transplanted onto artificial bases can be attached to the balance and weighed at predetermined intervals while they continue growth in the field. We also provide a set of simple equations for the volume and weight determinations required to calculate net growth rates. The new technique is highly accurate: low error of weight determinations due to variation of coral density (< 0.08%) and low standard error (< 0.01%) for repeated measurements of the same corals. We outline a transplantation technique for properly preparing corals for such long-term in situ experiments and measurements.