Spatial autocorrelation analysis of small-scale genetic structure in a clonal soft coral with limited larval dispersal

The philopatric larval dispesal and small effective population sizes characteristic of many clonal species should promote the development of significant small-scale genetic structure within populations as a result of isolation-by-distance. We used spatial autocorrelation statistics to detect genetic structure, arising from both clonal reproduction and philopatric dispersal of sexual propagules, for five allozyme loci within populations of the soft coral Alcyonium sp. In a population on Tatoosh Island, Washington, USA, sampled in 1991/1992, we found significant positive spatial autocorrelation at all loci among individuals separated by <40 cm, reflecting the presence of significant smallscale genetic structure due to associations among clonemates. For 4 of 5 loci, however, we detected no significant spatial autocorrelation among the different clones within this population over distances of 1 to 40 m. Analysis of soft-coral populations from six additional, topographically diverse sites in the north-east Pacific also did not reveal significant spatial autocorrelation among clones at any loci. This general lack of spatial autocorrelation of genotypes among clones suggests that significant small-scale genetic structure has not arisen in populations of Alcyonium sp. as a consequence of isolation-by-distance.     

Structural investigations on the mucus from six species of coral

The chemical composition of the mucus from three hard corals (Acropora formosa, Pachyseris speciosa and Fungia fungites) and three soft corals (Sarcophyton sp., Lemnalia sp., and Cespitularia sp.) collected on the Great Barrier Reef (1982–1985) was determined. Significant variation exists in the composition and structure of the six mucus samples, indicating the absence of a common structure for coral mucus. In all cases protein and/or carbohydrate polymers are the major components of the mucus, and lipids are present only in small amounts. The glycose composition varied between species, with fucose (F. fungites and Lemnalia sp.), arabinose (A. formosa), galactose (P. speciosa) and N-acetyl glucosamine (Sarcophyton sp.) being present in high concentrations. With the exception of Sarcophyton sp. and Lemnalia sp., all mucus samples were acidic and contained significant sulphate but no uronic or sialic acids. The amino acid composition of the mucus samples was not unusual, apart from A. formosa, which contained a high percentage of serine and threonine, and F. fungites, which had high levels of glutamic acid present.     

Effects of selective settlement and of aggression by residents on distribution of young recruits of two tropical damselfishes

The reef fishes that settled on an array of experimental corals at Lizard Island, Queensland, Australia, were counted during a pulse of recruitment in December 1986. NeitherPomacentrus sp. norP. amboinensis showed any evidence that harassment by residentDascyllus aruanus caused a decrease in persistence during the first day after settlement. LarvalPomacentrus sp. settled selectively on corals without residentD. aruanus. The results forP. amboinensis were ambiguous. Settlers of both species positioned themselves closer to the sand on corals with residentD. aruanus than on unoccupied corals. This could reduce access to planktonic food and increase the risk of predation. Adult aggression may be less important and active selection of settlement sites by larvae may be more important to the distribution of recruits than is suggested by the literature. The presence or absence of particular species should be included among the cues that larval reef fishes use to choose settlement sites.     

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.     

Dynamics of a coral reef community after mass mortality of branching <Emphasis Type="Italic">Acropora</Emphasis> corals and an outbreak of anemones

Dynamics of a coral reef community at Tiao-Shi Reef, southern Taiwan were studied using permanent transects to examine coral recovery and successive cascades to collapse stage resulting from chronic anthropogenic impacts and typhoons. Three distinct zones were recognized within a relatively small study area (250 m across) formerly dominated by large stands of branching Acropora corals. The first zone still retains the dominance of branching Acropora corals, although they show a significant decreasing tendency. The second zone exhibits recovery with a significant increase in branching Montipora stellata, which is recruited and grows faster than branching Acropora corals. The third zone is occupied by anemone, Condylactis sp., and demonstrates a stable phase of coral deterioration without recovery. Such differences in coral reef community dynamics within a small spatial scale illustrate mosaic dynamics which have resulted from degradation of the water quality, patchy mortality of large branching Acropora thickets caused by typhoons, the rapid asexual fragmentation and growth of M. stellata making it a successful colonizer, and occupation by anemone, Condylactis sp., together with unstable remnants of dead Acropora rubbles have not allowed coral recruits to survive.     

Association of <Emphasis Type="Italic">Waminoa</Emphasis> sp. (Acoela) with corals in the Wakatobi Marine Park,South-East Sulawesi,Indonesia

This is the first quantitative study on the prevalence of epizoic Waminoa sp. acoel worms and their association with corals in the Wakatobi Marine National Park (WMNP), South-East Sulawesi, Indonesia. Three replicate transects were laid on the reef crest, flat and slope at six sites in 2006 and eight sites in 2007. Four of the sites were common in both years. In total 69 transects were surveyed in 2006, and 87 transects in 2007. A total of 4.8% of all observed hard corals were associated with acoel worms in 2006 and 2.6% of hard and soft corals in 2007. Acoels were present on 16 and 21 of the coral taxa studied in 2006 and 2007 respectively. The worms were strongly associated with the azooxanthellate coral Tubastrea spp. and were rare or absent on the most abundant coral genera Montipora and Porites. The mean number of corals having acoels was highest on reef slopes, whereas acoels were virtually absent on reef flats. Corals that had a high and a medium cover of worms were more common in 2007 than in 2006. No significant trend in the adaptation of the zooxanthellae of Waminoa sp. to different depths at different sites was revealed. The impact of the worm on the coral is unknown, but high numbers may have a shading effect and a negative impact on the coral’s photophysiology. This acoel merits more study of its life cycle, its photophysiology, and its impact on its host corals. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Light harvesting in the green alga Ostreobium sp., a coral symbiont adapted to extreme shade

Ostreobium sp. (Chlorophyta: Siphonales) can be found as green bands within the skeletal material of a number of stony corals in the Indo-Pacific and Caribbean regions. Many of these corals also contain symbiotic dinoflagellates in the overlaying coral polyps that effectively screen out all the typical photosynthetically active radiation from the algae in the green bands below. Ostreobium sp., nevertheless, grows photosynthetically. Its action spectrum and absorption spectrum have been shown to extend much further into the near infra-red compared to other green algae. In the present study, carried out in 1987, fluorescence excitation and emission spectra were measured in Ostreobium sp. and compared to spectra obtained from the green alga Ulva sp. and the brown alga Endarachne sp. Xanthophylls, probably siphonein and an unidentified xanthophyll probably related to siphonaxanthin, are photosynthetically active in Ostreobium sp., and can sensitize Photosystem II fluorescence at 688 nm and Photosystem I (PS I) fluorescence at 718 nm. The fluorescence emission spectra of Ostreobium sp. measured at 25° C and 77 K were not remarkably different from those of the green alga Ulva sp. Absorbance changes induced by light were measured in Ostreobium sp. from 670 to 750 nm and were like those normally seen in green plants except that, in addition to the minimum expected for the reaction-center chlorophyll of PS I (P700) at 703 nm, another minimum was seen at 730 nm. It is possible that this spectrumreflects the functioning of a reaction center of Photosystem I that has adapted to function in light highly enriched in far-red wavelengths.CIW-DPB Publication No. 1021     

Three party symbiosis: acoelomorph worms,corals and unicellular algal symbionts in Eilat (Red Sea)

Epizoic worms were found to occur on certain coral colonies from reefs off the coast of Eilat (Red Sea). We identified 14 coral species infested by acoelomorph worms at a depth range of 2–50 m. The host corals were all zooxanthellate and included both massive and branching stony corals and a soft coral. Worms from all hosts were identified as belonging to the genus Waminoa and contained two distinct algal symbionts differing in size. The smaller one was identified as Symbiodinium sp. and the larger one is presumed to belong to the genus Amphidinium. Worm-infested colonies of the soft coral, Stereonephthya cundabiluensis, lacked a mucus layer and exhibited distinct cell microvilli, a phenotype not present in colonies lacking Waminoa sp. In most cases, both cnidarian and Acoelomorph hosts displayed high specificity for genetically distinctive Symbiodinium spp. These observations show that the epizoic worms do not acquire their symbionts from the “host” coral.     

Gas-liquid chromatograms of sesquiterpenes as finger prints for soft-coral identification

A possible additional means for aiding in the identification of soft corals based on their sesquiterpene composition, as determined by gas-liquid chromatography (GLC), is discussed. The use of this method for several species of Sinularia and Sarcophyton is illustrated. Several sesquiterpenes were identified, some of them for the first time from marine origin. Preliminary tests indicate that the sesquiterpene composition in the tested soft corals remained quite constant during different seasons of the year. It is suggested that such finger prints are produced by the corals themselves and not by the zooxanthellae, and that they are species-specific.     

Onset of symbiosis and distribution patterns of symbiotic dinoflagellates in the larvae of scleractinian corals

The establishment of symbiosis in early developmental stages is important for reef-building corals because of the need for photosynthetically derived nutrition. Corals spawn eggs and sperm, or brood planula larvae and shed them into the water. Some coral eggs or planulae directly inherit symbiotic dinoflagellates (Symbiodinium spp.) from their parents, while others acquire them at each generation. In most species examined to date, the larvae without dinoflagellates (aposymbiotic larvae) can acquire symbionts during the larval stage, but little is known regarding the timing and detailed process of the onset of symbiosis. We examined larval uptake of symbiotic dinoflagellates in nine species of scleractinian corals, the onset of symbiosis through the early larval stages, and the distribution pattern of symbionts within the larval host, while living and with histology, of two acroporid corals under laboratory conditions. The larvae acquired symbiotic dinoflagellates during the planktonic phase in all corals examined which included Acropora digitifera, A. florida, A. intermedia, A. tenuis, Isopora palifera, Favia pallida, F. lizardensis, Pseudosiderastrea tayamai, and Ctenactis echinata. The larvae of A. digitifera and A. tenuis first acquired symbionts 6 and 5 days after fertilization, respectively. In A. digitifera larvae, this coincided with the formation of an oral pore and coelenteron. The number of symbiotic dinoflagellates increased over the experimental periods in both species. To test the hypothesis that nutrients promotes symbiotic uptake, the number of incorporated dinoflagellates was compared in the presence and absence of homogenized Artemia sp. A likelihood ratio test assuming a log-linear model indicated that Artemia sp. had a significantly positive effect on symbiont acquisition. These results suggest that the acquisition of symbiotic dinoflagellates during larval stages is in common with many coral species, and that the development of both a mouth and coelenteron play important roles in symbiont acquisition.     

Population structure and feeding deterrence in three shallow-water antarctic soft corals

Alcyonium paessleri and Clavularia frankliniana are numerically abundant soft corals in the nearshore (12 to 33 m depth) benthic communities of eastern McMurdo Sound. They are much less abundant in western McMurdo Sound where a third species, Gersemia antarctica, co-occurs in low numbers. The body tissues of these three species are comprised mainly of organic material (53 to 70% dry wt), which is primarily dervied from NaOH-soluble protein and refractory material. The energetic contents of the whole-body tissues of A. paessleri, C. frankliniana and G. antarctica are 15.9, 17.3, and 14.5 kJ g^-1 dry wt, respectively. The mean biomass per individual is 1.81, 0.008, and 45 g dry wt for each respective species. Based on population densities of 7.3, 1337.3, and 0.04 soft corals m^-2 for A. paessleri, C. frankliniana and G. antarctica, respectively, the population energetic densities are estimated to be 210.1, 185.1, and 26.1 kJ m^-2. Despite the relatively rich energetic content of the tissue and apparent vulnerability to predators, very little predation occurs on these soft corals. Two potential predators, the antarctic sea stars Perknaster fuscus and Odontaster validus, exhibited significant chemotactic defensive tube-foot retractions to hexane, chloroform, methanol, and aqueous methanol extracts of each soft coral. In addition, wholebody tissue of each soft coral was rejected by the demersal fish Pseudotrematomus bernacchii and the cryopelagic fish Pagothenia borchgrevinki. In contrast, whole soft-coral tissues sequentially extracted in four increasingly polar solvents were readily ingested by these antarctic fishes, indicating that sclerites do not play a significant role in deterring predators. Our results indicate that these antarctic soft corals contain bioactive compounds which deter common predatory seastars and fishes.     

Habitat patchiness and predation modify the distribution of a coral-dwelling damselfish

Fish abundance is often better predicted by microhabitat variables on continuous reefs than on isolated patch reefs. Although this was suggested to stem from reduced post-recruitment relocation, this has not been shown experimentally. We found the relationship between the presence of a coral-dwelling fish, Dascyllus marginatus, and the size of its coral host to differ between corals on continuous reefs and the sparsely distributed corals on sandy bottoms. Empty transplanted corals were colonized exclusively by new recruits when on the sandy bottom, and both by new recruits and post-recruitment dispersal of adults when on the continuous reef. New recruits settled predominantly into small corals, although analyses of recruitment patterns were confounded by low recruitment in the studied years. Both tank experiments and field survey data suggest that the presence of recruits in small corals is at least partially driven by predation by the dottyback, Pseudochromis olivaceus, which lives predominantly in large corals within both habitats. Consequently, we suggest that the relationship between fish presence and coral size differs between the habitats due to coral size dependent predation on recruits and variability in the importance of direct recruitment to replenish fish populations.     

Stimulation of fat-body production in the polyps of the coralPocillopora damicornis by the presence of mutualistic crabs of the genusTrapezia

A mutualism exists between the xanthid crabs of the genusTrapezia and their host corals,Pocillopora damicornis. It has previously been established that these obligate coral residents benefit the coral hosts by defending them against echinoderm predators and by increasing the survival of polyps located deep between the coral branches. In turn, the corals apparently benefit the crabs by producing lipid-filled structures on which the trapezid crabs feed; these fat bodies may contain some of the lipid which in previous studies of coral metabolism has been termed excess. It was determined by experiments conducted at the Hawaii Institute of Marine Biology that the presence of crabs in colonies ofP. damicornis stimulates the polyps to produce the lipid-filled fat bodies; removal of crabs causes corals to cease producing fat bodies. A structure very similar to the fat bodies ofP. damicornis has been reported inAcropora durvillei. Both of these coral genera ordinarily possess xanthid-crab mutualists. This association between branching corals and crustaceans may have evolved because corals of these genera provide shelter among their branches and because these shallow-water corals are evidently capable of releasing lipid which is excess to the corals' metabolic needs, but which can be utilized by the crabs.     

Positive and Negative Effects of a Threatened Parrotfish on Reef Ecosystems

Species that are strong interactors play disproportionately important roles in the dynamics of natural ecosystems. It has been proposed that their presence is necessary for positively shaping the structure and functioning of ecosystems. We evaluated this hypothesis using the case of the world's largest parrotfish (Bolbometopon muricatum), a globally imperiled species. We used direct observation, animal tracking, and computer simulations to examine the diverse routes through which B. muricatum affects the diversity, dispersal, relative abundance, and survival of the corals that comprise the foundation of reef ecosystems. Our results suggest that this species can influence reef building corals in both positive and negative ways. Field observation and simulation outputs indicated that B. muricatum reduced the abundance of macroalgae that can outcompete corals, but they also feed directly on corals, decreasing coral abundance, diversity, and colony size. B. muricatum appeared to facilitate coral advancement by mechanically dispersing coral fragments and opening up bare space for coral settlement, but they also damaged adult corals and remobilized a large volume of potentially stressful carbonate sediment. The impacts this species has on reefs appears to be regulated in part by its abundance—the effects of B. muricatum were more intense in simulation scenarios populated with high densities of these fish. Observations conducted in regions with high and low predator (e.g., sharks) abundance generated results that are consistent with the hypothesis that these predators of B. muricatum may play a role in governing their abundance; thus, predation may modulate the intensity of the effects they have on reef dynamics. Overall our results illustrate that functionally unique and threatened species may not have universally positive impacts on ecosystems and that it may be necessary for environmental managers to consider the diverse effects of such species and the forces that mediate the strength of their influence. Efectos Positivos y Negativos de un Pez Loro Amenazado Sobre Ecosistemas Arrecifales     

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 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.     

Elevated temperature impairs onset of symbiosis and reduces survivorship in larvae of the Hawaiian coral, <Emphasis Type="Italic">Fungia scutaria</Emphasis>

Many corals obtain their obligate intracellular dinoflagellate symbionts from the environment as larvae or juveniles. The process of symbiont acquisition remains largely unexplored, especially under stress. This study addressed both the ability of Fungia scutaria (Lamarck 1801) larvae to establish symbiosis with Symbiodinium sp. C1f while exposed to elevated temperature and the survivorship of aposymbiotic and newly symbiotic larvae under these conditions. Larvae were exposed to 27, 29, or 31°C for 1 h prior to infection, throughout a 3-h infection period, and up to 72 h following infection. Exposure to elevated temperatures impaired the ability of coral larvae to establish symbiosis and reduced larval survivorship. At 31°C, the presence of symbionts further reduced larval survivorship. As sea surface temperatures rise, coral larvae exposed to elevated temperatures during symbiosis onset will likely be negatively impacted, which in turn could affect the establishment of future generations of corals.     

Uptake of dissolved inorganic nitrogen by the symbiotic clam Tridacna gigas and the coral Acropora sp.

Dissolved inorganic nitrogen flux was studied in the giant clam Tridacna gigas and the corals Acropora sp. and Tubastrea micrantha from the tropical reefs of Belau, Micronesia in 1983. T. micrantha, a nonsymbiotic coral, excreted ammonium. However, Tridacna gigas and Acropora sp., which contain symbiotic dinoflagellates (zooxanthellae) were able to take up both ammonium and nitrate. The requirement for a previous light exposure to sustain uptake by T. gigas is reported. The uptake kinetics of these symbioses are described and include the capacity of the zooxanthellae for surge uptake when given nutrient spikes.Contribution No. 417 of the Allan Hancock Foundation     

Marginal tentacles of the corallimorpharian Rhodactis rhodostoma. 1. Role in competition for space

The corallimorpharian Rhodactis rhodostoma (Ehrenberg, 1934) forms aggregations that dominate patches on some coral reef flats in the Red Sea. The outcomes and mechanisms of competition for space between this corallimorpharian and other sessile organisms are poorly understood. Polyps of R. rhodostoma were observed to overgrow zoanthids, hydrozoan corals, sponges and encrusting macroalgae on a fringing reef at Eilat, northern Red Sea. R. rhodostoma polyps also damaged, and in some cases overgrew, reef-building corals in the families Poritidae, Acroporidae and Pocilloporidae, most of which form branching colonies with small polyps that are subordinate in coral competitive hierarchies. In contrast, most stony corals in the families Faviidae and Mussidae had standoff interactions with R. rhodostoma, in which they prevented the corallimorpharians from damaging them or approaching closer than 1 to 3 cm. The latter corals are ranked at the top of competitive hierarchies for Indo-Pacific corals, and they form massive colonies of large polyps which may develop aggressive organs termed sweeper tentacles. Some soft corals that exude allelopathic chemicals also avoided overgrowth by the corallimorpharians. Tentacles along the oral disk margin of R. rhodostoma polyps were swollen and bulbous during contacts with cnidarians. These bulbous marginal tentacles had significantly thicker ectoderm and a higher proportion of holotrichous nematocysts than did the normally filiform marginal tentacles of R. rhodostoma polyps. It is concluded that, on the reef flat at Eilat, this corallimorpharian damages and overgrows a variety of sessile competitors, including branching stony corals, via the application of specialised marginal tentacles filled with penetrating nematocysts. R. rhodostoma is an intermediate competitor in the aggressive hierarchy among Indo-Pacific Anthozoa, including the reef-building corals. Received: 1 July 1998 / Accepted: 24 March 1999