Genetic structure of a scleractinian coral, Mycedium elephantotus, in Taiwan

The genetic structure of Mycedium elephantotus (Pallas, 1766) populations from three regions around Taiwan was examined using allozyme electrophoresis. Eight loci were scored from seven enzyme systems. Seven loci were polymorphic under the 95% criterion. The high ratio of observed to expected genotypic diversities at the collection sites (G _O:G _E=0.8 to 1.0) indicate that M. elephantotus propagates predominantly by sexual reproduction. Allele frequencies of M. elephantotus differed significantly among regions (D=0.024 to 0.256, F _ST=0.032 to 0.218, p < 0.001), while populations among collection sites within each region were homogeneous (D=0.000 to 0.015, F _ST=0.010 to 0.022, p > 0.05). Genetic differentiation between populations from southern Taiwan and the Penghu Islands is greater than that between southern and northern Taiwan populations, although the former two regions are much closer geographically. The UPGMA dendrogram based on Nei's unbiased genetic distance showed a clear subdivision of populations into two groupings, northern Taiwan/Penghu Islands and southern Taiwan. A higher level of gene flow was found between M. elephantotus populations in northern Taiwan and the Penghu Islands (N _m=7.56) than that between populations in southern Taiwan and other regions (N _m=0.90 to 1.72). The pattern of genetic subdivision among regions is consistent with the pattern of ocean currents, indicating that genetic differentiation is likely driven by surface circulation vicariance. Received: 2 December 1997 / Accepted: 15 September 1998     

Asynchronous deposition of dense skeletal bands in Porites lutea

Alternate dense and less-dense skeletal bands in massive corals have been used for many years to record the history of growth in species such as Porites lutea and Montastrea annularis, based on the assumption that one dense band and one less-dense band is equivalent to a year's growth. This report demonstrates that specimens of Porites lutea Edwards and Haime (collected from the same neighbourhood in Phuket, Thailand, from November 1983 through November 1984) produce skeletal bands asynchronously and that one year's growth in corals from certain sites may regularly consist of four bands of varying density. The annual banding pattern observed at all sites includes the deposition of a dense band in response to high sedimentation loads and, probably, reduced light levels.     

Oculina patagonica: a non-lessepsian scleractinian coral invading the Mediterranean Sea

The scleractinian coral Oculina patagonica De Angelis is a new immigrant from the Southwest Atlantic to the Mediterranean Sea, having established itself only recently along the Israeli coast. This species is the only scleractinian coral reported to have invaded a new region. In order to understand the swift establishment of this species along the Israeli coast, from 1994 to 1999 we studied its distribution, abundance, reproduction, recruitment, survival, and the effect of bleaching events on its population abundance. In addition, population studies of O. patagonica were performed at several localities along the eastern and western Mediterranean coasts. Highest abundance was recorded along the Spanish coast, reaching 30lj colonies per 10 m line transect in shallow water. Second in abundance was the Israeli coast, with a maximum of 10DŽ colonies per transect. O. patagonica was rare along the coast of Italy, and absent along the Mediterranean coast of France. During the study, both mortality and recruitment along the Israeli coast were very low. In contrast, recruitment along the Spanish coast was very high. Reproduction of the species was studied using gonadal histology. O. patagonica is gonochoric. Female gonads were first observed in May and male gonads in July, both reaching maturity in late August and early September. Naturally occurring azooxanthellate colonies of O. patagonica inhabiting small dark caves developed gonads and spawned in parallel to zooxanthellate colonies exposed to light. No gonads were found in zooxanthellate colonies that underwent bleaching during the reproduction season. The high incidence of bleaching events along the Israeli coast observed throughout the years of this study may explain the low recruitment of new colonies during the same period. In view of its current recruitment patterns, we expect further expansion of O. patagonica in range and abundance in the western Mediterranean, but very small expansion of the population in the eastern Mediterranean, due to repetitive annual bleaching events.     

Reproduction and recruitment of scleractinian corals in a high-latitude coral community, Amakusa, southwestern Japan

Reproduction and recruitment in high-latitude coral populations in Japan have been little studied. A comprehensive study of the reproduction and early life history was conducted on nine common scleractinian coral species in Amakusa, southwestern Japan (32°N) from 2001 to 2003 including; (1) fecundity (the proportion of colonies with mature eggs), (2) timing and synchrony of spawning, (3) initial larval settlement pattern, (4) recruitment, (5) post-settlement mortality. The fecundity was high (76.7–100%) in six of seven species examined in 2002 and 2003. Annual spawning of the seven species occurred from mid July to August in 2001–2003, when seawater temperature was at the annual maximum. Spawning was highly synchronised among conspecific colonies and species in 2002 and 2003, with five species spawning five to nine nights after the full moon and another two spawning around the new moon. Temporal patterns of larval settlement of three spawning species during the first 10 days after spawning were similar to those of other spawning species from low latitudes. The number of scleractinian recruits on settlement plates, deployed from July to October (the major recruitment period at the study site), was low (2 recruits/m²) for the three consecutive years. Post-settlement mortality of 1–1.5 month old spat of five species ranged between 88 and 100% over 3–10 months in the field, similar to the values reported for both high and low latitude species (>94–99%). Among the key stages examined, the low recruitment rate may be the most important step in limiting successful reproduction and recruitment of these high-latitude scleractinian populations. The low recruitment rate may be attributable to (1) the reduced influx of larval supply from other coral populations, which are smaller and more isolated at high-latitudes and (2) the longer precompetent larval phase of broadcast-spawning corals which results in an increased chance of larvae being dispersed away from parent populations.     

Light responses of a scleractinian coral (Plerogyra sinuosa)

During daytime Plerogyra sinuosa Dana displays globular expandable tentacles (bubbles) which foster the photosynthetic ability of the coral. Adaptational responses of this coral to different depths (5–25 m) and light conditions were investigated by photosynthetic pigment analysis, insitu measurements of oxygen production, transplantation and shading experiments. Pigment concentrations per unit tissue dry weight were variable, but unrelated to depth. Pigment concentrations per zooxanthellae cell remained constant and bubble size increased with depth. Light intensity at 25 m was 20 to 25% of the 5-m value, but daily integrated rates of photosynthesis were 65% of the 5-m rates, indicating a higher light utilization efficiency in deeper corals. Coral heads transplanted from 25 to 5 m died within 20 d if not protected against UV-radiation, but corals transplanted from 5 to 25 m acclimatized to the new light condition. Photosynthetic oxygen production and bubble size increased in shaded, sun-adapted corals within 60 min and decreased in sun-exposed, shade-adapted corals. The variable bubble size is interpreted as an adaptational mechanism to optimize light exposure of zooxanthellae.     

Experimental ecology of the temperate scleractinian coral Astrangia danae

The combined effects of temperature, light and symbiont density on the metabolic rate and calcification of the temperate coral Astrangia danae were studied experimentally using colonies containing different concentrations of zooxanthellae. After acclimation to five temperatures between 6.5° and 27°C, and incubation at three light levels and in darkness, respiration and photosynthesis were measured and corrected for rates due to commensals alone. Calcification rates were regressed on zooxanthellae concentration and production in order to define “symbiotic” and “non-symbiotic” averages, and the enhancement of calcification by symbiotic interactions in the polyps. Respiration by the polyparium varied less with temperature between 11.5° and 23°C than that of the commensals, suggesting physiological acclimation by the coral tissue. In-vivo zooxanthellae photosynthesis increased linearly with temperature and was near its maximum at 400 μEin m^?2 s^?1, but the photosynthesis of the endolithic algae of the corallum varied little between 11.5° and 27°C. Calcification at any given temperature was near its maximum at 40 μEin m^?2 s^?1 in both symbiotic and non-symbiotic corals. CaCO₃ deposition increased linearly with temperature in non-symbiotic colonies and in symbiotic colonies incubated in the dark. In symbiotic colonies, calcification in the light increased above these basic rates as temperature rose above 15°C. Below 15°C, symbiotic interactions failed to stimulate calcification, apparently due both to a lowering of zooxanthellae photosynthesis and to a decrease in the enhancing effect of any given level of primary production.     

Reproductive periodicity of the scleractinian coral Lophelia pertusa from the Trondheim Fjord, Norway

Between 2002 and 2008, samples of the cold-water scleractinian coral Lophelia pertusa were collected from the Trondheim Fjord in Norway to examine reproductive periodicity. Collections were made from three locations: Tautra, (63°35.36′N, 10°31.23′E at 40–70 m), Stokkbergneset (63°28.18′N, 09°54.73′E at 110–500 m), and Røberg (63°28.88′N, 09°59.50′E at 250 m). Populations of L. pertusa from the Trondheim Fjord initiated oogenesis in January and spawning occurred from late January to early March the following year. Gametogenic cycles of the female L. pertusa samples overlapped by approximately 2 months, with oogonia visible in January, but this was not evident in the males. This paper provides the most complete gametogenic cycle to date and spawning observations for this important structure-forming species. The results from fjord populations are compared with published and preliminary data from other regions and are discussed in the context of regional differences in physical and biological variables, particularly food supply. Differences in gametogenic cycles within a single species provide a rare opportunity (especially in deep-sea species) to examine potential drivers of reproduction.     

Photosynthesis, respiration and calcification of a zooxanthellate scleractinian coral under submerged and exposed conditions

An experimental set-up was designed to investigate photosynthesis, respiration and calcification of zooxanthellate scleractinian corals under submerged and exposed conditions. The results of experiments to determine the effect of exposure to air on the metabolism of Stylophora pistillata (Esper, 1797) revealed that: (1) maximum gross photosynthesis ( p _{g max}) is 3.6 times higher in water than in air; (2) an indicator of photoadaptation (I _k ) does not reveal any difference between water and air; (3) the difference between submerged and aerial respiration is not statistically significant; (4) aerial photosynthesis–irradiance (P–I) curves display lower initial slopes (α) than aquatic P–I curves; (5) there is no calcification in air. Received: 7 February 1997 / Accepted: 4 March 1997     

Experimental investigation into the effects of suspended sediment on fertilisation, larval survival and settlement in a scleractinian coral

Laboratory and field experiments were used to determine whether high (≃100 mg l⁻¹), low (≃50 mg l⁻¹) and control (≃0 mg l⁻¹) levels of suspended sediment affected fertilisation, larval survival, and larval settlement in the scleractinian coral Acropora digitifera (Dana, 1846). Both high- and low-sediment treatments significantly decreased fertilisation, but post-fertilisation embryonic development was not inhibited by suspended sediments. Larval survival and larval settlement were significantly reduced in high- and low-sediment treatments. No difference was found between high- and low-sediment treatments in any of the three post-spawning processes investigated, suggesting that they are susceptible to sediment concentrations which are not exceptionally high even under natural conditions (>50 mg l⁻¹). The introduction of an additional stress in the form of high levels of suspended sediments coupled with naturally high variability in recruitment may have a considerable effect on the successful supply and settlement of coral larvae to a reef. Given that many coral communities are open reproductive systems, the consequences of disturbance events are not likely to be restricted to the impact area. Recruitment to a population may be reduced significantly in the presence of high levels of suspended sediments because of effects on larval survival and settlement. Recruitment of larvae to adjacent populations may also be affected due to a decreased fertilisation success and potential increases in mortality of larvae passing through the affected site. Received: 13 August 1998 / Accepted: 22 July 1999     

Effects of pCO2 on the interaction between an excavating sponge,Cliona varians,and a hermatypic coral,Porites furcata

Rising dissolved pCO₂ is a mounting threat to coral reef ecosystems. While the biological and physiological impacts of increased pCO₂ are well documented for many hermatypic corals, the potential effects on bioerosion processes remain largely unknown. Increases in pCO₂ are likely to modify the direct interactions between corals and bioeroders, such as excavating sponges, with broad implications for the balance between biologically mediated deposition and erosion of carbonate in reef communities. This study investigated the effects of three levels of CO₂ (present-day, mid-century and end-of-century projections) on the direct interaction between a bioeroding sponge, Cliona varians, and a Caribbean coral, Porites furcata. Increased pCO₂ concentrations had no effect on the attachment rates of C. varians to the corals, and we observed no significant impact of pCO₂ on the survival of either the coral or sponges. However, exposure to end-of-century levels of CO₂-dosing (~750 μatm) reduced calcification in P. furcata and led to a significant increase in sponge-mediated erosion of P. furcata. These findings demonstrate that pCO₂ can enhance erosional efficiency without impacting survival or competitive vigor in these two species. While few studies have considered the influence of pCO₂ on the competitive outcomes of interactions between corals and other reef organisms, our study suggests that assessing the impacts of changing pCO₂ on species interactions is crucial to adequately predict ecosystem-level responses in the future.     

Genetic variation and clonal structure in the scleractinian coral Pocillopora damicornis in the Ryukyu Archipelago, southern Japan

Samples of the scleractinian coral Pocillopora damicornis were collected from six sites located around four islands in the Ryukyu Archipelago, southern Japan, and subjected to allozyme electrophoresis. Seven polymorphic loci were examined for their allelic patterns. The ratio of observed to expected genotypic diversity (0.30 < G _o :G _e  < 0.64), the ratio of the observed number of genotypes to the number of individuals (0.47 < N _g :N _i  < 0.75), and deviations from Hardy–Weinberg equilibrium indicated that asexual reproduction plays a major role in the maintenance of established populations. However, populations were not completely dominated by a single or a few clones, and most clones were represented by only a few individual samples. The high frequency of typhoons in the region suggests that, in P. damicornis, fragmentation caused through occasional exposure to powerful waves is a major mode of asexual reproduction, but asexual production of planulae may also be contributing to the maintenance of populations. A significant genetic differentiation (F ST) was found between the six populations examined (0.027 < F ST < 0.092, average F ST = 0.056). The moderate gene flow is discussed according to characteristics of the larval stage of the species, and to circulation patterns in the region. Received: 7 August 1998 / Accepted: 18 May 1999     

Genetic diversity,clonality and connectivity in the scleractinian coral Pocillopora damicornis: a multi-scale analysis in an insular,fragmented reef system

Clonality and genetic structure of the coral Pocillopora damicornis sensu lato were assessed using five microsatellites in 12 populations from four islands of the Society Archipelago (French Polynesia) sampled in June 2008. The 427 analysed specimens fell into 132 multilocus genotypes (MLGs), suggesting that asexual reproduction plays an important role in the maintenance of these populations. A haploweb analysis of ITS2 sequences of each MLG was consistent with all of them being conspecific. Genetic differentiation was detected both between and within islands, but when a single sample per MLG was included in the analyses, the populations turned out to be nearly panmictic. These observations provide further evidence of the marked variability in reproductive strategies and genetic structure of P. damicornis throughout its geographic range; comparison with results previously obtained for the congeneric species Pocillopora meandrina underlines the importance of life history traits in shaping the genetic structure of coral populations.     

Porites and the Phoenix effect: unprecedented recovery after a mass coral bleaching event at Rangiroa Atoll,French Polynesia

The 1997/1998 El Niño Southern Oscillation (ENSO) was the most severe coral bleaching event in recent history, resulting in the loss of 16 % of the world’s coral reefs. Mortality was particularly severe in French Polynesia, where unprecedented mortality of massive Porites was observed in lagoonal sites of Rangiroa Atoll. To assess the recovery of massive Porites 15 years later, we resurveyed the size structure and extent of partial mortality of massive Porites at Tivaru (Rangiroa). Surveys revealed an abundance of massive Porites colonies rising from the shallow lagoonal floor. Colony width averaged 2.65 m, reaching a maximum of 7.1 m (estimated age of ~391 ± 107 years old). The relative cover of recently dead skeleton within quadrats declined from 42.8 % in 1998 to zero in 2013, yet the relative cover of old dead skeleton increased only marginally from 22.1 % in 1998 to 26.1 % in 2013. At a colony level, the proportion of Porites dominated by living tissue increased from 34.9 % in 1998 to 73.9 % in 2013, indicating rapid recovery of recent dead skeleton to living tissue rather than transitioning to old dead skeleton. Such rapid post-bleaching recovery is unprecedented in massive Porites and resulted from remarkable self-regeneration termed the ‘Phoenix effect’, whereby remnant cryptic patches of tissue that survived the 1997/1998 ENSO event regenerated and rapidly overgrew adjacent dead skeleton. Contrary to our earlier predictions, not only are large massive Porites relatively resistant to stress, they appear to have a remarkable capacity for recovery even after severe partial mortality.     

Synchronous spawnings of 105 scleractinian coral species on the Great Barrier Reef

Following observations of mass spawning of hermatypic corals on the Great Barrier Reef in 1981 and 1982, spawning dates were successfully predicted and documented at five reefs on the Central and Northern Great Barrier Reef in 1983. During the predicted times, 105 species from 36 genera and 11 families were observed to spawn. Of these, 15 species were shown to have an annual gametogenic cycle. All but two of the species observed during mass spawnings shed gametes which underwent external fertilization and development. Synchronous spawning was observed both within and between the five reefs studied, which were separated by as much as 5° of latitude (500 km) or almost a quarter of the length of the Great Barrier Reef. The mass spawning of corals took place on only a few nights of the year, between the full and lastquarter moon in late spring. Maturation of gametes coincided with rapidly rising spring sea temperatures. Lunar and diel cycles may provide cues for the synchronization of gamete release in these species. The hour and night on which the greatest number of species and individuals spawned coincided with low-amplitude tides. Multispecific synchronous spawning, or mass spawning, of scleractinian and some alcyonacean corals represents a phenomenon which is, so far, unique in both marine and terrestrial communities.     

Allozyme electrophoresis demonstrates that the scleractinian coral Montipora digitata is two species

Allozyme electrophoresis at five loci was used to clarify the species status of two morphs of the scleractinian coral Montipora digitata (Dana, 1846), using specimens collected from three locations off the Queensland coast between October 1991 and April 1993. The two morphs, occurring sympatrically, were distinguished by one fixed gene difference at Locus LT-2 and frequency differences at three other loci, indicating reproductive isolation between them. Populations of both morphs were generally in Hardy-Weinberg equilibrium, suggesting that sexual reproduction does occur. Despite sampling designed to reduce the chance of collecting clonemates, the genetic diversity ratio (G _O:G_E) indicated that asexual reproduction also occurs. This is in agreement with the known modes of reproduction for this species. There was significant genetic differentiation between populations of one of the morphs of M. digitata. This could have been influenced by greater degrees of asexual reproduction in this morph, selection within sites, or variation in larval survival and recruitment patterns. Relatively low F _st (differentiation among populations) values found for a broadcast spawning species during this study are in accordance with the hypothesis that brooded planulae are adapted for rapid settlement, whereas planulae produced by broadcast spawners are adapted for widespread dispersal.     

Fungi in corals: black bands and density-banding of Porites lutea and P. lobata skeleton

 Dark coloration of coral skeleton forming black bands is commonly observed in fractured, massive-coral colonies (Porites lutea and P. lobata) collected from May- otte Island in the Mozambique Channel and Moorea Island in French Polynesia. Black-banding was similar in corals from the two areas and was associated with an assemblage of microbial endoliths: Ostreobium queketti, a common siphonal chlorophyte, and a type of Aspergillus-like fungus. Fungi of coral skeletons are capable of euendolithic growth entirely within the skeleton, and of cryptoendolithic growth whereby they spread from the skeleton into the skeletal pores. The morphology and size of fungal hyphae differs significantly between euendolithic, cryptoendolithic and reproductive phases. Reproductive phases involve formation of conidiophores. Insoluble residues in black bands involve a dark pigment and a dark membranous veil. When attacked by fungi, the algae are usually destroyed. They darken and are threaded by dense, dark-brown, fibrous excrescences. The fungi excrete a dark pigment that stains the surrounding skeletal carbonate black. The pigment is organic, and its presence correlates with higher concentrations of polysaccharides. Black bands match high-density bands of the coral skeleton. Both black bands and high-density bands form at the end of the rainy season in Mayotte. Thus, black-banding in the corals studied is caused by a series of events, beginning with an increase in the abundance of endolithic algae followed by an increase in skeletal density. The algae are then attacked by fungi, which produce more cryptoendolithic hyphae and conidia that are associated with production of the dark pigment. Received: 29 January 1999 / Accepted: 29 September 1999     

Evidence for broadcast spawning as well as brooding in the scleractinian coral Pocillopora damicornis

The reproductive activity of 88 colonies of Pocillopora damicornis at Rottnest Island, Western Australia, was examined over 12 mo (20 August 1988 to 20 August 1989). Larvae were found close to the three new moons of February, March and April, 1989, but in large numbers only during March; 67% of all corals produced larvae. There was no obvious connection between gametogenesis and the production of brooded larvae; however, those corals which did not produce, larvae also did not produce ova but did produce, sperm, and must be considered male colonies. All colonies which produced ova went on to produce planulae. Spermatogenesis occurred throughout the year, but oogenesis occurred between September 1988 and April 1989. Dramatic declines in the percentage of polyps containing eggs and sperm occurred in early February and early April 1989. On these dates the amount of lipid in the tissue of the corals decreased, which suggests that the decline in the percentage of polyps with sperm and eggs was not due to the gametes being resorbed by the polyps. Therefore, P. damicornis at Rottnest Island appears to be both a brooder and a broadcaster with asexual reproduction through brooded larvae and sexual reproduction through spawning of gametes. This accounts for earlier evidence in the literature that some input from sexual reproduction had been occurring in the populations of P. damicornis at Rottnest Island. There is no direct evidence for spawning, as broadcasting of gametes has not been observed, but the indirect evidence provides a strong argument for the existence of this dual reproduction.     

The production of sexual and asexual larvae within single broods of the scleractinian coral, Pocillopora damicornis

The genotypic compositions of two populations of the brooding coral, Pocillopora damicornis, were studied by exhaustive sampling of adult colonies on a fine scale in southern Taiwan. In addition, 100 larvae were randomly selected among more than 1,000 brooded larvae collected from a single broodparent within each population. Using 7 polymorphic microsatellite markers, both populations were found to be highly clonal, and 7 clonal lineages were characterized. One clonal lineage (C1) dominated both study areas and comprises 54.9% of all colonies sampled, while any of the other 6 clonal lineages represented no more than 5%. Among the 100 larvae randomly selected for genotyping from each broodparent, the extent of clonal reproduction was high, and only 29 and 6 larvae, respectively, were found to be genotypically different from their broodparent. Among the 35 genotypically distinct larvae, 33 were thought to be derived from sexual reproduction, and 2 were assumed to be clonal propagules that had undergone somatic mutations. Two genotypically identical larvae were also found in one of the 2 sexually derived larva arrays, indicating the possible existence of polyembryony. The high proportions of clonality in both adult colonies and brooded larvae suggest that asexually produced larvae might significantly contribute to the recruitment of local populations. The dense clonal population of P. damicornis in the study area favors the quick recolonization view of clonal propagules after disturbances.