Tissue age affects calcification in the scleractinian coral Madracis mirabilis

In this study, two factorial experiments were used to investigate the role of tissue age in affecting the phenotypic expression of calcification in scleractinian corals. Both experiments tested whether calcification was altered by tissue age and whether corals of different ages exploit plasticity to differing degrees by altering calcification rates under new environmental conditions. To isolate age and size effects, branches of the Caribbean coral Madracis mirabilis were broken into a distal portion that was functionally young and a proximal portion that was functionally old. Fragments were transplanted from a deep (17 m) to a shallow (9 m) site in a Jamaican lagoon to test whether age affected the plasticity of calcification. Both experiments demonstrated that calcification scaled isometrically in the two age groups, and although scaling exponents were indistinguishable statistically among ages, young fragments calcified faster than old fragments. Thus, the effect of age on calcification rate was absolute and independent of size. However, the interactive effect of age and depth was not significant, demonstrating that ability to alter calcification rate (i.e., the extent of phenotypic plasticity for this trait) was unaffected by age. Together, these patterns are consistent with the hypothesis that the proximal modules (i.e., polyps) of M. mirabilis are subject to physiological senescence, as has been reported for other clonal organisms, including algae, fungi, plants, bryozoans, ascidians, and other cnidarians.     

Simulation and analysis of flow patterns around the scleractinian coral Madracis mirabilis (Duchassaing and Michelotti)

Three-dimensional morphologies of Madracis mirabilis were obtained using X-ray computed tomography scanning techniques. The morphologies were used to simulate the flow patterns around the colony. In the simulations, the thin-branching low-flow morph with a relatively larger branch-spacing was compared with the more compact high-flow morph of M. mirabilis. For both morphologies, the inside-colony flow velocities were computed for Reynolds numbers ranging from 154 to 3840. In the high-flow morph, it was found that in the range of investigated Reynolds numbers a stagnant region develops within the colony, whereas in the low-flow morph the stagnant region disappeared. Experiments done under natural conditions suggest that a morph is adapted to a certain external flow velocity and develops a stagnant region below a particular threshold for the external flow velocity. When the external flow velocity exceeds a certain threshold, which is characteristic for the growth form, the core velocity becomes equal to the external velocity. A potential application of a profile of core velocities for a range of Reynolds numbers for a certain morph is the prediction of the optimal external flow velocity for a certain morph, and this can be used to assess the state of the physical (palaeo-) environment.     

A comparison between coral colonies of the genus Madracis and simulated forms

In addition to experimental studies, computational models provide valuable information about colony development in scleractinian corals. Using our simulation model, we show how environmental factors such as nutrient distribution and light availability affect growth patterns of coral colonies. To compare the simulated coral growth forms with those of real coral colonies, we quantitatively compared our modelling results with coral colonies of the morphologically variable Caribbean coral genus Madracis. Madracis species encompass a relatively large morphological variation in colony morphology and hence represent a suitable genus to compare, for the first time, simulated and real coral growth forms in three dimensions using a quantitative approach. This quantitative analysis of three-dimensional growth forms is based on a number of morphometric parameters (such as branch thickness, branch spacing, etc.). Our results show that simulated coral morphologies share several morphological features with real coral colonies (M. mirabilis, M. decactis, M. formosa and M. carmabi). A significant correlation was found between branch thickness and branch spacing for both real and simulated growth forms. Our present model is able to partly capture the morphological variation in closely related and morphologically variable coral species of the genus Madracis.     

Revisiting coral reef connectivity

A large river plume generated by anomalous precipitation and oceanic circulation associated with Hurricane Mitch was detected off Honduras in October 1998 using SeaWiFS ocean color images. This event provides the background for analyzing connectivity between coral reefs and land in the Meso-American reef system. We discuss the potential implications of such short-term events for disease propagation and nutrification, and their potential significance in evolutionary processes.     

Diurnal temporal patterns of the diversity and the abundance of reef fishes in a branching coral patch in New Caledonia

Small‐scale spatial and temporal variability in animal abundance is an intrinsic characteristic of marine ecosystems but remains largely unknown for most animals, including coral reef fishes. In this study, we used a remote autonomous unbaited video system and recorded reef fish assemblages during daylight hours, 10 times a day for 34 consecutive days in a branching coral patch of the lagoon of New Caledonia. In total, 50 031 fish observations belonging to 114 taxa, 66 genera and 31 families were recorded in 256 recorded videos. Carnivores and herbivore‐detritus feeders dominated the trophic structure. We found significant variations in the composition of fish assemblages between times of day. Taxa richness and fish abundance were greater in the early morning and in the late afternoon than during the day. Fourteen taxa displayed well‐defined temporal patterns in abundance with one taxon influenced by time of day, six influenced by tidal state and seven influenced by both time of day and tidal state. None of these 14 taxa were piscivores, 10 were herbivore‐detritus feeders, three were carnivores and one was plankton feeder. Our results suggest a diel migration from feeding grounds to shelter areas and highlight the importance of taking into account small‐scale temporal variability in animal diversity and abundance when studying connectivity between habitats and monitoring communities.     

Bioindication in coral reef ecosystems

The concept of bioindication in the sense of the use of organisms for detecting environmental stress has been employed in coral reef conservation and management for the past several years. Important tools are coral growth rates and various community parameters, notably hard coral cover. The present need is the optimal coordination of international efforts for the earliest possible institution of an effective monitoring system.     

A novel reef coral symbiosis

Reef building corals form close associations with unicellular microalgae, fungi, bacteria and archaea, some of which are symbiotic and which together form the coral holobiont. Associations with multicellular eukaryotes such as polychaete worms, bivalves and sponges are not generally considered to be symbiotic as the host responds to their presence by forming physical barriers with an active growth edge in the exoskeleton isolating the invader and, at a subcellular level, activating innate immune responses such as melanin deposition. This study describes a novel symbiosis between a newly described hydrozoan (Zanclea margaritae sp. nov.) and the reef building coral Acropora muricata (=A. formosa), with the hydrozoan hydrorhiza ramifying throughout the coral tissues with no evidence of isolation or activation of the immune systems of the host. The hydrorhiza lacks a perisarc, which is typical of symbiotic species of this and related genera, including species that associate with other cnidarians such as octocorals. The symbiosis was observed at all sites investigated from two distant locations on the Great Barrier Reef, Australia, and appears to be host species specific, being found only in A. muricata and in none of 30 other species investigated at these sites. Not all colonies of A. muricata host the hydrozoans and both the prevalence within the coral population (mean = 66%) and density of emergent hydrozoan hydranths on the surface of the coral (mean = 4.3 cm⁻², but up to 52 cm⁻²) vary between sites. The form of the symbiosis in terms of the mutualism–parasitism continuum is not known, although the hydrozoan possesses large stenotele nematocysts, which may be important for defence from predators and protozoan pathogens. This finding expands the known A. muricata holobiont and the association must be taken into account in future when determining the corals’ abilities to defend against predators and withstand stress.     

Perspectives in coral reef hydrodynamics

Knowledge of skeletogenesis in scleractinian corals is central to reconstructing past ocean and climate histories, assessing and counteracting future climate and ocean acidification impacts upon coral reefs, and determining the taxonomy and evolutionary path of the Scleractinia. To better understand skeletogenesis and mineralogy in extant scleractinian corals, we have investigated the nature of the initial calcium carbonate skeleton deposited by newly settling coral recruits. Settling Acropora millepora larvae were sampled daily for 10 days from initial attachment, and the carbonate mineralogy of their newly deposited skeletons was investigated. Bulk analyses using Raman and infrared spectroscopic methods revealed that the skeletons were predominantly comprised of aragonite, with no evidence of calcite or an amorphous precursor phase, although presence of the latter cannot be discounted. Sensitive selected area electron diffraction analyses of sub-micron areas of skeletal regions further consolidated these data. These findings help to address the uncertainty surrounding reported differences in carbonate mineralogy between larval and adult extant coral skeletons by indicating that skeletons of new coral recruits share the same aragonitic mineralogy as those of their mature counterparts. In this respect, we can expect that skeletogenesis in both larval and mature growth stages of scleractinian corals will be similarly affected by ocean acidification and predicted environmental changes.     

Model of a coral reef ecosystem

The results of modelling of a coral reef ecosystem at French Frigate Shoals and independent field measures of benthic primary productivity indicate relatively good agreement between food required by consumer trophic levels and organic carbon produced by primary producers. Based upon the high internal predation necessary for the model to match primary production estimates, we reason that the ecosystem is primarily regulated from the top down by forces of predation and that primary production appears to be controlled by nutrients, rate limits, and the distribution of space and habitat. In spite of relatively high primary productivity, potential yield at the top of the food chain is low because of high internal predation and high trophic complexity (6 trophic levels). Fishery yield might be maximized by harvesting low on the food chain particularly if top carnivores can be cropped to release predator pressure on selected prey. Agreement between field measures of metabolism and model (ECOPATH) results provides reasonable confidence that the model can be used as one tool for resource management.     

Experimental biology of coral reef ecosystems

Coral reef ecosystems are at the crossroads. While significant gaps still exist in our understanding of how “normal” reefs work, unprecedented changes in coral reef systems have forced the research community to change its focus from basic research to understand how one of the most diverse ecosystems in the world works to basic research with strong applied implications to alleviate damage, save, or restore coral reef ecosystems. A wide range of stressors on local, regional, and global spatial scales including over fishing, diseases, large-scale disturbance events, global climate change (e.g., ozone depletion, global warming), and over population have all contributed to declines in coral cover or phase shifts in community structure on time scales never observed before. Many of these changes are directly or indirectly related to anthropogenically induced changes in the global support network that affects all ecosystems. This review focuses on some recent advances in the experimental biology of coral reef ecosystems, and in particular scleractinian corals, at all levels of biological organization. Many of the areas of interest and techniques discussed reflect a progression of technological advances in biology and ecology but have found unique and timely application in the field of experimental coral reef biology. The review, by nature, will not be exhaustive and reflects the author's interests to a large degree. Because of the voluminous literature available, an attempt has been made to capture the essential elements and references for each topic discussed.     

The use of coral nubbins in coral reef ecotoxicology testing

While there is an urgent demand to establish reliable ecotoxicological assays for reef corals, there has not been yet an available source material that can supply the high number of colony replicates needed for reliable tests. In past experiments, the major obstacle to obtaining as many fragments as possible had been the damage inflicted to donor colonies by pruning. In this paper, we present the application of coral nubbins, a novel source material for coral ecotoxicology assays. Nubbins from the branching Red Sea coral Stylophora pistillata (n>450) were used for evaluating the impacts of water soluble fractions from a crude oil, an oil dispersant and dispersed oil. Coral nubbins (minute coral fragments in the size of one to several polyps) harvested from a single colony are genetically identical to each other, may be obtained in any quantity needed and whenever research activities demand their use. Several dozens of nubbins can be obtained from a single small branch in branching coral species, a procedure that has minimal impact on donor genotypes. Nubbins production is a low cost procedure and requires limited maintenance space. Results of short and long-term acute ecotoxicological tests are revealed and discussed here, indicating the advantageous use of nubbins as ubiquitous coral material for toxicology assays and physiological studies.     

Disturbance and the structure of coral reef fish communities on the reef slope

Three levels of physical disturbance were applied to corals in permanent 10x10 m quadrats along a section of fringing reef at Lizard Island on the Great Barrier Reef to investigate the response of fish assemblages. Tabular and corymbose corals were overturned and left in situ, reducing total hard coral cover from ˜55% to ˜47%, ˜43%, and ˜34%. Despite pre-existing associations with benthic cover, all fish groups examined (pomacentrids, labrids, chaetodontids, and acanthurids) were resistent to benthic disturbances at the level and scale at which they were applied. Partial Mantel's tests, in combination with partial Canonical Correspondence Analysis enabled spatial and temporal variation to be factored out from experimental effects. Most of the variation in the fish community could be assigned to spatio-temporal variables, indicating that spatial structure over the reef landscape may moderate localised disturbance effects. This study indicates that coral reef fish assemblages may be more resistant to disturbance than many correlative studies would suggest, and highlights a need for further information on levels and scales of natural habitat disturbance in order to apply a structured approach to the experimental investigation of the importance of habitat in structuring coral-reef fish assemblages.     

Extinction vulnerability of coral reef fishes

With rapidly increasing rates of contemporary extinction, predicting extinction vulnerability and identifying how multiple stressors drive non-random species loss have become key challenges in ecology. These assessments are crucial for avoiding the loss of key functional groups that sustain ecosystem processes and services. We developed a novel predictive framework of species extinction vulnerability and applied it to coral reef fishes. Although relatively few coral reef fishes are at risk of global extinction from climate disturbances, a negative convex relationship between fish species locally vulnerable to climate change vs. fisheries exploitation indicates that the entire community is vulnerable on the many reefs where both stressors co-occur. Fishes involved in maintaining key ecosystem functions are more at risk from fishing than climate disturbances. This finding is encouraging as local and regional commitment to fisheries management action can maintain reef ecosystem functions pending progress towards the more complex global problem of stabilizing the climate.     

Phylogeography and the conservation of coral reef fishes

Here we present a review of how the study of the geographic distribution of genetic lineages (phylogeography) has helped identify management units, evolutionary significant units, cryptic species, and areas of endemism, and how this information can help efforts to achieve effective conservation of coral reefs. These studies have confirmed the major biogeographic barriers that were originally identified by tropical species distributions. Ancient separations, identified primarily with mtDNA sequence comparisons, became apparent between populations on each side of the barriers. The general lack of correlation between pelagic larval duration and genetic connectivity across barriers indicates that life history and ecology can be as influential as oceanography and geography in shaping evolutionary partitions within ocean basins. Hence, conservation strategies require a recognition of ecological hotspots, those areas where habitat heterogeneity promotes speciation, in addition to more traditional approaches based on biogeography. Finally, the emerging field of genomics will add a new dimension to phylogeography, allowing the study of genes that are pertinent to recent and ongoing differentiation, and ultimately providing higher resolution to detect evolutionary significant units that have diverged in an ecological time scale.     

Augmented coral reef monitoring using a stationary reef monitoring system

Coral reef monitoring is a reliable tool to assess the effect of climate change as corals are sensitive to increases in water temperatures between 30 °C and 35 °C resulting in bleaching - a whitening process when the corals lose their color and the reefs begin to die. Existing satellite-based monitoring products facilitate coral bleaching monitoring over large spatial scales, but their use in predicting local scale stress that influences the bleaching severity across reefs is limited. In this paper, we describe a Stationary Reef Monitoring System (SRMS) that monitors the time evolution of coral reefs through the photography of nearby coral clusters. Simultaneously, the SRMS measures and records environmental parameters such as temperature, solar irradiance (PAR), and salinity in the waters surrounding the coral colonies. When deployed in the sea, the SRMS detected a 0.1–0.4 °C variability in temperature between the in situ and satellite datasets. The SRMS uses color photography along with quantitative data on environmental parameters to monitor the health of corals and eliminates the need for physical/visual verification of coral health by a diver. By this approach, one can determine the stress thresholds of corals and identify the vulnerable and resilient reefs so as to prioritize conservation efforts.     

Effects of variations in carbonate chemistry on the calcification rates of Madracis auretenra (= Madracis mirabilis sensu Wells, 1973): bicarbonate concentrations best predict calcification rates

Physiological data and models of coral calcification indicate that corals utilize a combination of seawater bicarbonate and (mainly) respiratory CO₂ for calcification, not seawater carbonate. However, a number of investigators are attributing observed negative effects of experimental seawater acidification by CO₂ or hydrochloric acid additions to a reduction in seawater carbonate ion concentration and thus aragonite saturation state. Thus, there is a discrepancy between the physiological and geochemical views of coral biomineralization. Furthermore, not all calcifying organisms respond negatively to decreased pH or saturation state. Together, these discrepancies suggest that other physiological mechanisms, such as a direct effect of reduced pH on calcium or bicarbonate ion transport and/or variable ability to regulate internal pH, are responsible for the variability in reported experimental effects of acidification on calcification. To distinguish the effects of pH, carbonate concentration and bicarbonate concentration on coral calcification, incubations were performed with the coral Madracis auretenra (= Madracis mirabilis sensu Wells, 1973) in modified seawater chemistries. Carbonate parameters were manipulated to isolate the effects of each parameter more effectively than in previous studies, with a total of six different chemistries. Among treatment differences were highly significant. The corals responded strongly to variation in bicarbonate concentration, but not consistently to carbonate concentration, aragonite saturation state or pH. Corals calcified at normal or elevated rates under low pH (7.6–7.8) when the seawater bicarbonate concentrations were above 1800 μm . Conversely, corals incubated at normal pH had low calcification rates if the bicarbonate concentration was lowered. These results demonstrate that coral responses to ocean acidification are more diverse than currently thought, and question the reliability of using carbonate concentration or aragonite saturation state as the sole predictor of the effects of ocean acidification on coral calcification.     

Sponge-mediated coral reef growth and rejuvenation

Sponges mediate consolidation of Porites furcata rubble on shallow Caribbean reefs by quickly adhering to rubble and stabilizing it until carbonate secreting organisms can grow and consolidate it to the reef. Experimental investigations demonstrate that the entire cycle from (1) temporary binding of rubble by sponges, through (2) rubble consolidation by encrusting coralline algae, to (3) colonization of consolidated rubble by corals, can be completed within 10 months. Bound rubble both adds to vertical reef growth and also provides stable substrata for colonization by corals. Corals that colonize stabilized rubble are damaged less and survive better than on unstable rubble. Rubble that is not temporarily stabilized by sponges does not become bound to the reef, because continuous movement disturbs the consolidation process, and does not provide suitable substrata for settlement and growth of corals. Sponge-mediated consolidation of rubble may increase rates of reef growth and enhance reef recovery after damage. This new role for sponges in reef growth is not obvious from examination of the internal fabric of a reef frame. Spongemediated consolidation may help to explain geographic and temporal differences in growth and morphology among shallow reefs of ramose corals.