Radiographic studies of reef coral exoskeletons: Rates and patterns of coral growth

X-radiographs were made of vertical slices through the centers of 47 hermatypic coral colonies collected at Eniwetok Atoll, Marshall Islands. The image thus obtained are useful for the study of colony geometry, development, and response to damage.Comparison of radioactive inclusions of known age with previously reported cyclic skeletal density variations normal to the axis of growth confirms the annual nature of the density banding. Growth rates based on density bands and radioactivity inclusions are calculated for all 47 specimens, and measurements of the individual ‘growth bands’ are presented for 25 of them.Bulk densities measured by X-ray transmission ranged from 1.0 to 2.2 g/cm³, with an average range of 1.3–1.6 g/cm³. Intra-specimen skeletal densities typically vary by 10–30%; the period of high density skeletal deposition appears to coincide with the season of higher rainfall and warmer surface water at Eniwetok. Pigment residues left by boring algae are more commonly found in low density portions of the skeletons, but this distribution is believed to result from rather than cause the variations in the density of the deposited aragonite.Linear growth rates for the same specimen vary by factors of two or more from year to year, but the 25 specimens studied did not show a common pattern in the linear growth rate. Other than showing some general trends in growth as a function of species and depth, linear growth rates do not appear to be a particularly informative parameter.The density and growth rate variations are important factors in the measurement of coral growth and metabolism, and to the study of environmental controls of coral growth.     

Food availability affects growth in a coral reef fish

Summary Pomacentrus amboinensis is common on small patch reefs within One Tree Lagoon (Great Barrier Reef), where it preferentially settles onto deep reefs. A preliminary experiment, in which juveniles were transplanted to identical reef structures at two sites, within two depth strata, indicated that juvenile growth and survivorship were better in deeper water. The hypothesis that this difference was due to food availability was tested by a supplemental feeding experiment, carried out at another two randomly chosen sites, within the same two depth strata. Fish were fed each day over a one month period, during which no mortality was observed. The growth rates of juveniles were markedly higher on all food-supplemented reefs, when compared to controls. Growth differed between depth strata, but there was no interaction between the food x depth factors, which would have suggested a greater effect of food supplementation in either habitat. Thus, although the difference between depths cannot be attributed to food availability, the results have a more general significance. Food appears to be a limiting resource (in terms of growth) in both the marginal shallow habitat, and the more suitable deeper habitat.     

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.     

Intraspecific competition in a reef coral: effects on growth and reproduction

Summary Growth rates and reproduction of a branching coral (Stylophora pistillata) were compared in the presence and in the absence of intraspecific competition. Field experiments demonstrated a significant decline in the growth rate of competing colonies compared to noncompeting control colonies; the growth rate slowed in all of the interacting individuals, irrespective of their place in the hierarchy of the intraspecific dominance or of their color morph. In case of immediate killing of the subordinate, the dominant colony grew at a normal rate. In addition to the marked decrease in the growth rate of interacting colonies, the typical symmetry shape of these colonies was changed to an abnormal growth form. The number of female gonads per polyp was significantly reduced in colonies competing intraspecifically, and the typical synchrony in reproduction among different branches of a given colony was changed and desynchronized. Again, these results did not correlate with the hierarchy of dominance. We conclude that intraspecific competition in reef corals involves great investment of energy. The ecological significance and the different pathways of this competition are discussed.     

Long-term cleaner fish presence affects growth of a coral reef fish

Cleaning behaviour is considered to be a classical example of mutualism. However, no studies, to our knowledge, have measured the benefits to clients in terms of growth. In the longest experimental study of its kind, over an 8 year period, cleaner fish Labroides dimidiatus were consistently removed from seven patch reefs (61-285 m(2)) and left undisturbed on nine control reefs, and the growth and parasite load of the damselfish Pomacentrus moluccensis determined. After 8 years, growth was reduced and parasitic copepod abundance was higher on fish from removal reefs compared with controls, but only in larger individuals. Behavioural observations revealed that P. moluccensis cleaned by L. dimidiatus were 27 per cent larger than nearby conspecifics. The selective cleaning by L. dimidiatus probably explains why only larger P. moluccensis individuals benefited from cleaning. This is the first demonstration, to our knowledge, that cleaners affect the growth rate of client individuals; a greater size for a given age should result in increased fecundity at a given time. The effect of the removal of so few small fish on the size of another fish species is unprecedented on coral reefs.     

Climate change and coral reef connectivity

This review assesses and predicts the impacts that rapid climate change will have on population connectivity in coral reef ecosystems, using fishes as a model group. Increased ocean temperatures are expected to accelerate larval development, potentially leading to reduced pelagic durations and earlier reef-seeking behaviour. Depending on the spatial arrangement of reefs, the expectation would be a reduction in dispersal distances and the spatial scale of connectivity. Small increase in temperature might enhance the number of larvae surviving the pelagic phase, but larger increases are likely to reduce reproductive output and increase larval mortality. Changes to ocean currents could alter the dynamics of larval supply and changes to planktonic productivity could affect how many larvae survive the pelagic stage and their condition at settlement; however, these patterns are likely to vary greatly from place-to-place and projections of how oceanographic features will change in the future lack sufficient certainty and resolution to make robust predictions. Connectivity could also be compromised by the increased fragmentation of reef habitat due to the effects of coral bleaching and ocean acidification. Changes to the spatial and temporal scales of connectivity have implications for the management of coral reef ecosystems, especially the design and placement of marine-protected areas. The size and spacing of protected areas may need to be strategically adjusted if reserve networks are to retain their efficacy in the future.     

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.     

Effects of herbivory,nutrients, and reef protection on algal proliferation and coral growth on a tropical reef

Maintaining coral reef resilience against increasing anthropogenic disturbance is critical for effective reef management. Resilience is partially determined by how processes, such as herbivory and nutrient supply, affect coral recovery versus macroalgal proliferation following disturbances. However, the relative effects of herbivory versus nutrient enrichment on algal proliferation remain debated. Here, we manipulated herbivory and nutrients on a coral-dominated reef protected from fishing, and on an adjacent macroalgal-dominated reef subject to fishing and riverine discharge, over 152 days. On both reefs, herbivore exclusion increased total and upright macroalgal cover by 9-46 times, upright macroalgal biomass by 23-84 times, and cyanobacteria cover by 0-27 times, but decreased cover of encrusting coralline algae by 46-100% and short turf algae by 14-39%. In contrast, nutrient enrichment had no effect on algal proliferation, but suppressed cover of total macroalgae (by 33-42%) and cyanobacteria (by 71% on the protected reef) when herbivores were excluded. Herbivore exclusion, but not nutrient enrichment, also increased sediment accumulation, suggesting a strong link between herbivory, macroalgal growth, and sediment retention. Growth rates of the corals Porites cylindrica and Acropora millepora were 30-35% greater on the protected versus fished reef, but nutrient and herbivore manipulations within a site did not affect coral growth. Cumulatively, these data suggest that herbivory rather than eutrophication plays the dominant role in mediating macroalgal proliferation, that macroalgae trap sediments that may further suppress herbivory and enhance macroalgal dominance, and that corals are relatively resistant to damage from some macroalgae but are significantly impacted by ambient reef condition.     

珊瑚礁生态保护与管理研究

珊瑚礁以其极高的生物多样性和生物生产力以及优美的自然景观 ,为人们提供了生活需要和游乐的资源 ,但同时也受到过度利用的破坏 ,尤其是近年来显得更为严重 ,因而珊瑚礁的生态保护与管理成为近 2 0年来倍受关注的问题。本文回顾了国内外珊瑚礁生态保护和管理的一些研究成果 ,通过自然和社会经济调查 ,并根据保护、研究和可持续利用的原则 ,将雷州半岛灯楼角珊瑚礁保护区划分为野生区 ,保护区、季节性封闭区和一般使用区 ,并强调公众参与、社区组织和领导组成、教育和培训、资源管理等为保护和管理中的措施     

Relationships between coral reef substrata and fish

 The objective of this work is to identify which substrata characteristics (such as coral morphology, coral diversity, coral species richness, percentage coverage by live coral or by algae) influence the structure and abundance of fish communities. The study was carried out at Reunion Island, Indian Ocean, where six sites were sampled in two zones (reef flat and outer reef slope). Quantitative data were collected by visual census techniques, along a linear transect of 50 m for the substratum, and a belt of 50×2 m for the fish communities. Correspondence analysis (CA) and an optimising cluster analysis, called dynamic clustering method (DCM) were used to describe and compare fish assemblages with the benthic composition. The relationships between benthic and fish communities were examined using the classes revealed by the partitioning of the substratum with DCM. This partitioning allowed us to derive four classes of substratum: the non-disturbed reef flat, the non-disturbed outer reef slope, the perturbed reef habitat and the reef pass. The analysis of the partitioning based on the coral variables suggests that there are significant relationships between benthic and fish assemblages. Accepted: 26 July 1996     

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.     

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.     

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.     

Functional versatility supports coral reef biodiversity

We explore the role of specialization in supporting species coexistence in high-diversity ecosystems. Using a novel ordination-based method to quantify specialist and generalist feeding structures and diets we examined the relationship between morphology and diet in 120 wrasses and parrotfishes from the Great Barrier Reef. We find that wrasses, despite their morphological diversity, exhibit weak links between morphology and diet and that specialist morphologies do not necessarily equate to specialized diets. The dominant pattern shows extensive overlap in morphology (functional morphospace occupation) among trophic groups; fish with a given morphology may have a number of feeding modes. Such trophic versatility may lay the foundation for both the origins and maintenance of high biodiversity on coral reefs.     

Measuring coral reef decline through meta-analyses

Coral reef ecosystems are in decline worldwide, owing to a variety of anthropogenic and natural causes. One of the most obvious signals of reef degradation is a reduction in live coral cover. Past and current rates of loss of coral are known for many individual reefs; however, until recently, no large-scale estimate was available. In this paper, we show how meta-analysis can be used to integrate existing small-scale estimates of change in coral and macroalgal cover, derived from in situ surveys of reefs, to generate a robust assessment of long-term patterns of large-scale ecological change. Using a large dataset from Caribbean reefs, we examine the possible biases inherent in meta-analytical studies and the sensitivity of the method to patchiness in data availability. Despite the fact that our meta-analysis included studies that used a variety of sampling methods, the regional estimate of change in coral cover we obtained is similar to that generated by a standardized survey programme that was implemented in 1991 in the Caribbean. We argue that for habitat types that are regularly and reasonably well surveyed in the course of ecological or conservation research, meta-analysis offers a cost-effective and rapid method for generating robust estimates of past and current states.