Transportation techniques for massive scleractinian corals

Transportation techniques for scleractinian corals have been described mainly for fragments and small colonies. As part of a recent study on captive sexual reproduction of the Caribbean species Montastrea annularis and Diploria strigosa, we transported relatively large (max. diameter of 21 cm), heavy (max. weight of 9,200 g) colonies of both species from Curaçao, Netherlands Antilles, to Rotterdam, The Netherlands. A new transportation technology was applied whereby the corals were transplanted to specially designed PVC crosses to provide stabilization during transport. In two transports (November 2001 and February 2002), 100 colonies were transported submerged, in a shipping time of >35 hr. The survival rate measured 2 weeks after transport was 100%. Four and 8 months after transport, respectively, two colonies of D. strigosa died without any obvious cause. In November 2002 we observed an outbreak of Dark Spots disease (DSD) affecting two‐thirds of the colonies of M. annularis. Although the colonies did not show any symptoms when they were collected, the disease most probably was transferred when the coral were transported from the field to the laboratory. The presented method is appropriate for transporting large, heavy corals–especially for scientific purposes. In general, species‐specific properties, colony size, and transportation time determine which transportation method should be applied. In the future, there may be a shift toward transports of fragments, coral larvae, and primary polyps to reduce collections in the field. Zoo Biol 23:165–176, 2004. © 2004 Wiley‐Liss, Inc.     

Morphological plasticity in scleractinian corals

When describing coral shape and form the term phenotypic plasticity, i.e. environment-induced changes in morphology, is often used synonymously with intraspecific variation. Variation, however, may simply be due to genetic differentiation (polymorphism). Of the 1314 extant scleractinian coral species, less than 20 have been tested for plastic responses. Morphological plasticity has important implications for coral identification, as skeletal features used in coral systematics are directly affected by environment. Furthermore, plastic changes can indicate how corals acclimatise to environmental change. The studies that have examined phenotypic plasticity in corals experimentally can be divided into two groups, i.e. 'non-clonal'—those that have transplanted whole colonies or fragments of colonies (but not treated the fragments as clones) to new environments, and 'clonal'—those that have transplanted colony fragments and used them as clone-mates. The use of clone-mates is preferable as it facilitates the identification of among-genotype variation for plasticity. The heterogeneous nature of the reef environment makes identifying the parameters that affect coral morphology difficult in the field, but there are also many problems conducting suitable aquarium experiments. Nevertheless, evidence to date suggests light and water movement are the most important variables inducing change. As these factors are known to be axiomatic to coral growth, it is possible that associated plastic changes in corals are adaptive; however, this hypothesis is yet to be tested rigorously.     

Tumor formations in scleractinian corals

A highly localized incidence of skeletal malformations (tumors) in the scleractinian coralsPlatygyra pini andP. sinensis on an inshore fringing reef at Cockle Bay, Magnetic Island within the Great Barrier Reef province is reported. These tumors are typified by a localized area of increased growth rate resulting in roughly circular protuberances extending up to 4.5 cm above the colony's surface. In both species, similar proportions of their populations carried tumors (24.1 % inP. pini and 18.7 % inP. sinensis). Larger colonies (>80 cm in diameter) are at least 7 times more likely to possess tumors than smaller colonies (<40 cm in diameter). X-radiographs of the skeletal malformations indicate a point of origin, presumably from a single budded polyp with subsequent, localized, accelerated growth. The mean radial growth rate of the tumorous area was 29 % greater than that of the surrounding normal regions. In contrast to the normal tissue, the tumorous tissue exhibited proliferation of cells, atrophied gastrodermal cells and mesenterial filaments which were larger and disordered in structure. The environmental conditions at Cockle Bay are relatively extreme with high turbidity, periodic exposure of the reef flat, abrupt changes in salinity during the wet season and mechanical damage to corals caused by unpredictable cyclonic storms. It is suggested that a combination of environmental stresses coupled with an injury inflicted on the corals are possible stimuli that initiate the development of these abnormal growth through either bacterial attack or the development of an aberrant polyp during tissue repair.     

Histological data in a combined phylogenetic analysis of scleractinian reef corals

Scleractinian systematics have undergone rapid changes due to increased use of molecular phylogenetics and new perspectives on skeletal morphology from micromorphology and microstructure. Despite this increase in characters there are still unresolved clades in the phylogeny, indicating that more characters are needed. This study investigates a new source of morphological data within the soft tissue of Indo‐Pacific scleractinian corals. Features of tissue layers, especially cnidocytes, are described in hematoxylin and eosin stained thin sections. Based on this new histological data source, a combined analysis with mitochondrial DNA and skeletal data is performed using parsimony and Bayesian analysis. Parsimony analysis yields three most‐parsimonious trees similar to trees based on Bayesian analysis. Character maps are also produced that show origination of histomorphological traits at deep nodes within the phylogeny. In general, both analyses retain the previously designated families Lobophylliidae and Merulinidae, but some genera are found to be paraphyletic. Nonetheless, the combined analysis produces a highly resolved and well‐supported phylogeny, which could lead to more effective use of biological conservation metrics based on evolutionary distinctiveness. These results show for the first time that inclusion of histomorphological characters improves the resolution of phylogenetic analyses of reef corals. J. Morphol. 277:494–511, 2016. © 2016 Wiley Periodicals, Inc.     

Hormones and reproduction in scleractinian corals

Most broadcast spawning scleractinian corals synchronously release gametes during a brief annual spawning period. In southern Taiwan, the mass spawning of scleractinians occurs in lunar mid-March. The exact cues triggering this annual phenomenon remain unclear. A scleractinian coral, Euphyllia ancora has been selected as a model for the hormones and reproduction studies. Testosterone (T) and estradiol (E2) in free and glucuronided forms were identified and consistently detected in coral polyps throughout the year. Peak levels of free E2, glucuronided E2 and T were obtained in the coral tissue just prior to spawning. The presence of specific aromatase activity was demonstrated in coral tissue. Higher concentrations of free E2 than glucuronided E2 were detected in the coral tissue throughout the year. In contrast, higher levels of glucuronided E2 than free E2 and glucuronided T were found in seawater during mass coral spawning. Furthermore, immunoreactivity and biological activity of immunoreactive gonadotropin-releasing hormone (irGnRH) was detected and quantified in coral tissue. Coral extracts (irGnRH) and mammalian (m)GnRH agonist had a similar dose-dependent effect on luteinizing hormone (LH) release in black porgy fish pituitary cells (in vitro). Peak levels of irGnRH were detected during the spawning period. In in vivo experiments, mGnRH agonist time- and dose-dependently stimulated aromatase activity, as well as the levels of T and E2 in free and glucuronided forms in coral. In conclusion, our data suggest that irGnRH and glucuronided E2 may play important roles in the control of reproduction and mass spawning in corals.     

Reassessing evolutionary relationships of scleractinian corals

The widely accepted family tree of Scleractinia published by Wells, based on a combination of morphological coral taxonomy and the fossil record, has recently been revised by Veron. It is now possible to test the validity of some of the conclusions reached by these and other authors by the use of molecular techniques. This paper reviews the results to date. Studies of ribosomal DNA have shown that the Scleractinia are monophyletic, i.e. derived from the same ancestral taxon. Extensions of this same data set now indicate that the Poritidae and Dendrophylliidae, with their fossil antecedents, may each warrant separate suborder status. They further suggest (a) that the Suborder Faviina (faviids, mussids and their allies) should probably be retained as a monophyletic group and (b) that Wells' original account of the isolated position of the Pocilloporidae and Astrocoeniidae is correct. These conclusions all accord with Veron's family tree. However, the Fungiina, even after removal of the Poritidae, are unlikely to be a monophyletic group at suborder level. The molecular data further show that externally observable morphological characters used in the taxonomy of extant corals distinguish families more reliably than do internal micro-skeletal characters frequently used in coral palaeontology.     

Differential physiological responses of two congeneric scleractinian corals to mineral accretion and an electric field

Despite increasing popularity of ‘electric’ reefs as a means for reef restoration, there is a distinct lack of quantitative evidence supporting the alleged benefits of this method. This study investigated the effects of an electric field versus an electric field in combination with a cathode on coral growth (skeletal extension) rates, coral survival, zooxanthella densities, chlorophyll a (chl a) concentrations, and chlorophyll fluorescence of Acropora pulchra and A. yongei. Coral transplants were grown for 4 months under three treatment conditions: (1) on an iron cathode, (2) on bamboo inside an electric field, or (3) on bamboo in the absence of an electric field. Contrary to predictions, coral growth rates of both species were highest inside the electric field and not on the cathode. Except for chl a concentrations, the cathode had a significant adverse effect on all measured variables for A. yongei but not for A. pulchra. Treatment had no effect on the survival of A. pulchra, while mortality rates of A. yongei were significantly higher in the presence of mineral accretion compared to the electric field and control. A. yongei on the cathode featured low zooxanthella densities, depressed electron transport rates (rETR) and maximum quantum yield (F _v/F _m), and reduced growth. By contrast, treatment had no effect on the fluorescence characteristics of A. pulchra, and zooxanthella densities were highest for corals on the cathode, coincident with high growth rates relative to the control. Overall, the data indicate that the proposed benefits of the mineral accretion technology to meet important objectives of reef rehabilitation with regard to colony growth and survival should be considered with caution.     

Larval development of certain gamete-spawning scleractinian corals

Embryogenesis and larval development were documented in 19 species of hermatypic scleractinians which release gametes during the summer coral spawning season on the Great Barrier Reef. Cleavage of fertilized eggs began approximately 2 h after spawning in all species, and gave rise to blastulae after 7–10 h. Endoderm formation in Platygyra sinensis was by invagination, and this appeared to occur in all species studied. All species observed at 36 h after spawing were mobile and full mobility was reached by 48 h. Settlement of planulae placed in aquaria occurred between 4 and 7 days after fertilization. These results suggest that larval corals produced by most gamete-releasing coral species are likely to be dispersed away from the parent reef.     

Skeletal ontogeny in basal scleractinian micrabaciid corals

The skeletal ontogeny of the Micrabaciidae, one of two modern basal scleractinian lineages, is herein reconstructed based on serial micro‐computed tomography sections and scanning electron micrographs. Similar to other scleractinians, skeletal growth of micrabaciids starts from the simultaneous formation of six primary septa. New septa of consecutive cycles arise between septa of the preceding cycles from unique wedge‐shaped invaginations of the wall. The invagination of wall and formation of septa are accompanied by development of costae alternating in position with septa. During corallite growth, deepening invagination of the wall results in elevation of septa above the level of a horizontal base. The corallite wall is regularly perforated thus invaginated regions consist of pillars inclined downwardly and outwardly from the lower septal margins. Shortly after formation of septa (S2 and higher cycles) their upper margins bend and fuse with the neighboring members of a previous cycle, resulting in a unique septal pattern, formerly misinterpreted as “septal bifurcation.” Septa as in other Scleractinia are hexamerally arranged in cycles. However, starting from the quaternaries, septa within single cycles do not appear simultaneously but are inserted in pairs and successively flank the members of a preceding cycle, invariably starting from those in the outermost parts of the septal system. In each pair, the septum adjacent to older septa arises first (e.g., the quinaries between S2 and S4 before quinaries between S3 and S4). Unique features of micrabaciid skeletal ontogeny are congruent with their basal position in scleractinian phylogeny, which was previously supported by microstructural and molecular data. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

A computational method for quantifying morphological variation in scleractinian corals

Morphological variation in marine sessile organisms is frequently related to environmental factors. Quantifying such variation is relevant in a range of ecological studies. For example, analyzing the growth form of fossil organisms may indicate the state of the physical environment in which the organism lived. A quantitative morphological comparison is important in studies where marine sessile organisms are transplanted from one environment to another. This study presents a method for the quantitative analysis of three-dimensional (3D) images of scleractinian corals obtained with X-ray Computed Tomography scanning techniques. The advantage of Computed Tomography scanning is that a full 3D image of a complex branching object, including internal structures, can be obtained with a very high precision. There are several complications in the analysis of this data set. In the analysis of a complex branching object, landmark-based methods usually do not work and different approaches are required where various artifacts (for example cavities, holes in the skeleton, scanning artifacts, etc.) in the data set have to be removed before the analysis. A method is presented, which is based on the construction of a medial axis and a combination of image-processing techniques for the analysis of a 3D image of a complex branching object where the complications mentioned above can be overcome. The method is tested on a range of 3D images of samples of the branching scleractinian coral Madracis mirabilis collected at different depths. It is demonstrated that the morphological variation of these samples can be quantified, and that biologically relevant morphological characteristics, like branch-spacing and surface/volume ratios, can be computed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Rethinking the phylogeny of scleractinian corals: a review of morphological and molecular data

Scleractinian corals, which include the architects of coral reefs, are found throughout the world's oceans and have left a rich fossil record over their 240 million year history. Their classification has been marked by confusion but recently developed molecular and morphological tools are now leading to a better understanding of the evolutionary history of this important group. Although morphological characters have been the basis of traditional classification in the group, they are relatively few in number. In addition, our current understanding of skeletal growth and homology is limited, and homoplasy is rampant, limiting the usefulness of morphological phylogenetics. Molecular phylogenetic hypotheses for the order, which have been primarily focused on reef-building corals, differ significantly from traditional classification. They suggest that the group is represented by two major lineages and do not support the monophyly of traditional suborders and most traditional families. It appears that once a substantial number of azooxanthellate taxa are included in molecular phylogenetic analyses, basal relationships within the group will be clearly defined. Understanding of relationships at lower taxonomic levels will be best clarified by combined analyses of morphological and molecular characters. Molecular phylogenies are being used to inform our understanding of the evolution of morphological characters in the Scleractinia. Better understanding of the evolution of these characters will help to integrate the systematics of fossil and extant taxa. We demonstrate how the combined use of morphological and molecular tools holds great promise for ending confusion in scleractinian systematics.     

Metabolism of estrogens and androgens by scleractinian corals

Estrogens and androgens are steroids that act as reproductive hormones in vertebrates. These compounds have also been detected in reef-building corals and other invertebrates, where they are hypothesized to act as bioregulatory molecules. Experiments were conducted using labeled steroid substrates to evaluate metabolism of estrogens and androgens by coral homogenates. GC-MS analysis of 13C-labeled steroids showed that Montipora capitata coral homogenates or fragments could convert estradiol to estrone and testosterone to androstenedione and androstanedione, evidence that M. capitata contains 17beta-hydroxysteroid dehydrogenase and 5alpha-reductase. When homogenates from three coral species and symbiotic dinoflagellates (zooxanthellae) were incubated with tritiated steroid substrates, metabolites separated by thin-layer chromatography confirmed that 17beta-hydroxysteroid dehydrogenase activity occurred in all species tested. NADP+ was the preferred cofactor in dehydrogenation reactions with coral homogenates. Reduction of estrone and androstenedione occurred at lower rates and aromatization of androgens was not observed. It is unclear whether estrogens detected previously in coral tissues are produced endogenously or sequestered in coral tissue from dietary or environmental sources. Previous studies have demonstrated that corals can take up estrogens from the water column overlying coral reefs. Considered in total, these observations suggest corals could alter the concentration or form of steroids available to reef organisms.     

Ontogeny of 'epithecal' and septal structures in scleractinian corals

The ontogenetic development of a solitary scleractinian coral, Flabellum distinctum Edwards & Haime, has been studied in serial thin section, with special attention being paid to epithecal nature in relation to septal growth. The term 'epitheca' has been confusingly used for two different skeletal structures: epitheca ( sensu stricto ) and marginotheca. The latter is here newly proposed. 'epitheca' is defined as a calcareous investment developed on the outside of other skeletal structures of a corallite. It can be distinguished from the marginotheca in section by lacking a dark line (calcification centre) and by being unrelated to the formation of septa. 'marginotheca' defines the outer margin of the main skeletal structures of a corallite. It has a dark line which functionally coincides with that of the eutheca. It is of primary origin, preceding formation of septa and provides the origin of the septa. The marginotheca is one of the more important and fundamental skeletal structures for coral classification.     

Regulation of release of antibacterials from stressed scleractinian corals

Recently, we showed that mechanical stress on scleractinian (stony) corals caused a rapid release of antibacterial material (referred to as coral antibacterial activity, or CAA), which killed various bacterial species, including the coral pathogen Vibrio coralliilyticus . We now report on studies on the regulation of CAA release from stressed scleractinian corals. Corals can repeatedly release highly active CAA as a result of sequential stress inductions. Coral fragments were transferred 19 times from one beaker into another with a stress induction each time after 10 min. There was a decrease in the level of antibacterial activity released during the first four to five transfers. After the fifth transfer, the corals kept releasing CAA for the rest of the experiment with no significant decrease. Apparently, the release of CAA is downregulated by feedback inhibition, depending on the concentration of CAA in the surrounding water. By separating CAA-treated V. coralliilyticus from the surrounding water, it was shown that CAA was bound irreversibly to bacterial cells in a stoichiometric manner. Approximately 4 × 10² bacterial cells were sufficient to bind 1 U of CAA. Resident coral bacteria were more resistant to CAA than bacteria isolated from seawater, suggesting an ecological role for CAA. CAA release was obtained from corals after removal of the mucus layer, and the mucus itself contained antibacterial activity.     

Negative indirect effects of neighbors on imperiled scleractinian corals

Predation pressure on an individual may be influenced by spatial associations with other organisms. In the case of rare and imperiled species, such indirect interactions may affect the persistence and recovery of local populations. This study examined the effects of coral neighborhood composition on the foraging behavior and impact of the corallivorous gastropod, Coralliophila abbreviata. We conducted a manipulative field experiment in which focal colonies of the threatened scleractinian coral Acropora cervicornis had no neighbors, conspecific neighbors, alternative prey (Orbicella faveolata) neighbors, or non-prey (Porites asteroides) neighbors. Individually tagged C. abbreviata were then seeded into the study area and allowed to colonize the experimental plots. Initial colonization was significantly affected by the species of neighboring corals and snail abundance after colonization was negatively correlated with focal colony growth. Snails exhibited a strong prey preference for A. cervicornis over O. faveolata and responded numerically to neighborhood quality (i.e., relative preference for neighboring corals). Thus, conspecific neighbors had the greatest predator-mediated negative effect on focal colony performance followed by O. faveolata neighbors. The results suggest that C. abbreviata mediate apparent competition between O. faveolata and A. cervicornis as both species contributed to the local abundance of their shared predator. Additionally, home range estimates for tagged C. abbreviata were calculated, compared among sexes, and found to be significantly greater for males than for females. Overall, this study sheds light on the foraging behavior of an important coral predator and highlights the potential importance of consumer-mediated indirect interactions in the dynamics of severely reduced populations. The results also have direct implications for conservation and population enhancement efforts.     

Endosymbiotic flexibility associates with environmental sensitivity in scleractinian corals

Flexibility in biological systems is seen as an important driver of macro-ecosystem function and stability. Spatially constrained endosymbiotic settings, however, are less studied, although environmental thresholds of symbiotic corals are linked to the function of their endosymbiotic dinoflagellate communities. Symbiotic flexibility is a hypothesized mechanism that corals may exploit to adapt to climate change. This study explores the flexibility of the coral–Symbiodinium symbiosis through quantification of Symbiodinium ITS2 sequence assemblages in a range of coral species and genera. Sequence assemblages are expressed as an index of flexibility incorporating phylogenetic divergence and relative abundance of Symbiodinium sequences recovered from the host. This comparative analysis reveals profound differences in the flexibility of corals for Symbiodinium, thereby classifying corals as generalists or specifists. Generalists such as Acropora and Pocillopora exhibit high intra- and inter-species flexibility in their Symbiodinium assemblages and are some of the most environmentally sensitive corals. Conversely, specifists such as massive Porites colonies exhibit low flexibility, harbour taxonomically narrow Symbiodinium assemblages, and are environmentally resistant corals. Collectively, these findings challenge the paradigm that symbiotic flexibility enhances holobiont resilience. This underscores the need for a deeper examination of the extent and duration of the functional benefits associated with endosymbiotic diversity and flexibility under environmental stress.     

Free-living scleractinian corals on reefs of the Seychelles Islands

Thirty-nine species of unattached scleractinian corals that belong to 22 genera of 9 families were found on the Seychelles reefs. Variations of the colony form of corals living on soft sediments under continuous wave action are described. Irrespective of their initial growth form and taxonomic position, corals assume a form close to spherical. Because of the worldwide deterioration of coral reefs, the adaptation to changing ecological conditions by reef-building corals needs to be studied.     

Nucleotide sequences of highly repetitive DNA from scleractinian corals

The staghorn coral genome contains 5% of a satellite DNA, consisting of 80 to 300 x 10(3) copies of a 118-bp repeat unit per haploid Acropora genome.     

First report of folliculinid ciliates affecting Caribbean scleractinian corals

This is the first report of a ciliate of the genus Halofolliculina infecting hard coral species of six families (Acroporidae, Agaricidae, Astrocoeniidae, Faviidae, Meandrinidae and Poritidae) and milleporids in the Caribbean. Surveys conducted during 2004–2005 in Venezuela, Panama and México confirmed that this ciliate affects up to 25 scleractinian species. The prevalence of this ciliate at the coral community level was variable across sites, being most commonly found at Los Roques, Venezuela, and at Bocas del Toro, Panama (prevalence 0.2–2.5%), but rarely observed in the Mexican Caribbean. Ciliates were more prevalent within populations of acroporids (Acropora palmata, Acropora cervicornis and Acropora prolifera) in Los Roques. Recent observations also corroborate the presence of these ciliates in Curacao and Puerto Rico. Our observations indicate that ciliates affecting corals have a wider distribution than previously thought, and are no longer exclusively found in the Indo-Pacific and Red Sea.