Reef coral diversity and global change

Regional anthropogenic processes such as pollution, dredging, and overfishing on coral reefs currently threaten the biodiversity of stony corals and other reef-associated organisms. Global climate change may interact with anthropogenic processes to create additional impacts on coral diversity in the near future. In order to predict these changes, it is necessary to understand the magnitude and causes of variation in scleractinian coral diversity throughout their 240 million year history. The fossil record documents long periods of speciation in corals, interrupted repeatedly by events of mass extinction. Some of these events relate clearly to changes in global climate. Diversity in reef corals has increased since their last period of extinction at the end of the Cretaceous (65 My bp ), and is still rising. During the last 8 million years, the fragmentation of the once pantropical Tethys Sea separated corals into two major biogeographical provinces. Periods of glaciation also have caused major changes in sea level and temperature. Accumulated evidence supports the theory that relative habitat area and changing patterns of oceanic circulation are mainly responsible for the two observed centres of recent coral diversity at the western tropical margins of the Atlantic and Pacific oceans. At predicted rates of climate change in the near future, coral reefs are likely to survive as an ecosystem. Increases in sea level may actually benefit corals and lead to regional increases in diversity if new habitat area on back reefs is opened to increased water circulation and thus coral dispersal. Rising temperature may cause higher rates of coral mortality and even local extinction in isolated, small populations such as those on oceanic islands. The effects of increases in ultraviolet radiation (UV) are largely unknown, but likely to be negative. UV may damage planktonic coral propagules in oceanic surface waters and thus decrease rates of gene flow between coral populations. This may result in increased local extinctions, again with the strongest impact on widely separated reefs with small coral populations. The largest threats to coral diversity are regional anthropogenic impacts, which may interact with global climate change to exacerbate rates of local species extinctions. Centres of high reef coral diversity coincide with human population centres in south-east Asia and the Caribbean, and thus the greatest potential for species loss lies in these geographical areas.     

The influence of colony size and coral health on the occupation of coral-associated gobies (Pisces: Gobiidae)

Fishes of the genus Gobiodon are habitat specialists by their association with Acropora corals. Little is known about the parameters that define host coral quality for these fishes, in particular their breeding pairs. Data were collected in the northern Red Sea using 10 × 1-m belt transects in different reefs and zones. Gobiid density was highly correlated with coral density over all sites and zones, and the more specialized goby species preferred coral species that are less vulnerable to environmental stress. Moreover, the occupation rate of corals by goby breeding pairs significantly increased with colony size and decreased with partial mortality of colonies. Logistic regression showed that both coral size (being most important) and partial mortality are key factors influencing the occupation by breeding pairs. This study provides the first evidence that breeding pairs of coral-associated gobiids have more advanced habitat requirements than con-specifics in other social states. As coral reefs are threatened worldwide and habitat loss and degradation increase, this information will help predict the potential effects on those reef fishes obligatorily associated with live corals.     

Relationships between habitat structure and fish communities on coral

The influence of habitat structure on reef-fish communities at Bar Reef Marine Sanctuary, Sri Lanka, was investigated. The relationship between habitat characteristics and the distribution and abundance of 135 species of fishes was examined on two reef types: coral and sandstone reefs. Results suggested that the reef-fish communities were strongly influenced by various aspects of reef structure. However, relationships between habitat variables and fish communities structure, varied between the two reef types. Fish species diversity was correlated with a number of habitat variables on the sandstone reefs, although structural complexity seemed to play the dominant role. There were no correlations between habitat structure and fish diversity on the coral reefs. Total abundance was not related to any one habitat parameter on either reef type. However, abundances of some species, families and trophic groups were correlated with habitat features. These specific correlations were commonly related to food or shelter availability. For example, coral feeders were correlated with live coral cover, and pomacentrid species, which used branching corals for protection, showed a significant relationship with the density of Acropora colonies. This shows that a summary statistic such as total abundance may hide important information. Effects of habitat structure on the distribution patterns of the fish communities was further investigated using multi-dimensional scaling ordination (MDS) and the RELATE-procedure. With the MDS ordinations for both habitat and fish-community composition it was possible to show that the multivariate pattern between the two ecological components was clearly correlated.     

Symbiotic crabs maintain coral health by clearing sediments

Stony corals are the foundation of coral reef ecosystems and form associations with other reef species. Many of these associations may be ecologically important and play a role in maintaining the health and diversity of reef systems, rendering it critical to understand the influence of symbiotic organisms in mediating responses to perturbation. This study demonstrates the importance of an association with trapeziid crabs in reducing adverse effects of sediments deposited on corals. In a field experiment, mortality rates of two species of branching corals were significantly lowered by the presence of crabs. All outplanted corals with crabs survived whereas 45–80% of corals without crabs died within a month. For surviving corals that lacked crabs, growth was slower and tissue bleaching and sediment load were higher. Laboratory experiments revealed that corals with crabs shed substantially more of the sediments deposited on coral surfaces, but also that crabs were most effective at removing grain sizes that were most damaging to coral tissues. The mechanism underlying this symbiotic relationship has not been recognized previously, and its role in maintaining coral health is likely to become even more critical as reefs worldwide experience increasing sedimentation.     

Carbonic anhydrases in anthozoan corals—A review

Coral reefs are among the most biologically diverse and economically important ecosystems on the planet. The deposition of massive calcium carbonate skeletons (biomineralization or calcification) by scleractinian corals forms the coral reef framework/architecture that serves as habitat for a large diversity of organisms. This process would not be possible without the intimate symbiosis between corals and photosynthetic dinoflagellates, commonly called zooxanthellae. Carbonic anhydrases play major roles in those two essential processes of coral’s physiology: they are involved in the carbon supply for calcium carbonate precipitation as well as in carbon-concentrating mechanisms for symbiont photosynthesis. Here, we review the current understanding of diversity and function of carbonic anhydrases in corals and discuss the perspective of theses enzymes as a key to understanding impacts of environmental changes on coral reefs.     

Importance of live coral habitat for reef fishes

Live corals are the key habitat forming organisms on coral reefs, contributing to both biological and physical structure. Understanding the importance of corals for reef fishes is, however, restricted to a few key families of fishes, whereas it is likely that a vast number of fish species will be adversely affected by the loss of live corals. This study used data from published literature together with independent field based surveys to quantify the range of reef fish species that use live coral habitats. A total of 320 species from 39 families use live coral habitats, accounting for approximately 8 % of all reef fishes. Many of the fishes reported to use live corals are from the families Pomacentridae (68 spp.) and Gobiidae (44 spp.) and most (66 %) are either planktivores or omnivores. 126 species of fish associate with corals as juveniles, although many of these fishes have no apparent affiliation with coral as adults, suggesting an ontogenetic shift in coral reliance. Collectively, reef fishes have been reported to use at least 93 species of coral, mainly from the genus Acropora and Porities and associate predominantly with branching growth forms. Some fish associate with a single coral species, whilst others can be found on more than 20 different species of coral indicating there is considerable variation in habitat specialisation among coral associated fish species. The large number of fishes that rely on coral highlights that habitat degradation and coral loss will have significant consequences for biodiversity and productivity of reef fish assemblages.     

Habitat associations of juvenile fish at Ningaloo Reef, Western Australia: the importance of coral and algae

Habitat specificity plays a pivotal role in forming community patterns in coral reef fishes, yet considerable uncertainty remains as to the extent of this selectivity, particularly among newly settled recruits. Here we quantified habitat specificity of juvenile coral reef fish at three ecological levels; algal meadows vs. coral reefs, live vs. dead coral and among different coral morphologies. In total, 6979 individuals from 11 families and 56 species were censused along Ningaloo Reef, Western Australia. Juvenile fishes exhibited divergence in habitat use and specialization among species and at all study scales. Despite the close proximity of coral reef and algal meadows (10's of metres) 25 species were unique to coral reef habitats, and seven to algal meadows. Of the seven unique to algal meadows, several species are known to occupy coral reef habitat as adults, suggesting possible ontogenetic shifts in habitat use. Selectivity between live and dead coral was found to be species-specific. In particular, juvenile scarids were found predominantly on the skeletons of dead coral whereas many damsel and butterfly fishes were closely associated with live coral habitat. Among the coral dependent species, coral morphology played a key role in juvenile distribution. Corymbose corals supported a disproportionate number of coral species and individuals relative to their availability, whereas less complex shapes (i.e. massive & encrusting) were rarely used by juvenile fish. Habitat specialisation by juvenile species of ecological and fisheries importance, for a variety of habitat types, argues strongly for the careful conservation and management of multiple habitat types within marine parks, and indicates that the current emphasis on planning conservation using representative habitat areas is warranted. Furthermore, the close association of many juvenile fish with corals susceptible to climate change related disturbances suggests that identifying and protecting reefs resilient to this should be a conservation priority.     

Phase shift to algal dominated communities at mesophotic depths associated with lionfish (<Emphasis Type="Italic">Pterois volitans</Emphasis>) invasion on a Bahamian coral reef

Mesophotic coral reefs (30–150 m) have been assumed to be physically and biologically connected to their shallow-water counterparts, and thus may serve as refugia for important taxonomic groups such as corals, sponges, and fish. The recent invasion of the Indo–Pacific lionfish (Pterois volitans) onto shallow reefs of the Caribbean and Bahamas has had significant, negative, effects on shallow coral reef fish populations. In the Bahamas, lionfish have extended their habitat range into mesophotic depths down to 91 m where they have reduced the diversity of several important fish guilds, including herbivores. A phase shift to an algal dominated (>50% benthic cover) community occurred simultaneously with the loss of herbivores to a depth of 61 m and caused a significant decline in corals and sponges at mesophotic depths. The effects of this invasive lionfish on mesophotic coral reefs and the subsequent changes in benthic community structure could not be explained by coral bleaching, overfishing, hurricanes, or disease independently or in combination. The significant ecological effects of the lionfish invasion into mesophotic depths of coral reefs casts doubt on whether these communities have the resilience to recover themselves or contribute to the recovery of their shallow water counterparts as refugia for key coral reef taxa.     

Mechanical vulnerability explains size‐dependent mortality of reef corals

Understanding life history and demographic variation among species within communities is a central ecological goal. Mortality schedules are especially important in ecosystems where disturbance plays a major role in structuring communities, such as coral reefs. Here, we test whether a trait‐based, mechanistic model of mechanical vulnerability in corals can explain mortality schedules. Specifically, we ask whether species that become increasingly vulnerable to hydrodynamic dislodgment as they grow have bathtub‐shaped mortality curves, whereas species that remain mechanically stable have decreasing mortality rates with size, as predicted by classical life history theory for reef corals. We find that size‐dependent mortality is highly consistent between species with the same growth form and that the shape of size‐dependent mortality for each growth form can be explained by mechanical vulnerability. Our findings highlight the feasibility of predicting assemblage‐scale mortality patterns on coral reefs with trait‐based approaches.     

Coral communities in the northern part of the Gulf of Tonkin, south China Sea

The reefs and benthic communities of 15 islands in Ha Long Bay, listed by UNESCO as a world heritage site, have been studied using scuba. Two reef types have been distinguished: structured and unstructured reefs. The changes in the composition and structure of reef-building corals takes place in these reefs under conditions of heavy silting in the surrounding waters. Poritids that primarily form the reef carcass become dominant scleractinians. More than 250 species of macrobenthos are found; the greatest species diversity was recorded in the reefs of Bo Hung and Kong Do islands, which we recommend be organized into a national nature reserve.     

Cleaner fish drives local fish diversity on coral reefs

Coral reefs are one of the most diverse habitats in the world, yet our understanding of the processes affecting their biodiversity is limited. At the local scale, cleaner fish are thought to have a disproportionate effect, in relation to their abundance and size, on the activity of many other fish species, but confirmation of this species' effect on local fish diversity has proved elusive. The cleaner fish Labroides dimidiatus has major effects on fish activity patterns and may indirectly affect fish demography through the removal of large numbers of parasites. Here we show that small reefs where L. dimidiatus had been experimentally excluded for 18 months had half the species diversity of fish and one-fourth the abundance of individuals. Only fish that move among reefs, however, were affected. These fish include large species that themselves can affect other reef organisms. In contrast, the distribution of resident fish was not affected by cleaner fish. Thus, many fish appear to choose reefs based on the presence of cleaner fish. Our findings indicate that a single small and not very abundant fish has a strong influence on the movement patterns, habitat choice, activity, and local diversity and abundance of a wide variety of reef fish species.     

Reef Fishes in Biodiversity Hotspots Are at Greatest Risk from Loss of Coral Species

Coral reef ecosystems are under a variety of threats from global change and anthropogenic disturbances that are reducing the number and type of coral species on reefs. Coral reefs support upwards of one third of all marine species of fish, so the loss of coral habitat may have substantial consequences to local fish diversity. We posit that the effects of habitat degradation will be most severe in coral regions with highest biodiversity of fishes due to greater specialization by fishes for particular coral habitats. Our novel approach to this important but untested hypothesis was to conduct the same field experiment at three geographic locations across the Indo-Pacific biodiversity gradient (Papua New Guinea; Great Barrier Reef, Australia; French Polynesia). Specifically, we experimentally explored whether the response of local fish communities to identical changes in diversity of habitat-providing corals was independent of the size of the regional species pool of fishes. We found that the proportional reduction (sensitivity) in fish biodiversity to loss of coral diversity was greater for regions with larger background species pools, reflecting variation in the degree of habitat specialization of fishes across the Indo-Pacific diversity gradient. This result implies that habitat-associated fish in diversity hotspots are at greater risk of local extinction to a given loss of habitat diversity compared to regions with lower species richness. This mechanism, related to the positive relationship between habitat specialization and regional biodiversity, and the elevated extinction risk this poses for biodiversity hotspots, may apply to species in other types of ecosystems.     

ECOLOGY AND MORPHOLOGY OF RECENT CORAL REEFS

1. The classical ‘coral reef problem’ concerned the geological relationships of reefs as major topographical features; modern coral studies consider reefs both as complex biological systems of high productivity and as geological structures forming a framework for and being modified by coral growth. 2. Deep borings in reefs have conclusively confirmed the general arguments of Darwin, that oceanic reefs developed by progressive subsidence of their foundations. Darwin failed to take account of Pleistocene changes in sea level and their effect on the present surface features of reefs. Daly's alternative ‘glacial control theory’ was based on false assumptions concerning marine erosion rates during glacial periods, but if sea level during the Holocene was higher than at present, as Daly also supposed, the effects on reef features would be profound. 3. Reefs are complex biological systems in tropical seas, dominated by scleractinian corals. Coral faunas are larger and more diverse in the Indo-Pacific than in the Atlantic. Hermatypic corals are restricted to shallow water by the light requirements of their symbiotic algae, but temperature is a major control of worldwide distributions. Temperature, salinity and sediment tolerances of corals are wider than formerly supposed, and corals can survive brief emersion except when it coincides with heavy rainfall. Water turbulence is an important ecological control, but difficult to measure. 4. The trophic status of corals is still unclear, but in spite of their anatomical and physiological specialization as carnivores it is likely that they derive some nutrient substances from zooxanthellae. Suggestions that filamentous algae in coral heads play a major part in the economy of the corals have not been supported by later work, but biomass pyramids constructed on the basis by Odum and Odum remain the only ones available. Most reefs are apparently autotrophic, with 1500–3500 g. Carbon being fixed per m.² per year. 5. Few animals eat corals, which may account for their success. Important predators are fish and the echinoderm Acanthaster. Quantitative estimates of biogenic erosion of organic skeletons on reefs are high. Fish affect not only corals but other invertebrates, algae and marine phanerogams. 6. Corals may be killed by ‘dark water’, intense rain or river floodwaters, earth movements, human interference and especially hurricanes. Reef recovery after hurricanes may take 10–20 years. 7. In addition to fringing, barrier and atoll reefs, intermediate types are recognised. The main types may consist of linear reefs or faros. Smaller lagoon reefs include pinnacles, patches and platforms, and submerged knolls. Complex cellular or mesh reef patterns are also found. 8. Reefs are conspicuously zoned, both laterally in response to changing exposure to waves to form windward and leeward reefs, and transversely, as a result of steep environmental gradients across reef flats from sea to lagoon. Topographic and ecological zones may be characterized by particular coral species, but these vary widely from reef to reef. A major distinction can be made between reefs with and without algal ridges, which are common on open-ocean trade-wind reefs, in the Indo-Pacific, but are absent on Caribbean reefs and on Indo-Pacific reefs in more sheltered waters. gorgonians are common on Caribbean reefs, alcyonaceans in the Indo-Pacific. 9. Much of the difficulty in comparing reefs stems from the lack of uniformity in surveying methods. Problems of describing the complex three-dimensional patterns of organisms on reefs have yet to be solved, and hence little progress has been made in explanation of these patterns. Explanation in terms of simple environmental controls is inadequate. 10. Understanding the distribution of corals is made more difficult both by taxo-nomic problems and by the plasticity of growth form in different situations. 11. Growth of corals and reefs may be estimated by measuring the growth of individual colonies, measuring rates of calcium carbonate deposition in the skeleton, measuring topographic change on the reef and deducing net rates of reef growth from geological evidence. Massive corals may increase in diameter by 1 cm./year, branches of branching corals may increase in length by 10 cm./year. Study of deposition rates shows variation within colonies, between species, in light and dark, and seasonally. Rates of reef growth extrapolated from colony measurements reach 2–5 cm./year, and contrast with figures as low as 0–02 cm/year averaged over 70 million years from borehole data. Both colony growth rates and geological data suggest worldwide variations in rates of reef growth. 12. In spite of clear evidence of long-continued subsidence, present surface features of reefs, often only thinly veneered by modern corals, have been much affected by recent sea level fluctuations. Many slightly raised reefs at 2–10 m. above sea level date at 90–160 thousand years B.P.; there is evidence for a sea level at about the present level at 30–35 thousand years B.P.; and controversy continues over whether sea level has stood higher than the present at any time since the last sea level rise began about 20,000 years ago. Evidence from many reefs suggests a slightly higher sea level in the last 4000 years, but on other reefs such evidence is lacking. 13. Several reef features (submerged terraces, groove-spur systems, algal ridge, reef flat, reef blocks and reef islands) have been interpreted either as relict features dating from a higher sea level in the last 5000 years, or contemporary features developed in response to present processes. In some cases the evidence is equivocal; in others it is clear that diverse features are being grouped together under the same name. If such features are referable to a higher sea level, this may have been of last Interglacial or even Interstadial age rather than Holocene. 14. A reef consists of a rigid framework defining several major depositional environments within and around it. Sediments are of biological, mainly skeletal origin, except in unusual environments such as the Bahama Banks. The characteristics of sediments derived from organisms depend partly on the breakdown patterns of particular skeletons, partly on transportation and sorting processes. Fine sediments may be either detrital, or physicochemical precipitates. 15. Organisms affect sediments after deposition, by disturbance, transportation and probably comminution. Fish and holothurians have been studied in detail. 16. While new theories of coral reefs are proposed from time to time, the need is less for new theories than for standardised procedures to ensure comparability of reef studies and the identification of variations in reefs both on local and regional scales. While reefs as biological systems adjust relatively rapidly to changes, reefs as geological systems adjust much more slowly. Because of the magnitude and recency of Pleistocene fluctuations in sea level, many biological features of reefs are out of phase with inherited geological features, and this had led to much controversy.     

Limited habitat and conservation value of a young artificial reef

Loss and degradation of natural habitats and their biodiversity may, arguably, be mitigated or compensated through the creation of human-engineered habitats: the underlying conservation tenet is that these artificial habitats compensate for diminished diversity caused by human impacts at local or regional scales. This approach is widely used in the sea by purposefully scuttling ships to create artificial reefs, but its performance as a conservation tool is seldom critically examined in these situations. Here we test if the diversity of sessile invertebrate assemblages on a large, but young (3 years), artificial reef, created by sinking a 133 m long battle ship off Eastern Australia, can mimic that of nearby natural reefs. We use this system as a model to test whether this artificial reef can form compensatory habitat of comparable quality and levels of biodiversity. Our assessment is based on the abundance, species richness, and species composition of sessile invertebrate assemblages, including corals. Despite some signs that temporal trajectories of ecological metrics, such as cover, began to approach natural conditions after 3 years, the ecological structure of sessile invertebrate assemblages on this young wreck remained fundamentally different from those on nearby natural reefs. In particular, large, long-lived corals were abundant on natural rocky reefs, but were rare and covered little area on the young wreck. These data demonstrate that when trajectories to community convergence with natural habitats are prolonged, as may be the case here, any compensatory effects of artificial habitats will have a considerable time lag. Such lags have implications for appraising the conservation value of wrecks and artificial reefs, and they emphasize the need to explicitly acknowledge temporal dynamics when using artificial habitats as complementary conservation tools to augment larger conservation efforts on natural systems.     

Effects of ultraviolet radiation on corals and other coral reef organisms

The discovery of the importance of solar ultraviolet radiation (UVR) as a factor affecting the biology of coral reefs dates only to about 1980. Interest has heightened during the past five years owing to the demonstration of loss of stratospheric ozone through human activities. We have only begun to document gross, qualitative effects of UVR on coral reef organisms, usually in experiments comparing the biological response to the presence or absence of UVR through the use of UV-cutoff filters, or to varying levels of UVR in transplantation studies. Most such studies have not distinguished between the effects of UVA (320–400 nm) and those of UVB (290–320 nm), although in the context of global change involving stratospheric ozone loss, it is the latter wavelengths that are relevant. To date we have been addressing physiological and ecological questions, not yet attempting to evaluate quantitatively the impact of forecast increases in solar UVB penetration. Interacting and synergistic effects of UVR with increased temperature, pollutants, sedimentation, visible light, etc. have scarcely been studied but will be essential to understanding and predicting the fate of coral reefs under conditions of global change. Here we comprehensively review the effects of UVR on corals and other reef macroorganisms, mindful that although much is known of proximal effects, little of this knowledge is directly useful in making long-term predictions regarding the health of coral reefs. We conclude that even small anthropogenic increases in UVB levels will have sublethal physiological manifestations in corals and other reef organisms, but that this will have relatively small impact on the distribution of reef corals and coral reefs, perhaps affecting their minimum depths of occurrence.     

Herbivory versus corallivory: are parrotfish good or bad for Caribbean coral reefs?

With coral cover in decline on many Caribbean reefs, any process of coral mortality is of potential concern. While sparisomid parrotfishes are major grazers of Caribbean reefs and help control algal blooms, the fact that they also undertake corallivory has prompted some to question the rationale for their conservation. Here the weight of evidence for beneficial effects of parrotfishes, in terms of reducing algal cover and facilitating demographic processes in corals, and the deleterious effects of parrotfishes in terms of causing coral mortality and chronic stress, are reviewed. While elevated parrotfish density will likely increase the predation rate upon juvenile corals, the net effect appears to be positive in enhancing coral recruitment through removal of macroalgal competitors. Parrotfish corallivory can cause modest partial colony mortality in the most intensively grazed species of Montastraea but the generation and healing of bite scars appear to be in near equilibrium, even when coral cover is low. Whole colony mortality in adult corals can lead to complete exclusion of some delicate, lagoonal species of Porites from forereef environments but is only reported for one reef species (Porites astreoides), for one habitat (backreef), and with uncertain incidence (though likely <<10%). No deleterious effects of predation on coral growth or fecundity have been reported, though recovery of zooxanthellae after bleaching events may be retarded. The balance of evidence to date finds strong support for the herbivory role of parrotfishes in facilitating coral recruitment, growth, and fecundity. In contrast, no net deleterious effects of corallivory have been reported for reef corals. Corallivory is unlikely to constrain overall coral cover but contraints upon dwindling populations of the Montastraea annularis species complex are feasible and the role of parrotfishes as a vector of coral disease requires evaluation. However, any assertion that conservation practices should guard against protecting corallivorous parrotfishes appears to be unwarranted at this stage.     

Effects of eutrophication on bioeroding sponge communities with the description of new West Indian sponges, Cliona spp. (Porifera: Hadromerida: Clionidae)

Abstract. Pieces of coral rubble ( Porites porites ), collected from across 3 fringing reefs that lie along a eutrophication gradient, were examined for the presence of clionid sponges. A similarity analysis of species composition showed that reef zone had less effect on clionid community composition than did other factors affecting the reefs as a whole. Except on the back reef, the Zones, distances, and depths within the reefs had no significant influence on the number of clionid invasions. Reef comparisons demonstrated that clionid abundance increased with increasing eutrophication. Clionids were found in 41% of the pieces collected from the most eutrophic site vs. 24% from the least eutrophic. Because clionids are the principal bio-eroders of coral reefs, any increase in their abundance will likely result in greater bioerosion rates. The mean abundance of Type 3 corals (in which fragmentation is the primary mode of propagation) is positively related to the frequency of boring sponge invasion. suggesting that increased bioerosion may be partly responsible for community shifts toward Type 3 corals in polluted waters. Cliona cf. vastifica , found for the first time in Barbados, flourishes on the most eutrophic reef and may become an important bioeroder under the highly eutrophic conditions that have begun to plague West Indian reefs. Two new species of Cliona (Porifera: Hadro-merida: Clionidae) are described.     

Comments on coral reef regeneration, bioerosion, biogeography, and chemical ecology: future directions

This paper represents a brief review of three processes operating on coral reefs and the results of studies of a fourth process, and how those results may be applied to the benefit of humankind. The areas are reef regeneration processes; bioerosion; dispersal, recruitment and biogeography of corals; and chemical ecology and natural products chemistry of reef organisms. Possible future directions for research will also be considered in each area. Regarding reef regeneration processes, coral reefs are degrading rapidly on a global scale due to over-fishing, fishing techniques causing habitat destruction, deforestation, mass mortalities of key reef species, nutrient enrichment and sedimentation. Seeding of reefs with the larvae of corals and other key reef organisms, such as echinoids, may help to promote and enhance reef regeneration in the future. Such techniques will be made possible by studies of the embryology, larval ecology, dispersal and recruitment processes, and related local physical oceanographic processes. Regarding bioerosion, both internal and external bioerosion are affected by grazers and predators. Bioerosion is also affected by nutrient enrichment, as shown through correlative studies (Great Barrier Reef) and studies of opportunity (Kaneohe Bay). Ongoing experiments such as ENCORE will help to answer questions about the role of dissolved nutrients in enhancing internal bioerosion. Questions still remain, however, regarding the role of particulates in promoting internal bioerosion and the resultant weakening of and negative growth in the reef framework. Regarding dispersal, recruitment and the biogeography of corals, it is now known that most species of coral reproduce via broadcasting, although there appear to be proportionally more brooders in the Caribbean than in the western Pacific. Differential extinctions in the western Pacific vs. the western Atlantic have contributed to the biogeographic distribution of corals we observe today and the concentric isoclines of species diversity in numerous reef organisms in the western Pacific. The role of reproductive mode in contributing to these patterns is, however, still not understood. Investigations into the roles of different larval longevities and reproductive modes may help us answer questions regarding their differential distribution and the potential effects of major perturbations such as global warming on future distributions. With respect to the chemical ecology of alcyonacean octocorals (soft corals), four functions have been determined thus far for secondary metabolites in this group, anti-predation, anti-competition (allelopathy), anti-fouling, and enhancement of reproductive success. Investigations of alcyonacean reproduction has revealed that it may be necessary for several secondary metabolites to be present simultaneously before a function may be realized or fully manifested. This raises questions regarding the manner in which novel compounds are tested by medical laboratories for bioactivity using a single compound. Simultaneously testing of multiple compounds derived from a single organism may be necessary in the future to reveal potential valuable synergistic bioactivity. Also, some novel secondary metabolites may have other valuable commercial applications, as is the case with the UV-absorbing compounds of corals and other reef organisms found on the Great Barrier Reef. In order to avoid overlooking medically or commercially valuable functions of these compounds, broader testing may be necessary.     

Overgrowth and killing of corals by the brown alga Lobophora hederacea (Dictyotales,Phaeophyceae) on healthy reefs in New Caledonia: A new case of the epizoism syndrome

Coral reef degradation is often associated with regime shifts from coral‐ to macroalgal‐dominated reefs. These shifts demonstrate that under certain conditions (e.g. coral mortality, decrease in herbivory, increased nutrients supply) some macroalgae may overgrow corals. The outcome of the competition is dependent on algal aggressiveness and the coral susceptibility. In undisturbed reefs, herbivore grazing is regulating macroalgal cover, thus preventing the latter from overgrowing corals. However, some macroalgae have evolved strategies not only to outcompete corals but also to escape herbivory to some extent, allowing overgrowth of some coral species in undisturbed reefs. Epizoism represents one of those successful strategies, and has been previously documented with red algae, cyanobacteria and Lobophora variegata (Dictyotales, Phaeophyceae). Here we report a new case of epizoism leading to coral mortality, involving a recently described species of Lobophora, L. hederacea, overgrowing the coral Seriatopora caliendrum (Pocilloporidae) in undisturbed reefs in New Caledonia.     

Comparison of coral diversity between big and small atolls: a case study of Yongle atoll and Lingyang reef,Xisha Islands,central of South China Sea

The South China Sea (SCS) includes large areas of extensive coral reef development but its reefs are still poorly known. Yongle atoll is the biggest typical atoll in the Xisha Islands, central of SCS. Lingyang Reef is an isolated small atoll within the whole big Yongle atoll. A total of 144 and 119 coral species were recorded at big Yongle atoll and small Lingyang Reef, respectively. The real coral richness might be higher because species accumulation curve did not saturate. The coral diversity pattern was similar between big Yongle atoll and small Lingyang Reef. Coral communities fell into three clusters, consistent with their habitats on reef slope, reef flat and lagoon slope. The highest coral diversity was observed on reef slopes and the lowest coral diversity was found on lagoon slope. Genera richness was a better proxy for representing coral species diversity on both the big and small atoll but percent live coral cover was not a robust proxy on the small atoll, which only explained 24% of species diversity. This study demonstrated high coral diversity with consistent pattern along habitat types, as has been shown from many other reefs. While far from exhaustive, the study allows first glimpses on how much biodiversity is contained on SCS coral reefs, and hopes to give an impetus to their conservation. The study also suggests that simplified surveys at a small scale and the use of genera richness as an effective proxy for overall diversity can indeed provide important information to rapidly monitor and evaluate the coral diversity in remote locations.