Sexual reproduction in the coral Pocillopora verrucosa (Cnidaria: Scleractinia) in KwaZulu-Natal, South Africa

The mode of reproduction in Pocillopora verrucosa Ellis and Solander, 1786 varies between geographically isolated regions. This scleractinian coral is common along the KwaZulu-Natal coast, and its reproductive mode and period of reproduction were assessed using histological preparations. The study was undertaken from 1992 to 1994 and showed that P. verrucosa is a simultaneous hermaphrodite and broadcast spawner in KwaZulu-Natal. Gametogenesis occurs from October to January, with the gametes maturing simultaneously so that spawning can be synchronised at new moon in January (mid-summer). Zooxanthellae are present in the mature oocytes. Received: 10 September 1997 / Accepted: 6 July 1998     

Highly repeated DNA sequences in the scleractinian coral genusAcropora: Evaluation of cloned repeats as taxonomic probes

A plasmid clone containing highly repeated DNA sequence fromAcropora formosa was used to probe slot blots of genomic DNA and Southern blots of Taq I digests from other Acroporidae. Homologous repeated DNA sequences were detected in 12 otherAcropora species, but not in three other species of Acroporidae (Astreopora sp.,Montipora digitata andM. aequituberculata). Slot-blot experiments indicated that the repeated sequences inAcropora latistella, A. tenuis, A. longicyathus andA. nobilis were distantly homologous with the clonedA. formosa repeat, whereas those inA. pulchra, A. millepora, A. valida, A. hyacinthus andA. microphthalma were highly homologous with this probe and those inA. digitifera were intermediate. Relatedness of the highhomology group toA. formosa was assessed by comparison of Taq I-digestion patterns. The predominant repeats inA. pulchra andA. hyacinthus had two Taq I sites per repeat unit (as didA. formosa), whereas repeats inA. digitifera, A. valida andA. microphthalma had one Taq I site per repeat; the pattern given byA. millepora implied that its highly homologous repeat units contained either one or two Taq I sites. Within the high-homology group, decreasing number of Taq I sites implies increasing taxonomic distance fromA. formosa. The relatedness series implied by these data differs from that based on morphological criteria.     

A study of mucus from the solitary coral Fungia fungites (Scleractinia: Fungiidae) in relation to photobiological UV adaptation

The ultraviolet (UV)-absorbance spectrum (300 to 360 nm) of mucus obtained from Fungia fungites (L. 1758, collected in Tahiti in 1991) after being exposed to air for up to 5 min was measured, and UV-absorbing compounds were demonstrated to be present in the mucus, with a peak at 332 nm. The concentration of these UV-absorbing compounds was at a maximum in the first 2 min of secretion and decreased thereafter. Concentration was significantly related to the weight of the coral. Also, as corals were adapted to bathymetric levels of UV radiation, mucus concentration of UV-absorbing compounds decreased significantly with increasing depth.     

Comparative aspects of sexual reproduction in the Caribbean coral genus Diploria (Scleractinia: Faviidae)

The increasing decline of Caribbean reef-building species underlines the importance of more information on their reproductive biology and ecology. This study compares the reproductive biology of the three species of Diploria, an important Caribbean reef-building genus affected by bleaching and disease, by: (a) characterizing their gametogenetic cycles simultaneously, (b) exploring the spatial and temporal variability in reproductive traits within and across species, (c) assessing fecundity and the minimum size of sexual reproduction, and (d) comparing information with other studies in the Caribbean. Starting in 1999, one tissue core was collected every month (for 17 months) from each of at least five large, tagged colonies of each species in La Parguera, Puerto Rico. Three more temporally spaced samplings were done during summer of 2000 and winter–spring of 2001 and 2002. Tissues were fixed in Helly’s solution, rinsed in fresh water, decalcified with 10% HCl, and preserved in 70% ethanol, embedded in paraplast, cut, and thin slides (7 μm) prepared and stained with Heidenhain’s Aniline-Blue method. Maximum egg size, number of eggs and spermaries were measured and gametogenetic cycles assessed. Microscopic observations confirmed that all three species were simultaneous hermaphrodites with a single, annual gametogenetic cycle. Spermatocytes and oocytes developed within the same mesentery but were not intermingled. Female gametes developed several months earlier than that of males, but both reached maturation simultaneously. Minor differences in the onset of gametogenesis were found for D. strigosa and D. clivosa, with both spawning after 10 p.m. between 8 and 10 days after the August and/or September full moons. D. labyrinthiformis on the other hand, began oogenesis in early July and spawned after 11 p.m. between 7 and 10 days after the April and/or May full moons. Spatial and temporal variability in mean number of spermaries and eggs/mesentery among colonies within and among species were found. D. labyrinthiformis had consistently significant higher mesenterial and polyp fecundity compared to the other two species over the 3 years. Average egg diameter varied between 240 and 246 μm and was similar among the three species. D. strigosa and D. clivosa sexual reproductive characteristics were similar to those of other large broadcast spawning Faviidae (i.e. Montastraea spp) in the region. In contrast, D. labyrinthiformis was a spring spawner (similar to Colpophyllia natans in Puerto Rico), which together with its significantly different micromorphology suggests that it might be phylogenetically more distant to the other taxa.     

Reproduction of Dendronephthya hemprichi (Cnidaria: Octocorallia): year-round spawning in an azooxanthellate soft coral

Sexual reproduction in the azooxanthellate octocoral Dendronephthya hemprichi Klunzinger, 1877 was studied in Eilat (Red Sea) for 2 years beginning March 1989. D. hemprichi is a gonochoric broadcasting species. Gonads at all developmental stages were found throughout the year. Small-sized oocytes and sperm sacs, 51 to 100 μm in diameter, are highly abundant, accompanied by numerous smaller primordial gonads. These features result from continuous gametogenesis, and lead to year-round oocyte and sperm maturation and release. Gamete release was observed in the laboratory on all monitoring dates, and most of the female colonies spawned repeatedly for several successive nights. The reproductive features of D. hemprichi constitute an exception to the generalization that free-spawning reef-corals have brief and synchronized broadcast-spawning episodes. This species demonstrates early onset of first reproduction, which further increases its reproductive output. Received: 7 May 1997 / Accepted: 15 May 1997     

Isolation and cloning of DNA from somatic tissue of soft corals (Cnidaria: Octocorallia)

A method is described for isolating high molecular weight DNA from somatic tissue of soft corals. The method avoids problems associated with the presence of nucleases, pigments and other secondary metabolites in soft corals. Tissue is frozen in liquid nitrogen, pulverized with a pestle and mortar, and the powder extracted with buffered sodium dodecyl sulphate at 4°C in the presence of phenol/chloroform. This technique has been applied toAlcyonium, Sinularia, Sarcophyton andLobophytum species. Repeated sequences have been cloned fromAlcyonium sp., and used to probe slot-blots of genomic DNA and Southern blots of restriction digests. Homologous repeated sequences were detected in threeAlcyonium sp., but not in threeSinularia sp., despite these genera being closely related.     

Sexual reproduction, larval development and benthic planulae of the solitary coral Monomyces rubrum (Scleractinia: Anthozoa)

The reproductive biology of the solitary ahermatypic coral Monomyces rubrum was studied in northeastern New Zealand between January 1996 and January 1998. The period of oogenesis lasted around 11 months, from late January to December, while spermatogenesis was more rapid, starting in late August and culminating in a spawning period in early December. Reproduction commenced at a polyp size of around 1,000 mm³ (5-6 years old) and the maximum estimated fecundity of the largest corals (7,000 mm³) was no more than 200 eggs. Oocytes were probably fertilized while within the mesentery and were shed into the coelenteron where they developed, via a solid blastula stage, for approximately 1 month. Planulae were relatively large, 3-4 mm in length and 1-2 mm diameter at the time of release, and crawled or swam immediately to the substratum. Peaks of planula shedding were semi-lunar in January 1997, but only one peak was observed in January 1998. The production of a few large rapidly settling larvae by this member of the family Flabellidae is consistent with the trend for solitary short-lived corals from other families to brood larvae rather than spawn gametes.     

Sexual reproduction of the reef coral Montastreacavernosa (Scleractinia: Faviidae) in the Santa Marta area, Caribbean coast of Colombia

Sexual reproduction of the reef-building coral Montastreacavernosa was studied in the Santa Marta area, Caribbean coast of Colombia, from October 1990 to October 1991. The area is subjected to a seasonal upwelling–outwelling regime. From microscopical analysis of tissue sections sampled each lunar month around the full moon, it was confirmed that this is a gonochoric, broadcasting species, with a single gametogenic cycle per year, and a 1:1 sex ratio. Oogenesis began a few weeks after spawning, and extended for 11 mo. The onset of spermatogenesis occurred just before the full moon of June, later than at other localities where sea-water temperatures during the first half of the year are not as low. The amount of reproductive tissue strongly increased in both sexes after July, in association with an increase in sea-water temperature and a decrease in photoperiod. The gonad index of fully mature female colonies was about four times lower than at other localities, perhaps due to the stressful seasonal regime. The male gonad index was lower than that of the female, indicating sex-related differences in the rates of biomass allocation to reproduction. As in other Caribbean localities, spawning occurred after the full moons of August and September. However, there was some evidence of a third spawning episode after the full moon of October, possibly associated with a delay in the occurrence of maximum sea-water temperatures in near-equatorial localities compared to higher latitudes. The probability of cross-fertilization in this species with a gonochoric breeding system and a broadcasting mode of reproduction is favored by its balanced sex ratio, its usually high within-reef zone abundance, and by localized and repeated spawning episodes, synchronized by lunar phase. Received: 19 April 1996 / Accepted: 24 September 1996     

DNA fingerprints of a gorgonian coral: a method for detecting clonal structure in a vegetative species

Clonal reproduction, a common life history strategy among sessile marine invertebrates, can lead to high local abundances of one to a few genotypes in a population. Analysis of the clonal structure of such populations can provide insight into the ecological and evolutionary history of the population, but requires markers that can identify individual genets. Forensic and demographic studies have demonstrated that DNA fingerprinting can provide markers that are unique for an individual genotype. We have generated DNA fingerprints for over 70 colonies of the clonal gorgonian, Plexaura A (Plexaura sp. A) collected from June 1990 through July 1991 in the San Blas Islands, Panama. DNA fingerprints within a singic individual were identical and fingerprinting resolved multiple genotypes within and among reefs. On one reef in the San Blas Islands, Panama, 59% of the colonies sampled were of one genotype and this genotype was not found on any other sampled reefs. A previous study using tissue grafts identified 13 putative clones on these reefs, while DNA fingerprints of the same colonies differentiated 17 genotypes. The present study demonstrates the utility of DNA fingerprinting for distinguishing clones and for identifying clonal structure of marine invertebrate populations.     

Use of electrical stimulation for discharging cnidom components of a species of Cerianthus (Anthozoa: Cnidaria)

Direct-current electrical stimulation was used as a means of discharging cnidom components from the marginal tentacles of a species of Cerianthus from Southern Queensland, Australia, in order to facilitate the study of cnidom components by light microscopy. The cnidom consists of b-rhabdoids (3 types), spirocysts, and ptychocysts. Threshold pulses of between 1 and 2 V with a frequency of 1 pulse per second for 2 sec discharge all types of cnidae. However, large numbers of certain cnida types discharge after a longer period of stimulation.     

Highly repetitive DNA sequences provide evidence for a lack of gene flow between two morphological forms of Herdmania momus (Ascidiacea: Stolidobranchia)

The genomic structure of two morphological forms of the pyurid ascidian Herdmania momus — H. momus forma curvata and H. momus forma grandis—are compared using a rapidly evolving highly repetitive element isolated from the genome of H. momus forma curvata. A 663 bp (base pair) Cla I satellite sequence is present only in the genome of H. momus forma curvata, and hence can be used as a form-specific marker. Low-stringency Southern blot analyses, using the Cla I satellite as a probe, revealed that the genomes of H. momus forma grandis and another pyurid ascidian, Pyura stolonifera, do not contain similar sequences to the H. momus forma curvata repetitive element. In addition to this genomic dissimilarity, there are a number of significant reproductive and developmental differences between the two H. momus forms, and interform fertilisation rates are significantly lower than intraform rates. The molecular, reproductive and developmental differences between H. momus forma curvata and H. momus forma grandis indicate the presence of strong barriers to gene flow.     

Mitochondrial DNA sequence variation in deep-sea bamboo coral (Keratoisidinae) species in the southwest and northwest Pacific Ocean

The Keratoisidinae are a poorly known, but phenotypically diverse group of deepwater corals. Recent developments in deepwater trawling in the southwest Pacific have provided many more specimens of bamboo corals. Two sub-regions of the mitochondrial genome were sequenced to test genetic relationships among specimens collected over a wide geographical range (27–50° S): a sub-region of the large-subunit rRNA (16S rRNA), characterized by a highly variable insertion/deletion (INDEL#2) region; and a non-coding region between COII and COI. Based on DNA haplotypes, 14 species of Keratoisidinae were recognized among 88 specimens from deep water in the southwest Pacific Ocean. The common haplotypes also appeared in specimens collected in the northwest Pacific Ocean and may indicate that some bamboo coral species are widespread in the Pacific, or that the mitochondrial markers are insensitive to recent speciation events. Many specimens were taken from flat bottom areas and, contrary to assumptions, the bamboo corals are not endemic to seamounts. The closure of some seamounts to trawling will protect bamboo corals from extinction, but not from local depletion.Communicated by M.S. Johnson, Crawley     

Effect of nutrient enrichment on the complementary (secondary) metabolite composition of the soft coral Sarcophyton ehrenbergi (Cnidaria: Octocorallia: Alcyonaceae) of the Great Barrier Reef

 A long-term study of the effects of nutrient enrichment on coral reefs (ENCORE Experiment) was carried out at One Tree Island (23°30′S; 152°96′E), Great Barrier Reef, between 1992 and 1996. The experiment involved the addition of water-soluble nutrients to 12 microatolls which contained a range of organisms and were situated within the larger lagoon. Three replicates of each of three nutrient treatments (nitrogen, phosphorus, nitrogen plus phosphorus) and an untreated set of three control atolls were prepared using the 12 selected microatolls. As part of the larger ENCORE experiment, changes in the chemical composition of the alcyonacean soft coral Sarcophyton ehrenbergi Marenzeller (Octocorallia: Alcyonacea), placed in the treatment microatolls, were monitored for a 1 yr period in an attempt to detect any responses attributable to nutrient enrichment. Multivariate analyses were performed to determine whether there were any patterns of response in the different nutrient treatments. At the level of individual metabolites, there were no clear treatment effects. However, the ratio of bioactive or stress metabolites (terpenes) to energy storage metabolites (lipids), referred to as the “physiological-change indicator”, revealed effects of nutrient enrichment. Nitrogen enrichment resulted in a trend towards higher physiological-change indicators than control or phosphorus treatments in the majority of cases, while phosphorus enrichment significantly decreased the ratio relative to controls. In most cases, the physiological-change indicator increased in soft soft corals relocated into contact with the scleractinian Pocillopora damicornis. The potential of soft corals to serve as indicators of a changed nutrient regime is discussed. Received: 23 November 1998 / Accepted: 23 September 1999     

Analysis of ribosomal ITS1 sequences indicates a deep divergence between Rhodactis (Cnidaria: Anthozoa: Corallimorpharia) species from the Caribbean and the Indo-Pacific/Red Sea

This paper explores the implications of ribosomal ITS1-sequence analysis for evolutionary relationships between four species of corallimorphs which are morphologically very similar but have distinct distributions. The analyses highlight the unreliability of morphological criteria in establishing systematic relationships amongst tropical corallimorphs. Both parsimony and distance methods of phylogenetic analysis strongly support a deep divergence between the Caribbean species Rhodactics sanctithoma and the three morphologically-related species R. howesii and R. indosinensis from the Indo-Pacific and R. rhodostoma from the Red Sea. The analyses also strongly support a closer relationship between the Red Sea species, R. rhodostoma, and one of the Indo-Pacific species, R. howesii, than between these two and the second Indo-Pacific species, R. indosinensis. The first of these results is presumably a consequence of the closure of the Central American Seaway, the event which led to the separate development of Atlantic and Indo-Pacific coral faunas. However, the lack of detailed distribution data represents a major barrier to reconstructing the evolutionary history of the genus.     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Chemical aspects of mass spawning in corals. II. (-)-Epi-thunbergol,the sperm attractant in the eggs of the soft coral Lobophytum crassum (Cnidaria: Octocorallia)

Sea turtles migrate between feeding and nesting areas that are often geographically separated by hundreds or thousands of kilometers. Observations of their aggregations at sea and at nesting beaches have led to the hypothesis that sea turtles migrate in socially structured groups. While this migratory strategy is common to many marine vertebrates, socially facilitated behavior is not well documented in testudines. In 1990 and 1991, we attached satellite transmitters to olive ridleys (Lepidochelys olivacea Eschscholtz) found ovipositing together during a mass nesting at Nancite Beach, Costa Rica, to determine whether they migrate independently or in groups after they leave the nesting beach. Results showed that the turtles were not spatially associated during the internesting period, were capable of re-establishing themselves as a group during a subsequent nesting emergence, and were not spatially associated during their postnesting migrations to oceanic feeding areas. We suggested that what appear to be socially structured groups of L. olivacea are in fact individual turtles simultaneously using the same habitat.     

Multivariate characterisation of the habitats of seven species of Malayan mudskippers (Gobiidae: Oxudercinae)

Mudskippers (Gobiidae: Oxudercinae) are amphibious gobies. The occurrence and habitats of seven sympatric species of Malayan mudskippers were recorded: Boleophthalmus boddarti, Boleophthalmus pectinirostris, Periophthalmodon schlosseri, Periophthalmus chrysospilos, Periophthalmus gracilis, Periophthalmus variabilis and Scartelaos histophorus. A total of 14 surveys were made at six localities along 120 km of the west coast of the Malay Peninsula in August 1996 and September 2006. A multivariate set of ordinal parameters were used to measure the quantity of environmental water in mudskippers’ habitats, and three guilds of species were discriminated. The resulting pattern mirrored the adaptive scenario depicted by physiological and anatomical studies of mudskippers’ terrestriality, although in one case a terminal taxon was less terrestrial than other more basal ones. Differentiation of trophic niches may also play a role in habitat selection by the herbivorous Boleophthalmus spp., which were only found in unvegetated, illuminated areas and on wet mud at low tide. With the more terrestrial guild, a significant negative co-occurrence suggested direct interspecific competition. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ecological,genetic, and morphological differences among three <Emphasis Type="Italic">Pavona</Emphasis> (Cnidaria: Anthozoa) species from the Pacific coast of Panama

Ecological, genetic, and morphological differences among three Panamanian Pacific Pavona species with strongly developed collines (Pavona varians, P. frondifera, and P. chiriquiensis) were examined. Ecological factors included geographical distributions of species, habitat preferences, interspecific interactions, reproductive ecology, and tolerance to bleaching. Genetic differences were based on the electrophoretic analysis of ten allozyme loci. Morphological analyses consisted of tissue coloration, colony morphology, and measurements and counts of ten macro- and micro-skeletal characters. P. varians, present on reefs or in coral communities, is the most widely distributed and shows considerable morphological variation. P. chiriquiensis, a recently described species, encrusts basalt rock and has little morphological variation. P. frondifera is a reef dweller with a compact foliose morphology. Tissue coloration varies from light to dark brown in P. varians, from pink to brown in P. frondifera, and from brick red to brown or silvery in Pavona chiriquiensis. Also, the white to silvery polyp mouths of the latter species are a diagnostic feature that allows an easy identification in the field. Aggressive dominance during short-term interspecific interactions were as follows: Pavona chiriquiensis>P. varians>P. frondifera. P. chiriquiensis and P. varians showed contrasting responses to sea warming during the 1997-1998 El Niño Southern Oscillation. Whereas entire P. chiriquiensis bleached and died within 4 weeks of exposure to 30-31°C, colonies of P. varians did so only on their upper surfaces. The response of P. frondifera to elevated temperatures was not observed because it is mainly present in the Gulf of Panama where coral bleaching was absent in 1997-1998. The genetic data indicated that P. chiriquiensis differed strongly from both P. varians and P. frondifera, with Nei's unbiased genetic distances of 0.434 and 0.379, respectively. A fixed difference between P. varians and P. frondifera, and P. chiriquiensis exists at the triose phosphate isomerase (TPI-2) locus. A nearly fixed difference between P. chiriquiensis and P. frondifera and between P. chiriquiensis and P. varians was found at the hexokinase (HK) locus. P. varians differed slightly from P. frondifera with Nei's unbiased genetic distance of 0.068. No fixed difference was found between P. varians and P. frondifera. There were strong differences between P. chiriquiensis and P. varians in spawning times and gamete characteristics. Spawning in P. varians and P. chiriquiensis is 12 h out of phase. Also, eggs of the former species are white to beige and positively buoyant whereas those of the latter species are dark green and neutrally to negatively buoyant. No reproductive data are yet available for P. frondifera. Calicular diameters are significantly greater in P. chiriquiensis than in the other two species. In contrast, corallum thickness is greater in P. varians and P. frondifera than in P. chiriquiensis. Canonical discriminant function analysis readily separated the three species.