Competition between scleractinian corals and macroalgae: An experimental investigation of coral growth, survival and reproduction

Macroalgae are a major component of many coral reef flat communities, and are potentially major competitors with corals. The influence of macroalgae on several demographic parameters of four species of scleractinian coral by means of an algal clearance experiment was examined to determine specifically if macroalgae are affecting coral cover, growth, fecundity, fission, survivorship and recruitment. Also investigated were patterns of natural encounters between corals and algae.

Algal cover at the study site ranged from 41 to 56%, and coral cover from 8 to 10%. In total, 92 ± 4 ( )% of coral colonies were in contact with one or more species of macroalgae. Changes in coral cover were significantly affected by the presence of macroalgae, with cover of Acropora species increasing faster in areas from which algae had been cleared compared to control areas where algae had not been removed, although this pattern did not occur for Pocillopora damicornis (Linnaeus). Similarly, growth of individual colonies was faster when macroalgae were absent for three Acropora species but not for P. damicornis. There were no differences detected in rates of fission or survivorship of corals between algal clearance and control treatments, although there were high levels of variability in both of these parameters. Fecundity of Acropora palifera (Lamarck), the only species examined, was approximately double in colonies in cleared plots compared to those in control plots with macroalgae present. As no recruitment occurred throughout the 2-yr study, it remains to be determined how macroalgae effect the settlement of coral larvae. The results show that macroalgae can have a major influence on the demography of scleractinian corals.     

Restoration of coral populations in light of genetic diversity estimates

Due to the importance of preserving the genetic integrity of populations, strategies to restore damaged coral reefs should attempt to retain the allelic diversity of the disturbed population; however, genetic diversity estimates are not available for most coral populations. To provide a generalized estimate of genetic diversity (in terms of allelic richness) of scleractinian coral populations, the literature was surveyed for studies describing the genetic structure of coral populations using microsatellites. The mean number of alleles per locus across 72 surveyed scleractinian coral populations was 8.27 (±0.75 SE). In addition, population genetic datasets from four species (Acropora palmata, Montastraea cavernosa, Montastraea faveolata and Pocillopora damicornis) were analyzed to assess the minimum number of donor colonies required to retain specific proportions of the genetic diversity of the population. Rarefaction analysis of the population genetic datasets indicated that using 10 donor colonies randomly sampled from the original population would retain >50% of the allelic diversity, while 35 colonies would retain >90% of the original diversity. In general, scleractinian coral populations are genetically diverse and restoration methods utilizing few clonal genotypes to re-populate a reef will diminish the genetic integrity of the population. Coral restoration strategies using 10–35 randomly selected local donor colonies will retain at least 50–90% of the genetic diversity of the original population. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Morphological variation in the polyps of the scleractinian coral Favia speciosa (Dana) around Singapore

A photographic technique was used to examine morphological differences in the living polyps of Favia speciosa sampled from three sites around Singapore. Eight characters were measured, seven of which differed significantly between the three study sites. Sedimentation rates and character size were much higher at the site closest to the mainland than at the two sites further from shore. Land reclamation and dredging contribute to high sediment rates in Singapore waters; these rates decrease with increasing distance from shore. Large polyps close to the main island of Singapore are possibly a plastic, or selected for, response to high levels of sediment.     

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.     

Microsatellite loci isolated from the scleractinian coral, Acropora nobilis

We report the isolation and characterization of eight microsatellite loci from the scleractinian coral, Acropora nobilis. The microsatellite loci were obtained using compound SSR primers or an enrichment protocol. All the loci were polymorphic with four to eight alleles per locus and observed heterozygosities ranging from 0.22 to 0.76. Some of the primers developed for the two congeners, Acropora palmata and Acropora millepora were applicable to A. nobilis. These loci are useful for studying the connectivity among A. nobilis populations in Okinawa, southern Japan.     

Effects of host nutritional status and seasonality on the nitrogen status of zooxanthellae in the temperate coral Plesiastrea versipora (Lamarck)

The nitrogen status of endosymbiotic dinoflagellates (zooxanthellae) in the temperate coral Plesiastrea versipora (Lamarck) was determined by measuring the extent to which ammonium (40 μM NH₄⁺) enhanced the rate of zooxanthellar dark carbon fixation above that seen in filtered seawater (FSW) alone; the enhancement ratio was expressed as [dark NH₄⁺ rate/dark FSW rate]. V_D′/V_L, a further index of nitrogen status, was also calculated where V_D′ = [dark NH₄⁺ rate − dark FSW rate] and V_L = rate of carbon fixation in the light. When corals were starved for 2-8 weeks, zooxanthellar nitrogen deficiency became apparent at ≥ 4 weeks, with NH₄⁺/FSW and V_D′/V_L averaging up to 2.08 and 0.0061, respectively. A decrease in light-saturated photosynthesis per zooxanthella also occurred, with the photosynthetic rate after 4-6 weeks being just 81% of that seen prior to starvation. In comparison, when corals were fed 5 times per week for 8 weeks the addition of ammonium had little effect, indicating nitrogen sufficiency; NH₄⁺/FSW and V_D′/V_L were 1.03 and 0.0003, respectively. Photosynthetic rates of zooxanthellae from well-fed corals were up to 1.7 times greater than those of zooxanthellae from starved corals. The nitrogen status of zooxanthellae from corals in the field exhibited seasonal differences. Autumn samples were nitrogen sufficient, with NH₄⁺/FSW = 1.003 and V_D′/V_L = 0.0001. In contrast, a small degree of nitrogen deficiency was seen in winter and spring, when NH₄⁺/FSW averaged 1.075 and 1.249, and V_D′/V_L averaged 0.0013 and 0.0014, respectively. The greatest degree of nitrogen deficiency was observed in summer, when NH₄⁺/FSW averaged 1.318 and V_D′/V_L averaged 0.0036. Given the clear links between food supply and nitrogen status seen under experimental conditions, and the likelihood that the zooxanthellae are also able to take up nutrients directly from the seawater, the fluctuations in nitrogen status may reflect temporal fluctuations in seawater nutrient concentrations and plankton abundance. The nutrient status of these temperate zooxanthellae in the field is in contrast to the marked nitrogen deficiency seen in zooxanthellae from nutrient-poor coral reef waters, and raises the possibility that temperate zooxanthellae can store nitrogen for use when exogenous nutrients and food are less readily available. This, in turn, may contribute to the considerable stability of temperate zooxanthellar populations under highly variable environmental conditions.     

A poecilostome copepod parasitic in a scleractinian coral in the moluccas

Wedanus inconstans, n. gen., n. sp., a lichomolgid copepod living in the polyps of the reef coral Goniopora tenuidens (Quelch), is described from Halmahera in the Moluccas. The genus is apparently related to Xenomolgus Humes & Stock, 1972.     

Impacts of bleaching on the soft coral Lobophytum compactum. I. Fecundity, fertilization and offspring viability

We document long-term effects of a simulated bleaching event on the reproductive output and offspring viability of the soft coral Lobophytum compactum. Corals were subjected to temperature and solar radiation treatments to produce both moderately (48–60%) and heavily (90–95%) bleached colonies. Although bleached colonies recovered their zooxanthellae within 10 to 18 weeks, impacts on reproductive output were significant for at least two annual spawning seasons. In the first year, both polyp fecundity and mean oocyte diameter were reduced and inversely correlated with the degree of bleaching, with complete failure of fertilization in the group of heavily bleached colonies. For moderately bleached soft corals, survival and growth of sexual offspring did not differ significantly from those of unbleached colonies. Although no further reductions in zooxanthellae densities in experimental soft corals were recorded throughout the subsequent second year, egg size and fecundity of the heavily bleached soft corals were still significantly reduced 20 months later. Severe bleaching clearly has long-term sub-lethal impacts, reducing overall reproductive output for at least two spawning seasons. Accepted: 1 June 2000     

Impacts of bleaching on the soft coral Lobophytum compactum. II. Biochemical changes in adults and their eggs

Experimental bleaching reduces the levels of important biochemical parameters in adult tissues and eggs of the soft coral Lobophytum compactum. Protein, lipid, mycosporine-like amino acids (MAAs) and carotenoid concentrations remained lower in bleached adults than in controls for at least 8 months. Reductions in concentrations of all four parameters were greater in eggs than in maternal tissues, potentially jeopardizing egg and larval viability. In particular, reductions in lipids, proteins and carotenoids in tissues of heavily bleached soft corals were amplified approximately twofold in eggs. In comparison, amplification of maternal tissue reductions were not as great for MAAs, suggesting that MAAs are given higher priority in egg provisioning. Our finding that MAA levels are normally three times higher in eggs than in unbleached maternal tissues supports the importance of MAAs for larval survival. Twenty months after experimental bleaching the biochemical composition of both adult tissues and their eggs were indistinguishable from those of control (unbleached) soft corals. Accepted: 1 June 2000     

Genetic variation and gene flow of broadcast spawning and planula brooding coral,Goniastrea aspera (Scleractinia) in the Ryukyu Archipelago,southern Japan

The scleractinian coral Goniastrea aspera (Verrill) undergoes both broadcast spawning and planulae brooding in the Ryukyu Archipelago of southern Japan. Genetic variation and gene flow in G. aspera were studied using allozyme electrophoresis. We tested the hypothesis that gene flow is determined by the competency period of the planulae. We also assessed the relative contributions of sexual and asexual reproduction to recruitment. For the five staining systems surveyed, G. aspera encoded five polymorphic loci and one monomorphic locus. The genotype frequencies in each population significantly differed from the expected Hardy-Weinberg equilibrium (HWE), indicating that the local populations of G. aspera are not fully panmictic. The high ratio of the observed number of genotypes to the number of individuals (0.90 +/- 0.07, mean NG:N +/- SD) and the observed to expected genotypic diversity (0.84 +/- 0.11, mean GO:GE +/- SD) suggested that each population is likely maintained by sexual reproduction. The genetic differentiation (FST) and value of average number of migrants per generation (Nem) among and within regions ranged from 0.025 to 0.104 and 2.2 to 9.6, respectively. Comparisons with other species demonstrated that larva survival rates also influence gene flow. In addition, gene flow on distant reefs by planulae originating from spawning might prevent divergence by planulae originating from brooding for short-distant dispersal among and within populations of G. aspera in the Ryukyu Archipelago.     

Nematodes from a fresh-water pool on a coral island in the Solomon Islands

The present paper deals with the nematode species collected from a small deep fresh-water pool on a coral island in the Solomon Islands. 50% of the specimens recorded belong to terrestrial forms (4 species), 45% were marine forms (6 species) and only 5% belonged to fresh-water (4 species). Three species are described as new to science.     

Small-scale variability in the size structure of scleractinian corals around Moorea, French Polynesia: patterns across depths and locations

The size structure of coral populations is influenced by biotic and physical factors, as well as species-specific demographic rates (recruitment, colony growth, mortality). Coral reefs surrounding Moorea Island are characterized by strong environmental gradients at small spatial scales, and therefore, we expected that the size structure of coral populations would vary greatly at this scale. This study aimed at determining the degree of spatial heterogeneity in the population size structure of two coral taxa, Pocillopora meandrina and massive Porites spp., among depths (6, 12, and 18 m) and among locations (Vaipahu, Tiahura and Haapiti) representing different exposure to hydrodynamic forces. Our results clearly underlined the strong heterogeneity in the size structure of both P. meandrina and massive Porites spp., with marked variation among depths and among locations. However, the lack of any consistent and regular trends in the size structure along depths or among locations, and the lack of correlation between size structure and mean recruitment rates may suggest that other factors (e.g., stochastic life history processes, biotic interactions, and disturbances) further modify the structure of coral populations. We found that the size structure of P. meandrina was fundamentally different to that of massive Porites spp., reflecting the importance of life history characteristics in population dynamics. Handling editor: I. Nagelkerken     

Genetic evidence supporting the existence of two distinct species in the genusGasterosteus around Japan

Synopsis The genetic and morphological features ofGasterosteus aculeatus were investigated for 29 populations around Japan. Allozyme analyses recognized two groups (Pacific Ocean group and Japan Sea group) that had distinct characteristic features, and showed high genetic differentiation between them (D = 0.482). The Pacific Ocean group had a wide range, from North America to Japan, along the Pacific coast. The distribution of the Japan Sea group was limited around the Sea of Japan and the Sea of Okhotsk. The distribution of these groups were found to be sympatric on the Pacific coast of Hokkaido Island, Japan. From this area, genetic analyses demonstrated that the sympatric populations of the two groups formed independent breeding stocks, and it is considered that the two groups were reproductively isolated from each other. Additionally, each group had distinctive morphological features of lateral plates and caudal keels in the sympatric area. These results suggested that these two groups of the threespine stickleback comprise different species and that the Japan Sea group is taxonomically distinguishable fromG. aculeatus.     

Genetic structure of Heliofungia actiniformis (Scleractinia: Fungiidae) populations in the Indo-Malay Archipelago: implications for live coral trade management efforts

The fungiid Heliofungia actiniformis is one of the most popular scleractinian coral species in the growing live aquarium trade, with the majority of specimens originating in Indonesia. Details on population connectivity may potentially provide important information with regards to harvest management efforts. Genetic structure was examined, using ribosomal ITS1, 5.8S and partial ITS2 sequences on a small scale among populations in the Spermonde Archipelago, South Sulawesi (up to 65 km distance, Φ_st = 0.09), and on a large scale throughout the Indo-Malay Archipelago (up to 2,900 km distance, Φ_st = 0.26). Significant genetic structuring was found at both scales. Within the Spermonde Archipelago isolation by distance as well as local oceanographic features shaped patterns of genetic connectivity. On the large scale, the data revealed genetically distinct populations in Tomini Bay, New Guinea and the Thousand Islands near Jakarta, and a lack of genetic differentiation among populations lying close to or directly in the path of the Indonesian throughflow: from the central Visayas to the Flores Sea (Φ_ct = 0.32). Whilst the influence of both historical and present day processes on genetic structuring of H. actiniformis populations was revealed, large scale results further emphasised the importance of oceanographic dynamics on larval dispersal patterns in this species. Potential for larval input from surrounding populations, and the increased vulnerability of upstream as well as isolated populations should be taken into consideration when setting future harvest quotas.     

华东地区青冈种群的遗传多样性及遗传分化

采用垂直板型聚丙烯酰胺凝胶电泳测定了华东地区6个青冈(Cyclobalanopsis glauca(Thunb.)Oerst.)种群的遗传多样性和遗传分化程度以及基因流。青冈种和种群水平都维持有较高的遗传多样性,期望杂合度分别为0.2252和0.2126,观察杂合度分别为0.1661和0.1771。种群间的遗传分化程度较低,分化度仅为5.6％,种群间的遗传一致度和遗传距离的均值分别为0.9729和0.0276。种群间的分化时间为1.4～27万年。基因流分别为4.21和20.49。     

The dynamics of the restoration of mechanical damage to colonies of the scleractinian coral Porites lutea under conditions of competition with algal settlers for substratum

Natural and anthropogenic catastrophes in the last decades of the 20th century and at the beginning of the new century have evoked significant changes in the biological diversity of coral reefs, viz., the mortality of hermatypic scleractinian corals that are replaced by communities of marine algae and seaweeds. In this connection, the study of the dynamics of algal colonization of a newly formed substratum and competition for the substratum between hermatypic corals and algae is of topical interest. This paper describes the results of investigations conducted in 2003 at the Sesoko Marine Biological Station of Ryukyu University (Okinawa, Japan). Colonies of the massive scleractinian coral Porites lutea were sampled from a fringing reef of Sesoko Island. Damages were inflicted on the upper portions of the colonies, the damaged coral fragments were placed in aquaria and maintained for six months under different light intensities in the absence of herbivorous and corallivorous animals. The experiments showed that under bright (70–90% of incident photosynthetically active radiation, PAR₀) and under moderate light (20–30% of PAR₀), the damaged parts of colonies of about 25 cm² were overgrown by newly formed polyps within six months. Under dim light (2–5% of PAR₀), this period was not sufficient for recovery from lesions. During the first month, the recovery rate of colonies was the highest and depended on the physiological state of the corals and light intensity: in bright and moderate light it was two times and more higher than under low light. During the subsequent months, the recovery rate was significantly lower and depended mainly on the degree of overgrowth of dead portions with algae and on their species composition. Under bright and moderate light, algal colonization of damaged colonies started with the settling of colonial and filamentous diatoms, cyanobacteria and green algae, followed by the formation of the algal turf community. Communities of red calcareous crusts and fleshy algae were mainly formed on damaged corals under low light. A comparison of experimental results showed that damaged colonies maintained in aquaria in the absence of herbivorous and corallivorous animals recovered 1.5 times faster than in the sea. We hypothesize that herbivorous fish have a negative rather than positive influence on the recovery of damaged colonies.     

Genetic variation in the Aleuts of the Pribilof Islands and the Eskimos of Kodiak Island

A sample of Aleuts residing in the Pribilof Islands of St. Paul (N = 163) and St. George (N = 62) and Eskimo residents of Kodiak Island (N = 294) have been typed for genetic variation at 31 discrete genetic markers. Of these, 16 were polymorphic and 15 were monomorphic. Several private polymorphisms previously reported in Eskimo or Alaskan Amerindian populations were absent in both the Aleuts and Kodiak Island Eskimos. Genetic distance analysis shows considerable genetic differentiation between Aleuts and Kodiak Island Eskimos.     

Quantifying the quality of coral bleaching predictions

Techniques that utilize sea surface temperature (SST) observations to predict coral reef bleaching are in common use and form the foundation for predicted global coral reef ecosystem demise within this century. Yet, quality assessments of these methods are typically qualitative or anecdotal. Quality is the correspondence of forecasts with observations and has standard quantitative measures. Here a forecast verification method, commonly used in meteorology, is presented and used to measure the quality of the degree heating weeks (DHW) technique as an exploration of insights that can be gleaned from this methodology. DHW values were calculated from NOAA Optimum Interpolation SST version 2 data and compared to a database of bleaching observations from 1990–2007. Quality is expressed with an objective measure, the Peirce Skill Score (PSS). The quality at varying DHW thresholds above which bleaching was projected to occur is calculated. By selecting the thresholds that maximize quality, the predictive technique is objectively optimized. This results in optimal threshold maps, showing reefs more prone and more resistant to bleaching. Optimization increases the quality of DHW as a predictor of bleaching from PSS = 0.55 to PSS = 0.83, in global average, but the optimal PSS and corresponding DHW values vary significantly from location to location. The coral reef research and management community are urged to adopt the simple, but rigorous tools of forecast verification routinely used in other disciplines so that bleaching forecasts can be quantitatively compared and their quality improved.