Environmental impacts of dredging and other sediment disturbances on corals: A review

A review of published literature on the sensitivity of corals to turbidity and sedimentation is presented, with an emphasis on the effects of dredging. The risks and severity of impact from dredging (and other sediment disturbances) on corals are primarily related to the intensity, duration and frequency of exposure to increased turbidity and sedimentation. The sensitivity of a coral reef to dredging impacts and its ability to recover depend on the antecedent ecological conditions of the reef, its resilience and the ambient conditions normally experienced. Effects of sediment stress have so far been investigated in 89 coral species (～10% of all known reef-building corals). Results of these investigations have provided a generic understanding of tolerance levels, response mechanisms, adaptations and threshold levels of corals to the effects of natural and anthropogenic sediment disturbances. Coral polyps undergo stress from high suspended-sediment concentrations and the subsequent effects on light attenuation which affect their algal symbionts. Minimum light requirements of corals range from <1% to as much as 60% of surface irradiance. Reported tolerance limits of coral reef systems for chronic suspended-sediment concentrations range from <10mgL(-1) in pristine offshore reef areas to >100mgL(-1) in marginal nearshore reefs. Some individual coral species can tolerate short-term exposure (days) to suspended-sediment concentrations as high as 1000mgL(-1) while others show mortality after exposure (weeks) to concentrations as low as 30mgL(-1). The duration that corals can survive high turbidities ranges from several days (sensitive species) to at least 5-6weeks (tolerant species). Increased sedimentation can cause smothering and burial of coral polyps, shading, tissue necrosis and population explosions of bacteria in coral mucus. Fine sediments tend to have greater effects on corals than coarse sediments. Turbidity and sedimentation also reduce the recruitment, survival and settlement of coral larvae. Maximum sedimentation rates that can be tolerated by different corals range from <10mgcm(-2)d(-1) to >400mgcm(-2)d(-1). The durations that corals can survive high sedimentation rates range from <24h for sensitive species to a few weeks (>4weeks of high sedimentation or >14days complete burial) for very tolerant species. Hypotheses to explain substantial differences in sensitivity between different coral species include the growth form of coral colonies and the size of the coral polyp or calyx. The validity of these hypotheses was tested on the basis of 77 published studies on the effects of turbidity and sedimentation on 89 coral species. The results of this analysis reveal a significant relationship of coral sensitivity to turbidity and sedimentation with growth form, but not with calyx size. Some of the variation in sensitivities reported in the literature may have been caused by differences in the type and particle size of sediments applied in experiments. The ability of many corals (in varying degrees) to actively reject sediment through polyp inflation, mucus production, ciliary and tentacular action (at considerable energetic cost), as well as intraspecific morphological variation and the mobility of free-living mushroom corals, further contribute to the observed differences. Given the wide range of sensitivity levels among coral species and in baseline water quality conditions among reefs, meaningful criteria to limit the extent and turbidity of dredging plumes and their effects on corals will always require site-specific evaluations, taking into account the species assemblage present at the site and the natural variability of local background turbidity and sedimentation.     

Is proximity to land-based sources of coral stressors an appropriate measure of risk to coral reefs? An example from the Florida Reef Tract

Localized declines in coral condition are commonly linked to land-based sources of stressors that influence gradients of water quality, and the distance to sources of stressors is commonly used as a proxy for predicting the vulnerability and future status of reef resources. In this study, we evaluated explicitly whether proximity to shore and connections to coastal bays, two measures of potential land-based sources of disturbance, influence coral community and population structure, and the abundance, distribution, and condition of corals within patch reefs of the Florida Reef Tract. In the Florida Keys, long-term monitoring has documented significant differences in water quality along a cross-shelf gradient. Inshore habitats exhibit higher levels of nutrients (DIN and TP), TOC, turbidity, and light attenuation, and these levels decrease with increasing distance from shore and connections to tidal bays. In clear contrast to these patterns of water quality, corals on inshore patch reefs exhibited significantly higher coral cover, higher growth rates, and lower partial mortality rates than those documented in similar offshore habitats. Coral recruitment rates did not differ between inshore and offshore habitats. Corals on patch reefs closest to shore had well-spread population structures numerically dominated by intermediate to large colonies, while offshore populations showed narrower size-distributions that become increasingly positively skewed. Differences in size-structure of coral populations were attributed to faster growth and lower rates of partial mortality at inshore habitats. While the underlying causes for the favorable condition of inshore coral communities are not yet known, we hypothesize that the ability of corals to shift their trophic mode under adverse environmental conditions may be partly responsible for the observed patterns, as shown in other reef systems. This study, based on data collected from a uniform reef habitat type and coral species with diverse life-history and stress-response patterns from a heavily exploited reef system, showed that proximity to potential sources of stressors may not always prove an adequate proxy for assigning potential risks to reef health, and that hypothesized patterns of coral cover, population size-structure, growth, and mortality are not always directly related to water quality gradients.     

Nutrient enrichment caused by in situ fish farms at Eilat, Red Sea is detrimental to coral reproduction

Recent studies report conflicting results concerning the effects of eutrophication on coral reproduction. The present study examines reproductive effort in the brooding coral Stylophora pistillata exposed to chronic eutrophication caused by in situ fish cages (FC) in the northern Gulf of Eilat (Aqaba). Histological studies of 20 S. pistillata colonies transplanted to each of two study sites, one close to the nutrient enriched FC site and the other at a reference site (IUI), 8 km southwest of the FC site, show that, overall, corals from the FC site have a significantly higher percentage of polyps containing oocytes and testes than corals from the IUI site. However, average oocyte size and the percentage of oocytes reaching the size at which fertilization occurs (i.e., >200 microm) were both significantly greater in colonies at the IUI site compared to the FC site. As the reproductive season progressed, colonies at the IUI site exhibited a decrease in the percentage of polyps containing oocytes, concomitant with an increase in the number of polyps containing planulae, indicating successful development of oocytes into planulae. In contrast, in colonies at the FC site oocyte numbers were greatest at the end of the reproductive season, and overall, numbers of planulae were significantly lower compared with the IUI colonies, suggesting relative failure of oocyte maturation, fertilization and ensuing larval development. The significantly higher lipid content found during the reproduction season in IUI colonies compared with FC colonies corroborates this assertion. This data strongly suggest that nutrients released from the fish farms have adverse effects on successful production of larvae of S. pistillata. In view of the recent severe deterioration of the coral reefs of Eilat and their present critical state of health, the only chance for their renewal is the use of immediate, prudent and rational protection measures against all man-made perturbations.     

Effects of coal contamination on early life history processes of a reef-building coral,Acropora tenuis

Successful reproduction and larval dispersal are important for the persistence of marine invertebrate populations, and these early life history processes can be sensitive to marine pollution. Coal is emerging as a contaminant of interest due to the proximity of ports and shipping lanes to coral reefs. To assess the potential hazard of this contaminant, gametes, newly developed embryos, larvae and juveniles of the coral Acropora tenuis were exposed to a range of coal leachate, suspended coal, and coal smothering treatments. Fertilisation was the most sensitive reproductive process tested. Embryo survivorship decreased with increasing suspended coal concentrations and exposure duration, effects on larval settlement varied between treatments, while effects on juvenile survivorship were minimal. Leachate exposures had negligible effects on fertilisation and larval settlement. These results indicate that coral recruitment could be affected by spills that produce plumes of suspended coal particles which interact with gametes and embryos soon after spawning.     

Inhibition of Fertilization and Larval Metamorphosis of the Coral Acropora millepora (Ehrenberg, 1834) by Petroleum Products

Accidental oil spills from ships or rigs and inputs of effluent such as production formation water (PFW) are key perceived threats to tropical biota from industry activities. Scleractinian corals are an important functional component of tropical reefs and the abundance, diversity and resilience of coral communities can be used as an indicator of ecosystem health. In this paper, we report the effects of petroleum products, including water accommodated fractions (WAF) of crude oil, PFW and dispersant (Corexit 9527), on fertilization and larval metamorphosis of the widespread scleractinian coral, Acropora millepora (Ehrenberg, 1834) in laboratory-based assays. At 20% v/v PFW fertilization was inhibited by 25%. This concentration was equivalent 0.0721 mg l⁻¹ total hydrocarbon (THC). In contrast, larval metamorphosis was more sensitive to this effluent, with 98% metamorphosis inhibited at the same concentration. Crude oil WAF did not inhibit fertilization of gametes until dispersant was introduced. Dispersed oil was slightly more toxic to fertilization than dispersant alone, suggesting toxicity to that event may be additive. The minimum concentration of dispersed oil which inhibited fertilization was 0.0325 mg l⁻¹ THC. Larval metamorphosis was more sensitive than fertilization to crude oil. Although crude oil and dispersant inhibited larval metamorphosis individually, this toxicity was magnified when larvae were exposed to combinations of both. Crude oil inhibited metamorphosis at 0.0824 mg l⁻¹ THC and at 0.0325 mg l⁻¹ THC when dispersed in 10% v/v (dispersant/oil). Management of petroleum-related risks to spawning corals should consider not only the occurrence of the annual coral spawning event, but also the subsequent 1–3-week period during which most larval metamorphosis and recruitment occur.     

Coral recruitment to the reefs of Eilat, Red Sea: temporal and spatial variation, and possible effects of anthropogenic disturbances

The accelerating deterioration of the coral reefs of Eilat has raised debate over the exact causes and how they affect the reefs. The hypothesis of the present study was that a low recruitment rate of reef-building coral species may play an important role in the decline of the Eilat reefs. Our goal was to assess spatial and temporal recruitment patterns in Eilat, focusing on examining the possible impact of human activities. The results of coral recruitment to 10 series of ceramic tiles on metal racks, revealed very low overall recruitment relative to other geographical regions. In addition, we found that recruitment rates and recruit survival were lowest at sites closest to the major eutrophication sources in Eilat. The low recruitment rates may be chronically too low to compensate for the elevated coral mortality rates of recent years. The significant differences between the present study and a similar study carried out during the same period using a different method, emphasize the crucial need for a standardized method for recruitment assessment in coral reefs worldwide.     

Partial mortality in massive reef corals as an indicator of sediment stress on coral reefs

Partial mortality and fission on colonies of four common massive coral species were examined at sites differing in their exposure to river sediments in St. Lucia, West Indies. Rates of partial mortality were higher close to the river mouths, where more sediments were deposited, than away from the rivers in two coral species. Frequency of fission showed no significant trend. The percent change in coral cover on reefs from 1995 to 1998 was negatively related to the rate of partial mortality estimated in 1998 in all species. This suggests that partial mortality rates could reflect longer-term temporal changes in coral communities. Similar conclusions could also be reached using a less precise measure and simply recording partial mortality on colonies as <50% and >/=50% dead tissue. We conclude that partial mortality in some species of massive reef corals, expressed as the amount of dead tissue per colony, could provide a rapid and effective means of detecting sediment stress on coral reefs.     

River discharge reduces reef coral diversity in Palau

Coral community structure is often governed by a suite of processes that are becoming increasingly influenced by land-use changes and related terrestrial discharges. We studied sites along a watershed gradient to examine both the physical environment and the associated biological communities. Transplanted corals showed no differences in growth rates and mortality along the watershed gradient. However, coral cover, coral richness, and coral colony density increased with increasing distance from the mouth of the bay. There was a negative relationship between coral cover and mean suspended solids concentration. Negative relationships were also found between terrigenous sedimentation rates and the richness of adult and juvenile corals. These results have major implications not only for Pacific islands but for all countries with reef systems downstream of rivers. Land development very often leads to increases in river runoff and suspended solids concentrations that reduce coral cover and coral diversity on adjacent reefs.     

Metals in Pocillopora damicornis exposed to tin smelter effluent

Metal concentrations in tissues and skeleton of the coral Pocillopora damicornis were compared from intertidal reefs adjacent to a tin smelter at Phuket, Thailand. Tissue concentrations of ten metals were significantly higher for corals collected at the tin smelter than at control reefs, 1 km distant. Elevated tissue metal concentrations were considered not to be derived from particulate material but probably from solution. Analyses of skeletal material revealed no differences in metal concentrations between study sites. The implications of elevated tissue metal concentrations are discussed in relation to reduced growth rates and low abundance of branching coral species at the reef adjacent to the tin smelter when compared to the control reefs.     

Deterioration Index (DI): a suggested criterion for assessing the health of coral communities

The extensive deterioration of coral reefs worldwide highlights the importance of creating efficient monitoring methods to best assess their state of health. At present, several suggested parameters serve such indicators. None of these, however, is well accepted as reliably representing reef community health. In the present study we examine a new approach based on the ratio between mortality and recruitment rates of branching corals, which we term 'Deterioration Index' (DI). It aims at providing a quantitative indication of the state of health of reef-building coral communities. The method was developed and tested on 16 coral communities on artificially laid rocks along the coast of Eilat, Red Sea (Gulf of Aqaba). In contrast to frequently used indices (i.e. mortality rate, abundance and species richness), which did not demonstrate a consistent result in comparing disturbed vs. undisturbed coral communities, the DI revealed significant differences between these communities. Our results suggest that the use of the DI may enable the detection of disturbed coral communities in one instance monitoring, where the other parameters had failed. The DI, therefore, may provide a comparable quantitative assessment of the deterioration process and its intensity in a coral community. We propose the DI approach as an efficient and applicable tool for coral reef monitoring.     

Modern microbialites and their environmental significance, Meiji reef atoll, Nansha (Spratly) Islands, South China Sea

Meiji (Mischief) coral atoll, in Nansha (Spratly) Islands, South China Sea, consists of an annular reef rim surrounding a central lagoon. On the atoll rim there are either protuberant 'motu' (small coral patch reefs on the rim of atoll) islets or lower sandy cays that contain modern microbialite deposits on the corals in pinnacles and surrounding bottoms of the atoll. Microbialites, including villiform, hairy, and thin spine growth forms, as well as gelatinous masses, mats and encrustation, developed on coral colonies and atoll rim sediments between 0 and 15 m deep-water settings. The microbialites were produced by natural populations of filamentous cyanobacteria and grew on (1) bulbous corals together with Acropora sp., (2) on massive colonies of Galaxea fascicularis, (3) on dead Montipora digitata, and (4) on dead Acropora teres, some hairy microbialite growing around broken coral branches. This study demonstrates that microbial carbonates are developed in coral reefs of South China Sea and indicates that microbial processes may be important in the construction of modern reef systems. The results have significance in the determination of nature and composition in microorganisms implied in the formation ancient microbialites, and permit evaluation of the importance of microbial deposits in mo-dern coral reefs and of 'microbialites' in biogeochemical cycles of modern coral reef systems. The re-sults also provide evidence of modern analogues for ancient microbialites in shallow-water settings, and combine with sedimentological studies of ancient microbialites to understand their controls.     

Long-term community changes on a high-latitude coral reef in the Greater St Lucia Wetland Park, South Africa

South African coral reefs are limited in size but, being marginal, provide a model for the study of many of the stresses to which these valuable systems are being subjected globally. Soft coral cover, comprising relatively few species, exceeds that of scleractinians over much of the reefs. The coral communities nevertheless attain a high biodiversity at this latitude on the East African coast. A long-term monitoring programme was initiated in 1993, entailing temperature logging and image analysis of high resolution photographs of fixed quadrats on representative reef. Sea temperatures rose by 0.15 degrees C p.a. at the site up to 2000 but have subsequently been decreasing by 0.07 degrees C p.a. Insignificant bleaching was encountered in the region during the 1998 El Nino Southern Oscillation (ENSO) event, unlike elsewhere in East Africa, but quantifiable bleaching occurred during an extended period of warming in 2000. Peak temperatures on the South African reefs thus appear to have attained the coral bleaching threshold. While this has resulted in relatively little bleaching thus far, the increased temperatures appear to have had a deleterious effect on coral recruitment success as other anthropogenic influences on the reefs are minimal. Recruitment success diminished remarkably up to 2004 but appears again to be improving. Throughout, the corals have also manifested changes in community structure, involving an increase in hard coral cover and reduction in that of soft corals, resulting in a 5.5% drop in overall coral cover. These "silent" effects of temperature increase do not appear to have been reported elsewhere in the literature.     

Inhibition of larval swimming activity of the coral (Platygyra acuta) by interactive thermal and chemical stresses

This study examined the interactive effects of copper and elevated temperature and subsequent depuration on Platygyra acuta coral larvae. Larval mortality and motility were significantly affected by copper alone (70% and 100% inhibition respectively). Their respective lethal/inhibitory concentration (LC₅₀/IC₅₀) were 10–130% and 86–193% higher than those reported for other larvae. Temperature (ambient, 27 °C and elevated, 30 °C) alone and the combination of temperature and copper did not significantly affect both endpoints. This study provides the first quantitative data on depuration effect on resumption of larval motility after copper exposure, although no sign of larval recovery was observed. These findings suggest that the effects of copper pollution outweigh the thermal tolerance of coral larvae. High LC₅₀ and IC₅₀ recorded may be unique for corals from marginal reefs like Hong Kong which have already been exposed to high levels of copper pollution.     

Coral bleaching and mortality on artificial and natural reefs in Maldives in 1998, sea surface temperature anomalies and initial recovery

The bleaching and subsequent mortality of branching and massive corals on artificial and natural reefs in the central atolls of Maldives in 1998 are examined with respect to sea surface temperature (SST) anomalies. SST normally peaks in April-May in Maldives. The UK Meteorological Office's Global sea-Ice and SST data set version 2.3 b shows that in 1998 monthly mean SST was 1.2-4 S.D. above the 1950-1999 average during the warmest months (March-June), with the greatest anomaly in May of +2.1 degrees C. Bleaching was first reported in mid-April and was severe from late April to mid-May with some recovery evident by late-May. At least 98% of branching corals (Acroporidae, Pocilloporidae) on artificial structures deployed on a reef flat in 1990 died whereas the majority of massive corals (Poritidae, Faviidae, Agariciidae) survived the bleaching. The pre-bleaching coral community on the artificial reefs in 1994 was 95% branching corals and 5% massives (n = 1589); the post-bleaching community was 3% branching corals and 97% massives (n = 248). Significant reductions in live coral cover were seen at all natural reefs surveyed in the central atolls, with average live coral cover decreasing from about 42% to 2%, a 20-fold reduction from pre-bleaching levels. A survey of recruitment of juvenile corals to the artificial structures 10 months after the bleaching event showed that 67% of recruits (> or = 0.5 cm diameter) were acroporids and pocilloporids and 33% were from massive families (n = 202) compared to 94% and 6%, respectively, in 1990-1994 (n = 3136). Similar post-bleaching dominance of recruitment by branching corals was seen on nearby natural reef (78% acroporids and pocilloporids; 22% massives). A linear regression of April mean monthly SST against year was highly significant (p < 0.001) and suggests a rise of 0.16 degree C per decade. If this trend continues, by 2030 mean April SST in the central atolls will normally exceed the anomaly level at which corals appear there are susceptible to mass bleaching.     

Effects of algal turfs and sediment on coral settlement

Successful settlement and recruitment of corals is critical to the resilience of coral reefs. Given that many degraded reefs are dominated by benthic algae, recovery of coral populations after bleaching and other disturbances requires successful settlement amidst benthic algae. Algal turfs often accumulate sediments, sediments are known to inhibit coral settlement, and reefs with high inputs of terrestrial sediments are often dominated by turfs. We investigated the impacts of two algal turf assemblages, and of sediment deposits, on settlement of the coral Acropora millepora (Ehrenberg). Adding sediment reduced coral settlement, but the effects of different algal turfs varied. In one case, algal turfs inhibited coral settlement, whereas the other turf only inhibited settlement when combined with sediments. These results provide the first direct, experimental evidence of effects of filamentous algal turfs on coral settlement, the variability in those effects, and the potential combined effects of algal turfs and trapped sediments.     

Recent progress on signalling molecules of coral-associated microorganisms

Coral reefs have high primary productivity and are one of the most important ecosystems in the ocean. However, the health and stability of coral reefs are constantly threatened by climate change and human activities. The coral holobiont is a complex symbiosis between the coral animal, zooxanthellae, and the associated community of microorganisms including bacteria, archaea, viruses, etc. Coral-associated microorganisms are found to be important for the maintenance of coral health,and they are proposed to contribute to the acclimatization and adaptation of reef-building corals under rapid climate change. The coral-microbe interaction mediated by chemical signalling molecules is an important driving force for shaping the microbial communities. Herein, we summarize our current knowledge of the chemical signalling molecules involved in the interaction of the coral holobiont. Specifically, the cooperation and competition among microbes mediated by N-acyl homoserine lactones(AHLs), the interrelationship between microbes and hosts mediated by dimethylsulfoniopropionate(DMSP) and nitric oxide(NO), and the response of corals and microorganisms to reactive oxygen species(ROS) under environmental stresses are highlighted in this review. We further discuss the potential of manipulating the coral microbiome using signalling molecules to restore and protect coral reefs.     

Photosystem II herbicide pollution in Hong Kong and its potential photosynthetic effects on corals

Photosystem II (PSII) herbicides have been shown to affect the photosynthesis of corals at low, environmentally relevant concentrations. The recent detection of the PSII herbicide Irgarol-1051 in coastal waters of Hong Kong at concentrations above the EC(50) for reduction of photosynthesis of corals prompted further investigation into the extent of PSII herbicide pollution in coral reefs of Hong Kong. Snap-shot and passive samples were taken from coral reef sites and evaluated via HPLC/MS-MS and a novel bioanalytical technique. Low concentrations (less than 10 ng L(-1)) of diuron and atrazine were found at all study sites. Extracts from these samples concentrated by a factor of 10 were found to reduce the photosynthetic yield of zooxanthellae. It appears unlikely that herbicide pollution is a key issue in isolation but may act synergistically with other stressors to reduce the viability of Hong Kong's coral reefs. The study has also demonstrated the feasibility of combining sample extraction techniques with a coral specific bioanalytical technique for a sensitive assessment of risks associated with herbicide exposure in corals.     

Coral recruitment onto an experimental pulverised fuel ash-concrete artificial reef

An experimental artificial reef was deployed in December 1993 at Hoi Ha Wan Marine Park, Hong Kong. This is the first study documenting natural scleractinian coral recruitment onto a stabilised pulverised fuel ash (PFA)-concrete artificial reef. Visible recruits were first recorded 9-10 months after the placement of reef blocks, i.e., in the autumn of 1994. Two scleractinians, Oulastrea crispata and Culicia japonica, were recruited. The recruit density of the former was much greater than the latter. The spatial recruitment pattern of the corals was observed to be affected by the orientation of the attaching surface. O. crispata settled predominantly on the undersides of the reef blocks. There was an edge effect on O. crispata recruitment. C. japonica, however, had a preference for exposed surfaces. O. crispata did not show a preference for block composition whereas C. japonica favoured blocks with high (75% by volume) PFA levels. This shows that PFA-concrete is a potential substratum for artificial reef construction, especially when such reefs aim at rehabilitating corals.     

ENCORE: the effect of nutrient enrichment on coral reefs. Synthesis of results and conclusions

Coral reef degradation resulting from nutrient enrichment of coastal waters is of increasing global concern. Although effects of nutrients on coral reef organisms have been demonstrated in the laboratory, there is little direct evidence of nutrient effects on coral reef biota in situ. The ENCORE experiment investigated responses of coral reef organisms and processes to controlled additions of dissolved inorganic nitrogen (N) and/or phosphorus (P) on an offshore reef (One Tree Island) at the southern end of the Great Barrier Reef, Australia. A multi-disciplinary team assessed a variety of factors focusing on nutrient dynamics and biotic responses. A controlled and replicated experiment was conducted over two years using twelve small patch reefs ponded at low tide by a coral rim. Treatments included three control reefs (no nutrient addition) and three + N reefs (NH4Cl added), three + P reefs (KH2PO4 added), and three + N + P reefs. Nutrients were added as pulses at each low tide (ca twice per day) by remotely operated units. There were two phases of nutrient additions. During the initial, low-loading phase of the experiment nutrient pulses (mean dose = 11.5 microM NH4+; 2.3 microM PO4(-3)) rapidly declined, reaching near-background levels (mean = 0.9 microM NH4+; 0.5 microM PO4(-3)) within 2-3 h. A variety of biotic processes, assessed over a year during this initial nutrient loading phase, were not significantly affected, with the exception of coral reproduction, which was affected in all nutrient treatments. In Acropora longicyathus and A. aspera, fewer successfully developed embryos were formed, and in A. longicyathus fertilization rates and lipid levels decreased. In the second, high-loading, phase of ENCORE an increased nutrient dosage (mean dose = 36.2 microM NH4+; 5.1 microM PO4(-3)) declining to means of 11.3 microM NH4+ and 2.4 microM PO4(-3) at the end of low tide) was used for a further year, and a variety of significant biotic responses occurred. Encrusting algae incorporated virtually none of the added nutrients. Organisms containing endosymbiotic zooxanthellae (corals and giant clams) assimilated dissolved nutrients rapidly and were responsive to added nutrients. Coral mortality, not detected during the initial low-loading phase, became evident with increased nutrient dosage, particularly in Pocillopora damicornis. Nitrogen additions stunted coral growth, and phosphorus additions had a variable effect. Coral calcification rate and linear extension increased in the presence of added phosphorus but skeletal density was reduced, making corals more susceptible to breakage. Settlement of all coral larvae was reduced in nitrogen treatments, yet settlement of larvae from brooded species was enhanced in phosphorus treatments. Recruitment of stomatopods, benthic crustaceans living in coral rubble, was reduced in nitrogen and nitrogen plus phosphorus treatments. Grazing rates and reproductive effort of various fish species were not affected by the nutrient treatments. Microbial nitrogen transformations in sediments were responsive to nutrient loading with nitrogen fixation significantly increased in phosphorus treatments and denitrification increased in all treatments to which nitrogen had been added. Rates of bioerosion and grazing showed no significant effects of added nutrients. ENCORE has shown that reef organisms and processes investigated in situ were impacted by elevated nutrients. Impacts were dependent on dose level, whether nitrogen and/or phosphorus were elevated and were often species-specific. The impacts were generally sub-lethal and subtle and the treated reefs at the end of the experiment were visually similar to control reefs. Rapid nutrient uptake indicates that nutrient concentrations alone are not adequate to assess nutrient condition of reefs. Sensitive and quantifiable biological indicators need to be developed for coral reef ecosystems. The potential bioindicators identified in ENCORE should be tested in future research on coral reef/nutrient interactions. Synergistic and cumulative effects of elevated nutrients and other environmental parameters, comparative studies of intact vs. disturbed reefs, offshore vs. inshore reefs, or the ability of a nutrient-stressed reef to respond to natural disturbances require elucidation. An expanded understanding of coral reef responses to anthropogenic impacts is necessary, particularly regarding the subtle, sub-lethal effects detected in the ENCORE studies.