Species and population genomic differentiation in Pocillopora corals (Cnidaria, Hexacorallia).

Correctly delimiting species and populations is a prerequisite for studies of connectivity, adaptation and conservation. Genomic data are particularly useful to test species differentiation for organisms with few informative morphological characters or low discrimination of cytoplasmic markers, as in Scleractinians. Here we applied Restriction site Associated DNA sequencing (RAD-sequencing) to the study of species differentiation and genetic structure in populations of Pocillopora spp. from Oman and French Polynesia, with the objectives to test species hypotheses, and to study the genetic structure among sampling sites within species. We focused here on coral colonies morphologically similar to P. acuta (damicornis type ß). We tested the impact of different filtering strategies on the stability of the results. The main genetic differentiation was observed between samples from Oman and French Polynesia. These samples corresponded to different previously defined primary species hypotheses (PSH), i.e., PSHs 12 and 13 in Oman, and PSH 5 in French Polynesia. In Oman, we did not observe any clear differentiation between the two putative species PSH 12 and 13, nor between sampling sites. In French Polynesia, where a single species hypothesis was studied, there was no differentiation between sites. Our analyses allowed the identification of clonal lineages in Oman and French Polynesia. The impact of clonality on genetic diversity is discussed in light of individual-based simulations.

Juvenile corals underpin coral reef carbonate production after disturbance.

Sea-level rise is predicted to cause major damage to tropical coastlines. While coral reefs can act as natural barriers for ocean waves, their protection hinges on the ability of scleractinian corals to produce enough calcium carbonate (CaCO3 ) to keep up with rising sea levels. As a consequence of intensifying disturbances, coral communities are changing rapidly, potentially reducing community-level CaCO3 production. By combining colony-level physiology and long-term monitoring data, we show that reefs recovering from major disturbances can produce 40% more CaCO3 than currently estimated due to the disproportionate contribution of juvenile corals. However, the buffering effect of highly productive juvenile corals is compromised by recruitment failures, which have been more frequently observed after large-scale, repeated bleaching events. While the size structure of corals can bolster a critical ecological function on reefs, climate change impacts on recruitment may undermine this buffering effect, thus further compromising the persistence of reefs and their provision of important ecosystem services.

Recent and old duplications in crustaceans "Internal Transcribed Spacer 1″: structural and phylogenetic implications.

Internal Transcribed Spacer structures are important in preserving accessibility to specific enzymes for the maturation of rRNAs. ITS1 sequences reported in the literature in Crustaceans range between 182 and 820 bp and are characterized by the absence of repeats or the presence of only a limited number of microsatellites. Here, we sequenced ITS1 for a range of shrimp families (infraorder Caridea) and show that most taxa have much larger ITS1 sequences. We find a high number of microsatellites in Alpheus hebes and Crangon crangon and we report repeat units in Pandalidae, Palaemonidae and mainly in Alpheidae species. Up to four repeats were found in A. vanderbilti (1915 bp), A. rostratus (1635 bp) and A. lottini (1625 bp). In general, four helices were found in ITS1. Repeat units led to extra hairpins and loops. No conserved positions occurred except in helix 4. Three clades were defined in A. lottini for the first time. We estimated the ITS1 divergence rate for the three clades of A. lottini collected in French Polynesia using existing calibrations of substitution rates. Rates of sequence evolution are largely influenced by repeat units, which likely evolve separately. By comparison with COI marker, we estimated the divergence rate of the whole ITS1 sequence to range from 0.5 to 1.4% Pmy and between 0.12 and 0.5% for the 3' end of ITS1 located outside the repeat units. Given the degree of identity between repeats, we suggest that a duplication event recently occurred in A. floridanus (98% identity) whereas an ancient duplication happened in A. sulcatus (50% identity) early at the origination of the group Alpheidae, approximately 50 mya ago. In conclusion, our results highlight an over representation of shorter ITS1 sequences in public repositories, and underlines the importance to further understand patterns of molecular evolution of this functionally important gene.

Response of coral assemblages to thermal stress: are bleaching intensity and spatial patterns consistent between events?

Mass bleaching events resulting in coral mortality are among the greatest threats to coral reefs, and are projected to increase in frequency and intensity with global warming. Achieving a better understanding of the consistency of the response of coral assemblages to thermal stress, both spatially and temporally, is essential to determine which reefs are more able to tolerate climate change. We compared variations in spatial and taxonomic patterns between two bleaching events at the scale of an island (Moorea Island, French Polynesia). Despite similar thermal stress and light conditions, bleaching intensity was significantly lower in 2007 (approximately 37 % of colonies showed signs of bleaching) than in 2002, when 55 % of the colonies bleached. Variations in the spatial patterns of bleaching intensity were consistent between the two events. Among nine sampling stations at three locations and three depths, the stations at which the bleaching response was lowest in 2002 were those that showed the lowest levels of bleaching in 2007. The taxonomic patterns of susceptibility to bleaching were also consistent between the two events. These findings have important implications for conservation because they indicate that corals are capable of acclimatization and/or adaptation and that, even at small spatial scales, some areas are consistently more susceptible to bleaching than others.

Coral assemblages in Tonga: spatial patterns, replenishment capacities, and implications for conservation strategies.

Coral reefs in Tonga have been confronted by multiple threats of various origins, including large-scale disturbances and human-induced stressors. These reef communities have been poorly studied, and efficient conservation actions are urgently needed. The aim of this study was to: (1) examine the spatial distribution of coral assemblages in the lagoon of Tongatapu; (2) determine the degree to which spatial heterogeneity of adult corals is influenced by recruitment processes; and (3) examine the implications of these results in terms of conservation actions. We recorded a total of 37 adult and 28 juvenile coral genera, a mean density of 11.6 adult and 5.5 juvenile colonies m(-2), and a dominance of Montipora, Acropora, and Porites. For seven of the 10 dominant genera, spatial patterns of adults were linked to the short-term recruitment pattern history. Despite a reduced diversity and abundance of adult corals in some areas, the lagoon of Tongatapu retains the potential for replenishment through recruitment of young corals. Consequently, we suggest that conservation actions should focus on reducing factors causing coral mortality and maintain suitable conditions for the establishment and growth of juvenile corals, thus increasing the probability that they will reach maturity and participate to the maintenance of local populations. Rather than establishing a large marine protected area, which will almost certainly suffer from a lack of control and poor enforcement, alternative conservation measures could be successfully implemented through the establishment of several small village-based marine reserves, as has been undertaken in other South Pacific islands with promising results.

Effects of Madagascar marine reserves on juvenile and adult coral abundance, and the implication for population regulation.

Recruitment is a critical component in the dynamics of coral assemblages, and a key question is to determine the degree to which spatial heterogeneity of adults is influenced by pre-vs. post-settlement processes. We analyzed the density of juvenile and adult corals among 18 stations located at three regions around Madagascar, and examined the effects of Marine Protected Areas (MPAs). Our survey did not detect a positive effect of MPAs on juveniles, except for Porites at the study scale. The MPA effect was more pronounced for adults, notably for Acropora, Montipora, Seriatopora, and Porites at the regional scale. For most dominant genera, densities of juveniles and adults were positively correlated at the study scale, and at least at one of the three regions. These outcomes suggest recruitment-limitation relationships for several coral taxa, although differences in post-settlement events may be sufficiently strong to distort the pattern established at settlement for other populations. The modest benefits of MPAs on the density of juvenile corals demonstrated here argue in favor of strengthening conservation measures more specifically focused to protect recruitment processes.

Coral reef structural complexity loss exposes coastlines to waves.

Coral reefs offer natural coastal protection by attenuating incoming waves. Here we combine unique coral disturbance-recovery observations with hydrodynamic models to quantify how structural complexity dissipates incoming wave energy. We find that if the structural complexity of healthy coral reefs conditions is halved, extreme wave run-up heights that occur once in a 100-years will become 50 times more frequent, threatening reef-backed coastal communities with increased waves, erosion, and flooding.

Innate immune responses of a scleractinian coral to vibriosis.

Scleractinian corals are the most basal eumetazoan taxon and provide the biological and physical framework for coral reefs, which are among the most diverse of all ecosystems. Over the past three decades and coincident with climate change, these phototrophic symbiotic organisms have been subject to increasingly frequent and severe diseases, which are now geographically widespread and a major threat to these ecosystems. Although coral immunity has been the subject of increasing study, the available information remains fragmentary, especially with respect to coral antimicrobial responses. In this study, we characterized damicornin from Pocillopora damicornis, the first scleractinian antimicrobial peptide (AMP) to be reported. We found that its precursor has a segmented organization comprising a signal peptide, an acidic proregion, and the C-terminal AMP. The 40-residue AMP is cationic, C-terminally amidated, and characterized by the presence of six cysteine molecules joined by three intramolecular disulfide bridges. Its cysteine array is common to another AMP and toxins from cnidarians; this suggests a common ancestor, as has been proposed for AMPs and toxins from arthropods. Damicornin was active in vitro against Gram-positive bacteria and the fungus Fusarium oxysporum. Damicornin expression was studied using a combination of immunohistochemistry, reverse phase HPLC, and quantitative RT-PCR. Our data show that damicornin is constitutively transcribed in ectodermal granular cells, where it is stored, and further released in response to nonpathogenic immune challenge. Damicornin gene expression was repressed by the coral pathogen Vibrio coralliilyticus. This is the first evidence of AMP gene repression in a host-Vibrio interaction.

The war of corals: patterns, drivers and implications of changing coral competitive performances across reef environments.

Amidst global environmental changes, predicting species' responses to future environments is a critical challenge for preserving biodiversity and associated human benefits. We explored the original idea that coral competitive performances, the ability of corals to preempt ecological space on the reef through territorial warfare, serve as indicators of species' ecological niches and environmental windows, and therefore, responses to future environments. Our surveys indicated that coral performances varied with taxonomic identity, size and position along environmental gradients, highlighting complex interplays between life-history, warfare-strategy and niche segregation. Our results forewarn that growing alterations of coastal environments may trigger shifts in coral dominance, with the decline of major reef-building taxa like acroporids, and emphasize the importance of limiting human impacts for coastal resilience. Our empirical approach untangles the complexity of species' battle-like interactions and can help identify winners and losers in various communities caught in the interplay between ecological niches, environmental windows and global changes.

A contemporary baseline of Madagascar's coral assemblages: Reefs with high coral diversity, abundance, and function associated with marine protected areas.

Madagascar is a major hotspot of biodiversity in the Western Indian Ocean, but, as in many other regions, coral reefs surrounding the island confront large-scale disturbances and human-induced local stressors. Conservation actions have been implemented with encouraging results for fisheries, though their benefit on coral assemblages has never been rigorously addressed. In this context, we analyzed the multiscale spatial variation of the composition, generic richness, abundance, life history strategies, and cover of coral assemblages among 18 stations placed at three regions around the island. The potential influences of marine protected areas (MPAs), algal cover, substrate rugosity, herbivorous fish biomass, and geographic location were also analyzed. Our results highlight the marked spatial variability, with variation at either or both regional and local scales for all coral descriptors. The northeast coastal region of Masoala was characterized by the high abundance of coral colonies, most notably of the competitive Acropora and Pocillopora genera and stress-tolerant taxa at several stations. The southwest station of Salary Nord was distinguished by lower abundances, with depauperate populations of competitive taxa. On the northwest coast, Nosy-Be was characterized by higher diversity and abundance as well as by high coral cover (~42-70%) recorded at unfished stations. Results clearly underline the positive effects of MPAs on all but one of the coral descriptors, particularly at Nosy-Be where the highest contrast between fished and unfished stations was observed. Biomass of herbivorous fishes, crustose coralline algae cover, and substrate rugosity were also positively related to several coral descriptors. The occurrence of reefs with high diversity, abundance, and cover of corals, including the competitive Acropora, is a major finding of this study. Our results strongly support the implementation of locally managed marine areas with strong involvement by primary users, particularly to assist in management in countries with reduced logistic and human resources such as Madagascar.

Scaling up calcification, respiration, and photosynthesis rates of six prominent coral taxa.

Coral reefs provide a range of important services to humanity, which are underpinned by community-level ecological processes such as coral calcification. Estimating these processes relies on our knowledge of individual physiological rates and species-specific abundances in the field. For colonial animals such as reef-building corals, abundance is frequently expressed as the relative surface cover of coral colonies, a metric that does not account for demographic parameters such as coral size. This may be problematic because many physiological rates are directly related to organism size, and failure to account for linear scaling patterns may skew estimates of ecosystem functioning. In the present study, we characterize the scaling of three physiological rates - calcification, respiration, and photosynthesis - considering the colony size for six prominent, reef-building coral taxa in Mo'orea, French Polynesia. After a seven-day acclimation period in the laboratory, we quantified coral physiological rates for three hours during daylight (i.e., calcification and gross photosynthesis) and one hour during night light conditions (i.e., dark respiration). Our results indicate that area-specific calcification rates are higher for smaller colonies across all taxa. However, photosynthesis and respiration rates remain constant over the colony-size gradient. Furthermore, we revealed a correlation between the demographic dynamics of coral genera and the ratio between net primary production and calcification rates. Therefore, intraspecific scaling of reef-building coral physiology not only improves our understanding of community-level coral reef functioning but it may also explain species-specific responses to disturbances.

Physiological responses of the scleractinian coral Pocillopora damicornis to bacterial stress from Vibrio coralliilyticus.

As the effects of climate change have become increasingly visible over the past three decades, coral reefs have suffered from a number of natural and anthropogenic disturbances that have caused a critical decline in coral populations. Among these disturbances are coral diseases, which have appeared with increasing frequency and severity, often in correlation with increases in water temperature. Although the crucial role played by Vibrio species in coral disease has been widely documented, the scientific community does not yet fully understand the infection process of Vibrio or its impact on coral physiology and immunology. Here, we investigated the physiological and transcriptomic responses of a major reef-building coral, Pocillopora damicornis, when exposed to a specific pathogen (Vibrio coralliilyticus) under virulent (increasing water temperature) and non-virulent (constant low temperature) conditions. The infection process was examined by electron microscopy and quantitative reverse-transcription PCR, and coral health was monitored by visual observations and measurements of zooxanthellar density. The results obtained suggest that coral tissue invasion occurs upon increasing water temperature only. Transcriptomic variations were investigated using a suppression-subtractive-hybridization approach, and the expression levels of six candidate immune-related genes were examined during bacterial exposure. These genes correspond to three lectin-like molecules putatively involved in the recognition of pathogens, two metal-binding proteins putatively involved in antibacterial response and one cystein protease inhibitor. The transcription patterns of these selected genes provide new insights into the responses of coral colonies to virulent versus non-virulent bacteria.

Diversity, structure and demography of coral assemblages on underwater lava flows of different ages at Reunion Island and implications for ecological succession hypotheses.

Understanding colonization of new habitats and ecological successions is key to ecosystem conservation. However, studies on primary successions are scarce for reef-building corals, due to the rarity of newly formed substratum and the long-term monitoring efforts required for their long life cycle and slow growth rate. We analysed data describing the diversity, structure and demography of coral assemblages on lava flows of different ages and coral reefs at Reunion Island, to evaluate the strength and mechanisms of succession, and its agreement to the theoretical models. No significant differences were observed between the two habitats for most structure and demographic descriptors. In contrast, species richness and composition differentiated coral reefs from lava flows, but were not related to the age of the lava flow. We observed a strong dominance of Pocillopora colonies, which underline the opportunistic nature of this taxa, with life-history traits advantageous to dominance on primary and secondary successional stages. Although some results argue in favor of the tolerance model of succession, the sequences of primary successions as theorized in other ecosystems were difficult to observe, which is likely due to the high frequency and intensity of disturbances at Reunion, that likely distort or set back the expected successional sequences.

Temperature affects the reproductive outputs of coral-eating starfish Acanthaster spp. after adult exposure to near-future ocean warming and acidification.

Outbreaks of the coral-eating crown-of-thorns starfish Acanthaster spp. (COTS) have become to be amongst the most severe threats to coral reefs worldwide. Although most research has focused on COTS early development, it remains unclear how COTS populations will keep pace with changing ocean conditions. Since reproduction is a key process contributing to outbreaks, we investigated the reproductive success of adult COTS acclimated for 3-4 months to different treatment combinations of ambient conditions, ocean warming (+2 °C) and acidification (-0.35 pH). Our results suggest that the optimal breeding season in New Caledonia is concentrated around the end of the calendar year, when water temperature reaches >26 °C. We found negative effects of temperature on egg metrics, fertilisation success, and GSI, conflicting with previously documented effects of temperature on echinoderm reproductive outputs. Fertilisation success dropped drastically (more than threefold) with elevated temperature during the late breeding season. In contrast, we detected no effects of near-future acidification conditions on fertilisation success nor GSI. This is the first time that COTS reproduction is compared among individuals acclimated to different conditions of warming and acidification. Our results highlight the importance of accounting for adult exposure to better understand how COTS reproduction may be impacted in the face of global change.

Citizen Science, a promising tool for detecting and monitoring outbreaks of the crown-of-thorns starfish Acanthaster spp.

Monitoring potentially devastating coral-eating crown-of-thorns starfish (COTS) populations at scales relevant to management is a challenging task. Here, we investigated a citizen science approach to detect COTS outbreaks and prioritize management responses. Between 2014 and 2018, 38 000 COTS were recorded through 641 online observation reports submitted across New Caledonia, Vanuatu and Fiji by private stakeholders (51%), NGOs (22%), business operator (11%), research/government agencies (16%). COTS were observed in multiple areas, including in remote/inhabited reefs where they had never been reported. A three-level classification was developed to discriminate risk areas and propose operational guidelines to streamline management actions. About two-thirds of reports had low abundances (<10 starfish sighted) and could be addressed with low priority. Verification surveys at 65 reef sites confirmed outbreaks in half of the cases, along with high peak densities (7 000 ind.ha-1). Combining professional and non-professional observers increased the detection range (+27%) and the number of COTS detections (+129%). Citizen reports were eventually followed by removal campaigns organized within diverse institutional frameworks. While citizen monitoring has intrinsic limitations, we advocate that it constitutes a complementary and promising approach to support the ongoing management efforts in all countries affected by COTS.

Quantifying and addressing the prevalence and bias of study designs in the environmental and social sciences.

Building trust in science and evidence-based decision-making depends heavily on the credibility of studies and their findings. Researchers employ many different study designs that vary in their risk of bias to evaluate the true effect of interventions or impacts. Here, we empirically quantify, on a large scale, the prevalence of different study designs and the magnitude of bias in their estimates. Randomised designs and controlled observational designs with pre-intervention sampling were used by just 23% of intervention studies in biodiversity conservation, and 36% of intervention studies in social science. We demonstrate, through pairwise within-study comparisons across 49 environmental datasets, that these types of designs usually give less biased estimates than simpler observational designs. We propose a model-based approach to combine study estimates that may suffer from different levels of study design bias, discuss the implications for evidence synthesis, and how to facilitate the use of more credible study designs.

Coral bleaching under thermal stress: putative involvement of host/symbiont recognition mechanisms.

BACKGROUND: Coral bleaching can be defined as the loss of symbiotic zooxanthellae and/or their photosynthetic pigments from their cnidarian host. This major disturbance of reef ecosystems is principally induced by increases in water temperature. Since the beginning of the 1980s and the onset of global climate change, this phenomenon has been occurring at increasing rates and scales, and with increasing severity. Several studies have been undertaken in the last few years to better understand the cellular and molecular mechanisms of coral bleaching but the jigsaw puzzle is far from being complete, especially concerning the early events leading to symbiosis breakdown. The aim of the present study was to find molecular actors involved early in the mechanism leading to symbiosis collapse. RESULTS: In our experimental procedure, one set of Pocillopora damicornis nubbins was subjected to a gradual increase of water temperature from 28 degrees C to 32 degrees C over 15 days. A second control set kept at constant temperature (28 degrees C). The differentially expressed mRNA between the stressed states (sampled just before the onset of bleaching) and the non stressed states (control) were isolated by Suppression Subtractive Hybridization. Transcription rates of the most interesting genes (considering their putative function) were quantified by Q-RT-PCR, which revealed a significant decrease in transcription of two candidates six days before bleaching. RACE-PCR experiments showed that one of them (PdC-Lectin) contained a C-Type-Lectin domain specific for mannose. Immunolocalisation demonstrated that this host gene mediates molecular interactions between the host and the symbionts suggesting a putative role in zooxanthellae acquisition and/or sequestration. The second gene corresponds to a gene putatively involved in calcification processes (Pdcyst-rich). Its down-regulation could reflect a trade-off mechanism leading to the arrest of the mineralization process under stress. CONCLUSION: Under thermal stress zooxanthellae photosynthesis leads to intense oxidative stress in the two partners. This endogenous stress can lead to the perception of the symbiont as a toxic partner for the host. Consequently, we propose that the bleaching process is due in part to a decrease in zooxanthellae acquisition and/or sequestration. In addition to a new hypothesis in coral bleaching mechanisms, this study provides promising biomarkers for monitoring coral health.

Multiscale variability in coral recruitment in the Mascarene Islands: From centimetric to geographical scale.

Coral recruitment refers to the processes allowing maintenance and renewal of coral communities. Recruitment success is therefore indispensable for coral reef recovery after disturbances. Recruitment processes are governed by a variety of factors occurring at all spatial and temporal scales, from centimetres to hundreds of kilometres. In the present context of rising disturbances, it is thus of major importance to better understand the relative importance of different scales in this variation, and when possible, the factors associated with these scales. Multiscale spatio-temporal variability of scleractinian coral recruitment was investigated at two of the Mascarene Islands: Reunion and Rodrigues. Recruitment rates and taxonomic composition were examined during three consecutive six-month periods from regional to micro-local scales (i.e. from hundreds of kilometres to few centimetres) and between two protection levels (no-take zones and general protection zones). Very low recruitment rates were observed. Rodrigues displayed lower recruitment rates than Reunion. Recruit assemblage was dominated by Pocilloporidae (77.9%), followed by Acroporidae (9.9%) and Poritidae (5.2%). No protection effect was identified on coral recruitment, despite differences in recruitment rates among sites within islands. Recruits were patchily distributed within sites but no aggregative effect was detected, i.e. the preferentially colonised tiles were not spatially grouped. Recruits settled mainly on the sides of the tiles, especially at Rodrigues, which could be attributed to the high concentration of suspended matter. The variability of recruitment patterns at various spatial scales emphasises the importance of micro- to macro-local variations of the environment in the dynamics and maintenance of coral populations. High temporal variability was also detected, between seasons and years, which may be related to the early 2016 bleaching event at Rodrigues. The low recruitment rates and the absence of protection effect raise questions about the potential for recovery from disturbances of coral reefs in the Mascarene Islands.

Social-environmental drivers inform strategic management of coral reefs in the Anthropocene.

Without drastic efforts to reduce carbon emissions and mitigate globalized stressors, tropical coral reefs are in jeopardy. Strategic conservation and management requires identification of the environmental and socioeconomic factors driving the persistence of scleractinian coral assemblages-the foundation species of coral reef ecosystems. Here, we compiled coral abundance data from 2,584 Indo-Pacific reefs to evaluate the influence of 21 climate, social and environmental drivers on the ecology of reef coral assemblages. Higher abundances of framework-building corals were typically associated with: weaker thermal disturbances and longer intervals for potential recovery; slower human population growth; reduced access by human settlements and markets; and less nearby agriculture. We therefore propose a framework of three management strategies (protect, recover or transform) by considering: (1) if reefs were above or below a proposed threshold of >10% cover of the coral taxa important for structural complexity and carbonate production; and (2) reef exposure to severe thermal stress during the 2014-2017 global coral bleaching event. Our findings can guide urgent management efforts for coral reefs, by identifying key threats across multiple scales and strategic policy priorities that might sustain a network of functioning reefs in the Indo-Pacific to avoid ecosystem collapse.

Recovery of coral assemblages despite acute and recurrent disturbances on a South Central Pacific reef.

Coral reefs are increasingly threatened by various types of disturbances, and their recovery is challenged by accelerating, human-induced environmental changes. Recurrent disturbances reduce the pool of mature adult colonies of reef-building corals and undermine post-disturbance recovery from newly settled recruits. Using a long-term interannual data set, we show that coral assemblages on the reef slope of Moorea, French Polynesia, have maintained a high capacity to recover despite a unique frequency of large-scale disturbances which, since the 1990s, have caused catastrophic declines in coral cover and abundance. In 2014, only four years after one of the most extreme cases of coral decline documented, abundance of juvenile and adult colonies had regained or exceeded pre-disturbance levels, and no phase-shift to macroalgal dominance was recorded. This rapid recovery has been achieved despite constantly low coral recruitment rates, suggesting a high post-disturbance survivorship of recruits. However, taxonomic differences in coral susceptibility to disturbances and contrasting recovery trajectories have resulted in changes in the relative composition of species. In the present context of global coral reef decline, our study establishes a new benchmark for the capacity of certain benthic reef communities to sustain and recover their coral cover from repeated, intense disturbances.