Major bleaching events can lead to increased thermal tolerance in corals

Climate change is a major threat to coral reef ecosystems worldwide. A key determinant of the fate of reef corals in a warming climate is their capacity to tolerate increasing thermal stress. Here, an increase in thermal tolerance is demonstrated for three major coral genera (Acropora, Pocillopora and Porites) following the extensive mass bleaching event that occurred on the Great Barrier Reef (Australia) in 1998. During the subsequent and more severe thermal stress event in 2002, bleaching severity was 30–100% lower than predicted from the relationship between severity and thermal stress in 1998, despite higher solar irradiances during the 2002 thermal event. Coral genera most susceptible to thermal stress (Pocillopora and Acropora) showed the greatest increase in tolerance. Although bleaching was severe in 1998, whole-colony mortality was low at most study sites. Therefore, observed increases in thermal tolerance cannot be explained by selective mortality alone, suggesting a capacity for acclimatization or adaptation. Although the vulnerability of coral reefs remains largely dependent on the rate and extent of climate change, such increase in thermal tolerance may delay the onset of mass coral mortalities in time for the implementation of low-emission scenarios and effective management.     

In situ thermal dynamics of shallow water corals is affected by tidal patterns and irradiance

We studied the diel variation of in situ coral temperature, irradiance and photosynthetic performance of hemispherical colonies of Porites lobata and branching colonies of Porites cylindrica during different bulk water temperature and tidal scenarios on the shallow reef flat of Heron Island, Great Barrier Reef, Australia. Our study presents in situ evidence that coral tissue surface temperatures can exceed that of the surrounding water under environmental conditions typically occurring during low tide in shallow reef or lagoon environments. Such heating may be a regular occurrence on shallow reef flats, triggered by the combined effects of high irradiance and low water flow characteristic of low Spring tides. At these times, solar heating of corals coincides with times of maximum water temperature and high irradiance, where the slow flow and consequent thick boundary layers impede heat exchange between corals and the surrounding water. Despite similar light-absorbing properties, the heating effect was more pronounced for the hemispherical P. lobata than for the branching P. cylindrica. This is consistent with previous laboratory experiments showing the evidence of interspecific variation in coral thermal environment and may result from morphologically influenced variation in convective heat transfer and/or thermal properties of the skeleton. Maximum coral surface warming did not coincide with maximum irradiance, but with maximum water temperature, well into the low-tide period with extremely low water flow in the partially drained reef flat, just prior to flushing by the rising tide. The timing of low tide thus influences the thermal exposure and photophysiological performance of corals, and the timing of tidally driven coral surface warming could potentially have different physiological impacts in the morning or in the afternoon.     

Algal overgrowth of alcyonacean soft corals

Colonies of the soft coral Lobophytum pauciflorum (Ehrenberg, 1834) (Coelenterata: Octocorallia: Alcyonacea: Alcyoniidae), some of which were heavily overgrown by the algae Ceranium flaccidum (Kuetzing) Ardissone and Enteromorpha sp., and other minor epizoites, were collected at Taylor Reef (17°50S; 146°35E) in the Great Barrier Reef. Overgrown colonies contained the diterpene 2-epi-sarcophytoxide as the major secondary metabolite, while conspecific colonies with clean polyparies contained two diterpenes in approximately equal amounts: 14-hydroxycembra-1,3,7,11-tetraene and 15-hydroxycembra-1,3,7,11-tetraene. By contrast, twenty conspecific pairs of overgrown and clean colonies of other alcyoniid soft corals collected from Pelorus Channel, Palm Island Group (18°34S; 146°29E), showed no chemical differences within in the pairs. Cultures of a common species of Ceramium [C. codii (Richards) Mazoyer] were incubated with different concentrations of nine soft-coral-derived diterpenes and significant algal growth inhibition was observed in many cases. It appears that terpenoids from soft coral may contribute to the lack of epizoic organisms on soft-coral polyparies.     

Effects of eutrophication on reef-building corals

Fourteen environmental variables were monitored at seven locations along the west coast of Barbados on a weekly basis over a one-year period, 1981 to 1982. The physicochemical and biological data indicate that an environmental gradient exists as a result of increased eutrophication of coastal waters. Growth rates (linear extension) of Montastrea annularis (Ellis and Solander), measured along the environmental gradient, exhibit high correlation with a number of water quality variables. Concentration of suspended particulate matter is the best univariate estimator of M. annularis skeletal extension rates (r ²=0.79, P<0.0001). The results suggest that suspended particulate matter may be an energy source for reef corals, increasing growth up to a certain maximum concentration. After this, reduction of growth occurs due to smothering, reduced light levels and reduced zooxanthellae photosynthesis.     

Antimicrobial activity of Red Sea corals

Scleractinian corals and alcyonacean soft corals are the two most dominant groups of benthic marine organisms inhabiting the coral reefs of the Gulf of Eilat, northern Red Sea. Antimicrobial assays performed with extracts of six dominant Red Sea stony corals and six dominant soft corals against marine bacteria isolated from the seawater surrounding the corals revealed considerable variability in antimicrobial activity. The results demonstrated that, while the majority (83%) of Red Sea alcyonacean soft corals exhibited appreciable antimicrobial activity against marine bacteria isolated from the seawater surrounding the corals, the stony corals had little or no antimicrobial activity. From the active soft coral species examined, Xenia macrospiculata exhibited the highest and most potent antimicrobial activity. Bioassay-directed fractionation indicated that the antimicrobial activity was due to the presence of a range of compounds of different polarities. One of these antibiotic compounds was isolated and identified as desoxyhavannahine, with a minimum inhibitory concentration (MIC) of 48 μg ml⁻¹ against a marine bacterium. The results of the current study suggest that soft and hard corals have developed different means to combat potential microbial infections. Alcyonacean soft corals use chemical defense through the production of antibiotic compounds to combat microbial attack, whereas stony corals seem to rely on other means.     

Patterns of Oil-Sediment rejection in corals

The patterns of oil-sediment rejection of 19 Caribbean hermatypic corals are identical to theri patterns of rejection of clean sediments. The rejection pattern is typical for coral species, and displays maximum and minimum rates dependent on the size and density of the oil-sediment particles. The viscosity of the oil determines the size of the oil-sediment particles. A coral's efficiency of rejection of sediment depends on the size and amount of the sediment particles. Oil drops 0.06 mm are removed by the coral's tissues. Physical contact with oil-sediment particles appears to be less harmful to corals than the toxic effects of oils.     

Temperature and pressure tolerance of larvae of Crepidula fornicata suggest thermal limitation of bathymetric range

The extant deep-sea fauna is thought to result from recolonisation of this environment by shallow-water organisms following climate-driven mass extinctions. Planktonic larval tolerance to high pressure is considered an important preadaptation for successful deep-sea invasion. In this study, the pressure and temperature tolerance of a species without any known confamilial deep-sea relative were assessed for the first time. Early- and late-veliger larvae of the shallow-water species Crepidula fornicata were subjected to a temperature/hydrostatic pressure regime from 5 to 25 °C and from 0.1 to 40 MPa. Although early and late veliger survived pressures equivalent to 2,000 m water depth or greater at all temperatures, decreased larval activity indicated significant sublethal temperature and pressure effects. Reduced larval activity of early veliger at low temperatures suggests that the bathymetric range of this species may be thermally constrained. A mechanistic model is proposed to explain the emerging pattern of ontogenetic shifts in pressure tolerance of shallow-water benthic invertebrates.     

Antimicrobial activity of extracts of Caribbean gorgonian corals

Extracts of 39 species of Caribbean gorgonians were tested for antimicrobial activity against 15 strains of marine bacteria. The bacteria consisted of three opportunistic pathogens, Vibrio parahaemolyticus, Leucothrix mucor, and Aerococcus viridans, and 12 strains isolated from either healthy or decayed gorgonians. Overall, only 15% (79 out of 544) of the tests resulted in antibacterial activity with 33% (13 out of 39) of the gorgonians inhibiting only one bacterial strain and 23% (9 out of 39) showing no activity. The extracts of four Pseudopterogorgia species showed relatively high levels of activity, inhibiting 43 to 86% of the bacterial strains. The potency of the active Pseudopterogorgia species was variable, however, and three additional Pseudopterogorgia species were inactive against all bacterial strains. With the exception of one sensitive strain, Vibrio species were resistant to gorgonian metabolites. Our results indicate that organic extracts of most Caribbean gorgonians do not possess potent, broad-spectrum antibacterial activity inhibitory to the growth of opportunistic marine pathogens and bacteria associated with healthy and decayed gorgonian surfaces. These findings suggest that the inhibition of bacterial growth is not the primary ecological function of gorgonian secondary metabolites and that bacteria may not be important selective agents in the evolution of gorgonian secondary chemistry.     

Chemical resistance of gorgonian corals against fungal infections

The frequency and impact of diseases affecting corals throughout the Caribbean have been increasing but little is known about the factors promoting the emergence and outbreak of disease. A disease caused by a fungal pathogen [Aspergillus sydowii (Thom et Church)] which affects Caribbean sea fan corals provided an opportunity to examine the efficacy of coral crude extracts in disease resistance. Minimum inhibitory concentration (MIC) assays showed that of the 20 common gorgonian species in the Florida Keys, extracts from 15 species had MICs < 15 mg ml⁻¹ against A. sydowii pathogenic to sea fans. Extracts from several species in two gorgonian genera (Pseudoplexaura and Pseudopterogorgia) were among the most active, with MICs < 10 mg ml⁻¹. Gorgonia ventalina L., one of two sea fan species known to be hosts to A. sydowii in the field, had an MIC < 10 mg ml⁻¹, suggesting that complete disease resistance requires more active extracts. For the antifungal compounds to be effective in situ, they must also occur in sufficiently high concentrations in living coral tissue. For example, Pseudopterogorgia americana (Gmelin) had comparatively potent extracts but did not have sufficient concentrations in the tissue to be effective. Conversely, Plexaura homomalla Esper extracts were less potent but occurred in high enough concentrations in the tissue to be effective against A. sydowii. When potency and extract concentration are considered together (i.e. potency × concentration), several other gorgonian corals emerge as likely hosts to A. sydowii. Crude extracts from the most active gorgonian species were also effective against two geographic variants of A. sydowii pathogenic to sea fans, a non-pathogenic terrestrial strain of A. sydowii, and three strains of A. flavus Link known to be human, plant, and insect pathogens (MIC range, 7.5 to > 15 mg ml⁻¹). Although the potency in these assays did not attain a clinically significant level, the potency is comparable to a known antifungal agent, hygromycin B, which had an MIC ≤7.5 mg ml⁻¹ in our assays, highlighting the potential of these gorgonian corals for bioprospecting. Received: 29 May 1999 / Accepted: 22 May 2000     

Turnover of photosynthetically fixed carbon in reef corals

Siderastrea siderea and Montastrea annularis were labeled in situ with NaH¹⁴CO₃. The corals were sampled over a period of 11 days and the radioactivity remaining in the ethanolsoluble and ammonia-soluble fractions measured. Total radioactivity in the corals fell to about 1/3 after one night and then to about 1/3 in the next 10 days. The ethanol-soluble radioactivity is probably converted to the less soluble, ammonia-extractable, material in the dark.Contribution No. 1514 from the University of Miami, Rosenstiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

Stress-induced rapid release of antibacterials by scleractinian corals

Mechanical stress on the coral Pocillopora damicornis caused the release of material that killed the coral pathogen Vibrio coralliilyticus. The bactericidal material was released into the surrounding seawater rapidly, reaching a maximum killing activity within 1 min of the stress. The coral antibacterial activity, referred to as CAA, was retained following filter sterilization and storage at –20°C. Exposure of V. coralliilyticus to CAA for 30 s, 1 min and 6 min resulted in the death of 82%, 89% and 99% of the bacteria, respectively. Release of CAA following mechanical stress was also observed with four other coral species tested. P. damicornis CAA was bactericidal to a wide variety of Gram-negative and Gram-positive bacteria. This is the first report that hard corals rapidly release fast-acting bactericidal material following mechanical stress. The release of CAA was demonstrated with both aquarium corals and corals taken directly from the sea. It is suggested that CAA is part of hard corals host defense system against infection, the natural stimulation for release of CAAs being the bite of a predator. Previous failures to detect antibacterial activity in hard corals can be attributed to a lack of understanding of the sensitive mechanism by which they are released.Communicated by M. Kühl, Helsingør     

Effects of eutrophication and sedimentation on juvenile corals

Settlement of juvenile scleractinian corals was investigated from 1987 to 1990 on eutrophic and less eutrophic fringing reefs on the west coast of Barbados, West Indies. The number of coral recruits and number of recruiting coral species on cement blocks decreased with increasing eutrophication of the reefs. This may suggest lower settlement rates on eutrophic reefs, but could also liave resulted from higher post-settlement mortality, since blocks were examined only once after 3 yr of immersion. Coral settlement rates to artificial plates that were checked monthly were also lower on the more eutrophic reefs. This could result from lower local availability of larvae caused by fewer adult corals and/or lower reproductive rates of corals on eutrophic reefs. However, the ratio of coral recruits to adult coral abundance was considerably lower on eutrophic reefs, suggesting that local coral abundance alone can not explain lower settlement rates on eutrophic reefs. The lower rates on eutrophic reefs may result from a lower probability of coral larvae settlin when present, perhaps because of a limited availability of suitable settlement substrate. Colonization of settlement plates by non-coralline organisms was heavier on eutrophic reefs, and unoccupied space was lower, supporting the suggestion that suitable coral settlement substrate may be limiting on eutrophic reefs. Moreover, coralline algae, which facilitate metamorphosis and settlement of coral larvae, were less abundant on settlement plates on eutrophic reefs.     

Spatial arrangement of competitors influences coexistence of reef-building corals

Spatial aggregation among strong competitors has been identified as a putative mechanism promoting the coexistence of weak competitors in intensely competitive communities. With notable exceptions in plant communities, few investigators have tested this hypothesis experimentally. In this study, we manipulated the spatial arrangement of corals to test whether within-patch aggregation of a strong coral competitor enhances the success of a weaker coral competitor. Corals grown in simple aggregated arrangements, where the number and type of competitors were held constant, grew almost twice as much as those in non-aggregated arrangements. These growth results suggest that species coexistence is promoted by aggregation within competitive neighborhoods. Thus spatial aggregation may be one of several important mechanisms contributing to the persistence of weak competitors and species coexistence on coral reefs.     

Some aspects of the thermal tolerance of Trichomya hirsuta (Mollusca: Bivalvia) of Eastern Australia

The upper temperature tolerance range for Trichomya hirsuta L. is described for acclimation temperatures of 10° to 32°C at 30% S. Changes in thermal resistance with time are described along with the incipient lethal temperatures. The eurythermicity for the species is 240.7 C°-squared. The ultimate upper incipient lethal temperature is 35.1°C. Definition of the incipient sublethal temperature is given, and is equal to 32°C. Diet effects are shown not to be significant in short-term lethal experiments, but indicate that unfed mussels reduce their resistance after prolonged exposure to high temperatures. Size was also shown to have no effect on resistance and tolerance. A comparison of the tolerance triangle of T. hirsuta with those of the temperate species Mytilus edulis and Mya arenaria revealed that the 24-h LT₅₀ minus 2.2 C° approximates the incipient lethal temperature. The eurythermicities of the temperate species are Mytilus edulis, 363 C°-squared; and Mya arenaria, 415 C°-squared.     

Palatability of invertebrate larvae to corals and sea anemones

Risk of larval mortality is an underlying theme in debates and models concerning the ecology and evolution of the differing reproductive characteristics among marine benthic invertebrates. In these discussions, predation is often assumed to be a major source of larval mortality. Previous studies, focused primarily on planktotrophic larvae, suggested that marine larvae generally were susceptible to, and poorly defended against, planktivorous fishes and invertebrates. Larval-planktivore interactions involving larger and more conspicuous lecithotrophic larvae that are typical of many brooding sessile invertebrates have not been well studied. This lack of data for diverse larval types has hindered testing broad generalities about marine larvae and planktivore prey-preferences. This study demonstrates that lecithotrophic larvae of many Caribbean and temperate western Atlantic invertebrates are distasteful to co-occurring corals and anemones. These larval predators frequently rejected larvae of sponges (6 of 9 species), gorgonians (7 of 9 species), corals (3 of 3 species), hydroids (2 of 2 species) and a bryozoan. Larvae of three temperate colonial ascidians were readily consumed. Frequencies of survivorship for larvae captured but rejected by corals and anemones were generally high and, in 20 of 24 assays, were not statistically different from those of unattacked control larvae. Levels of metamorphosis (when it occurred) of rejected larvae also rarely differed significantly from those of unattacked controls. These results provide further evidence that larval palatability to predators may not be as high as once thought, particularly for brooded larvae of sessile colonial invertebrates. The means by which larvae may avoid or deter predators, and the demographic consequences for marine invertebrates and for the evolution of invertebrate life-history patterns, need to be assessed.     

Effects of sand deposition on scleractinian and alcyonacean corals

The ability of corals to withstand experimental sand deposition was investigated for two experimental periods (17 h and 6 wk) in eight scleractinia (Favia favus, Favites pentagona, Platygyra daedalea, Gyrosmilia interrupta, Galaxea fascicularis, Cyphastrea chalcidicum, Favites abdita, Goniopora dijboutensis) and five alcyonacea (Lobophytum depressum, L. venustum, Sinularia dura, S. leptoclados, Sarcophyton glaucum) collected in 1992 from Natal, South Africa. Scleractinia were active sediment shedders, alcyonacea passive, relying on water motion and gravity. Short-term sand clearing efficiency was primarily dependent on corallum shape. Sand application led to hydrostatic inflation of polyps in scleractinia and the entire colony in alcyonacea as well as to increased tentacular action in the scleractinian Gyrosmilia interrupta. Under continuous sand application, inflation remained while other activities, such as tentacular motion, ceased completely. In scleractinia and alcyonacea, tissue necroses appeared after the first week of continuous sand application. Death of entire colonies and partial bleaching of continually sandcovered areas were observed in alcyonacea only. Different grain sizes of sand had no influence on clearing reaction or efficiencies.     

Sediment-rejection efficiency of 22 species of Australian scleractinian corals

Sediment-rejection rates of 22 Australian midshelf coral species were studied in situ between March and July 1988 at Lizard Island, Great Barrier Reef. Rejection rates of non-branching species were positively correlated with calice size and were faster for fine (63 to 250 m) than for coarse (500 to 1000 m) sediment at influxes of 200 mg/cm². Increasing water turbulence was a more important influence on rejection rates for some species than for others. Most replicates of most species cleared all sediment in 2 d. Of those that did not, some Favia stelligera and Leptoria phrygia showed partial tissue death within 48 h, and Gardineroseris planulata showed partial tissue death within 6 d. Montipora aequituberculata, Porites lobata and P. lutea tolerated sediment for at least 6 d, and exhibited extensive tissue bleaching, but these tissues recovered after sediment removal. Sediment-rejection efficiency and sediment tolerance are not directly related, and wide interspecific differences may occur in both. These findings are correlated with differences in sediment-rejection mechanisms and with ecological distributions on the reef.     

Experimental tests of suspension feeding in Atlantic reef corals

The ability of 15 species of Atlantic reef corals to act as suspension feeders was demonstrated by their removal of suspended particles from sea water in culture vessels. Mean clearance rates varied from 16.6 to 145.5 ml water cleared/h/cm² of live coral tissue. The lowest rates was found in Porites porites which is primarily a tentacle feeder, and the highest in Diploria clivosa which acts as both a tentacle feeder and suspension feeder. Rates of particle clearance in Agaricia agaricites, which is primarily a suspension feeder, were influenced by current velocity and type of food.