Ecological condition of coral reef assemblages in the Cuban Archipelago

The condition of coral reefs in the Cuban Archipelago is poorly known. We aimed to analyse coral assemblages across 199 reef sites belonging to 12 localities. Crest and fore reefs were assessed using six metrics: species richness, density, coral cover, mortality, coral size and reef complexity. The condition of reefs varied across the archipelago from healthy to depleted reefs. The localities with best scores were Cienfuegos, Bahía de Cochinos and Cazones. These reefs have values of living coral cover (>20%) and complexity (>50?cm) similar to the best preserved Caribbean reefs. However, the majority of crest biotopes suffered important deterioration with old mortality of Acropora palmata populations and moderate coral cover (15%); although crest reefs still maintained their structural complexity. Despite moderate levels of coral cover in fore reefs (18%), their condition was alarming because 25% of the sites had cover below the recovery threshold of 10%, accumulated mortality and structural flattening. Compared with the 1980s, the species richness was roughly the same (42) for crest and fore reefs, although dominance has changed to widespread tolerant species. Coral reef assemblages varied at local and regional scales in similar magnitude, suggesting the combined effects of natural and anthropogenic drivers.     

Climatological context for large-scale coral bleaching

Large-scale coral bleaching was first observed in 1979 and has occurred throughout virtually all of the tropics since that time. Severe bleaching may result in the loss of live coral and in a decline of the integrity of the impacted coral reef ecosystem. Despite the extensive scientific research and increased public awareness of coral bleaching, uncertainties remain about the past and future of large-scale coral bleaching. In order to reduce these uncertainties and place large-scale coral bleaching in the longer-term climatological context, specific criteria and methods for using historical sea surface temperature (SST) data to examine coral bleaching-related thermal conditions are proposed by analyzing three, 132 year SST reconstructions: ERSST, HadISST1, and GISST2.3b. These methodologies are applied to case studies at Discovery Bay, Jamaica (77.27°W, 18.45°N), Sombrero Reef, Florida, USA (81.11°W, 24.63°N), Academy Bay, Galápagos, Ecuador (90.31°W, 0.74°S), Pearl and Hermes Reef, Northwest Hawaiian Islands, USA (175.83°W, 27.83°N), Midway Island, Northwest Hawaiian Islands, USA (177.37°W, 28.25°N), Davies Reef, Australia (147.68°E, 18.83°S), and North Male Atoll, Maldives (73.35°E, 4.70°N). The results of this study show that (1) The historical SST data provide a useful long-term record of thermal conditions in reef ecosystems, giving important insight into the thermal history of coral reefs and (2) While coral bleaching and anomalously warm SSTs have occurred over much of the world in recent decades, case studies in the Caribbean, Northwest Hawaiian Islands, and parts of other regions such as the Great Barrier Reef exhibited SST conditions and cumulative thermal stress prior to 1979 that were comparable to those conditions observed during the strong, frequent coral bleaching events since 1979. This climatological context and knowledge of past environmental conditions in reef ecosystems may foster a better understanding of how coral reefs will respond in future, ocean warming scenarios.     

The effects of sea surface temperature anomalies on oceanic coral reef systems in the southwestern tropical Atlantic

In 2010, high sea surface temperatures that were recorded in several parts of the world and caused coral bleaching and coral mortality were also recorded in the southwest Atlantic Ocean, between latitudes 0°S and 8°S. This paper reports on coral bleaching and diseases in Rocas Atoll and Fernando de Noronha archipelago and examines their relationship with sea surface temperature (SST) anomalies recorded by PIRATA buoys located at 8°S30°W, 0°S35°W, and 0°S23°W. Adjusted satellite data were used to derive SST climatological means at buoy sites and to derive anomalies at reef sites. The whole region was affected by the elevated temperature anomaly that persisted through 2010, reaching 1.67 °C above average at reef sites and 1.83 °C above average at buoys sites. A significant positive relationship was found between the percentage of coral bleaching that was observed on reef formations and the corresponding HotSpot SST anomaly recorded by both satellite and buoys. These results indicate that the warming observed in the ocean waters was followed by a warming at the reefs. The percentage of bleached corals persisting after the subsidence of the thermal stress, and disease prevalence increased through 2010, after two periods of thermal stress. The in situ temperature anomaly observed during the 2009–2010 El Niño event was equivalent to the anomaly observed during the 1997–1998 El Niño event, explaining similar bleaching intensity. Continued monitoring efforts are necessary to further assess the relationship between bleaching severity and PIRATA SST anomalies and improve the use of this new dataset in future regional bleaching predictions.     

Augmented coral reef monitoring using a stationary reef monitoring system

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

Extreme temperature events will drive coral decline in the Coral Triangle

In light of rapid environmental change, quantifying the contribution of regional‐ and local‐scale drivers of coral persistence is necessary to characterize fully the resilience of coral reef systems. To assess multiscale responses to thermal perturbation of corals in the Coral Triangle (CT), we developed a spatially explicit metacommunity model with coral–algal competition, including seasonal larval dispersal and external spatiotemporal forcing. We tested coral sensitivity in 2,083 reefs across the CT region and surrounding areas under potential future temperature regimes, with and without interannual climate variability, exploring a range of 0.5–2.0°C overall increase in temperature in the system by 2054. We found that among future projections, reef survival probability and mean percent coral cover over time were largely determined by the presence or absence of interannual sea surface temperature (SST) extremes as well as absolute temperature increase. Overall, reefs that experienced SST time series that were filtered to remove interannual variability had approximately double the chance of survival than reefs subjected to unfiltered SST. By the end of the forecast period, the inclusion of thermal anomalies was equivalent to an increase of at least 0.5°C in SST projections without anomalies. Change in percent coral cover varied widely across the region within temperature scenarios, with some reefs experiencing local extinction while others remaining relatively unchanged. Sink strength and current thermal stress threshold were found to be significant drivers of these patterns, highlighting the importance of processes that underlie larval connectivity and bleaching sensitivity in coral networks.     

Predicting coral bleaching hotspots: the role of regional variability in thermal stress and potential adaptation rates

Sea surface temperature fields (1870–2100) forced by CO₂-induced climate change under the IPCC SRES A1B CO₂ scenario, from three World Climate Research Programme Coupled Model Intercomparison Project Phase 3 (WCRP CMIP3) models (CCSM3, CSIRO MK 3.5, and GFDL CM 2.1), were used to examine how coral sensitivity to thermal stress and rates of adaption affect global projections of coral-reef bleaching. The focus of this study was two-fold, to: (1) assess how the impact of Degree-Heating-Month (DHM) thermal stress threshold choice affects potential bleaching predictions and (2) examine the effect of hypothetical adaptation rates of corals to rising temperature. DHM values were estimated using a conventional threshold of 1°C and a variability-based threshold of 2σ above the climatological maximum Coral adaptation rates were simulated as a function of historical 100-year exposure to maximum annual SSTs with a dynamic rather than static climatological maximum based on the previous 100 years, for a given reef cell. Within CCSM3 simulations, the 1°C threshold predicted later onset of mild bleaching every 5 years for the fraction of reef grid cells where 1°C > 2σ of the climatology time series of annual SST maxima (1961–1990). Alternatively, DHM values using both thresholds, with CSIRO MK 3.5 and GFDL CM 2.1 SSTs, did not produce drastically different onset timing for bleaching every 5 years. Across models, DHMs based on 1°C thermal stress threshold show the most threatened reefs by 2100 could be in the Central and Western Equatorial Pacific, whereas use of the variability-based threshold for DHMs yields the Coral Triangle and parts of Micronesia and Melanesia as bleaching hotspots. Simulations that allow corals to adapt to increases in maximum SST drastically reduce the rates of bleaching. These findings highlight the importance of considering the thermal stress threshold in DHM estimates as well as potential adaptation models in future coral bleaching projections.     

Thermal history influences lesion recovery of the threatened Caribbean staghorn coral Acropora cervicornis under heat stress

Anthropogenic climate change is the biggest threat to coral reefs, but reef restoration efforts are buying time for these ecosystems. Lesion recovery, which can be a determinant of colony survival, is particularly important for restored species. Here, we evaluate lesion recovery of 18 genets of Acropora cervicornis from Florida reefs with different thermal regimes in a temperature challenge experiment. Genets demonstrated significant variability in healing, which greatly slowed under heat stress. Only 35% of fragments healed at 31.5 °C compared to 99% at 28 °C. Donor reef thermal regime significantly influenced lesion recovery under heat stress with corals from warmer reefs demonstrating greater healing than corals from cooler reefs, but did not influence recovery under ambient conditions. These findings should encourage practitioners to utilize rapidly healing genets, avoid fragmentation in high temperatures, and incorporate assisted relocation by moving corals from warmer to cooler reefs, where they might succeed under future climate conditions.

     

Caribbean mesophotic coral ecosystems are unlikely climate change refugia

Deeper coral reefs experience reduced temperatures and light and are often shielded from localized anthropogenic stressors such as pollution and fishing. The deep reef refugia hypothesis posits that light‐dependent stony coral species at deeper depths are buffered from thermal stress and will avoid bleaching‐related mass mortalities caused by increasing sea surface temperatures under climate change. This hypothesis has not been tested because data collection on deeper coral reefs is difficult. Here we show that deeper (mesophotic) reefs, 30–75 m depth, in the Caribbean are not refugia because they have lower bleaching threshold temperatures than shallow reefs. Over two thermal stress events, mesophotic reef bleaching was driven by a bleaching threshold that declines 0.26 °C every +10 m depth. Thus, the main premise of the deep reef refugia hypothesis that cooler environments are protective is incorrect; any increase in temperatures above the local mean warmest conditions can lead to thermal stress and bleaching. Thus, relatively cooler temperatures can no longer be considered a de facto refugium for corals and it is likely that many deeper coral reefs are as vulnerable to climate change as shallow water reefs.     

Forecasting decadal changes in sea surface temperatures and coral bleaching within a Caribbean coral reef

Elevated sea surface temperature (SST) caused by global warming is one of the major threats to coral reefs. While increased SST has been shown to negatively affect the health of coral reefs by increasing rates of coral bleaching, how changes to atmospheric heating impact SST distributions, modified by local flow environments, has been less understood. This study aimed to simulate future water flow patterns and water surface heating in response to increased air temperature within a coral reef system in Bocas del Toro, Panama, located within the Caribbean Sea. Water flow and SST were modeled using the Delft3D-FLOW© computer simulation package. Locally measured physical parameters, including bathymetry, astronomic tidal forcing, and coral habitat distribution were input into the model and water flow, and SST was simulated over a four-month period under present day, as well as projected warming scenarios in 2020s, 2050s, and 2080s. Changes in SST, and hence the thermal stress to corals, were quantified by degree heating weeks. Results showed that present-day reported bleaching sites were consistent with localized regions of continuous high SST. Regions with highest SST were located within shallow coastal sites adjacent to the mainland or within the interior of the bay, and characterized by low currents with high water retention times. Under projected increases in SSTs, shallow reef areas in low flow regions were found to be hot spots for future bleaching.     

Morphologie, Ökologie und Zonierung von Korallenriffen bei Aqaba, (Golf von Aqaba,Rotes Meer)

The coral reefs of the Gulf of Aqaba are among the most northern ones of the world. This study, the first concerning the east coast of this topographically and hydrographically peculiar sea, considers relationships of biophysiographical and structural reef zones to fundamental abiotic environmental factors. An introduction to paleogeography, geology, petrography, topography, climate and hydrography is followed by terminological definitions used to describe the different reef areas. The investigations were carried out on two transects crossing fringing reefs of different shape. Each transect was 20 m wide and run from the shore over nearly 200 m to the fore reef in about 30 m depth. One reef, a “coastal-fringing reef”, represents an unaltered straight reef flat from shore to the reef edge 60 m away; two large pinnacles reach the surface some 125 m off the shore. The other reef, a “lagoon-fringing reef”, is divided into a 100 m wide lagoon of 0.5–2.3 m depth and a reef crest separated from the former by a rear reef. The reef platform of the lagoon-fringing reef is cut by a system of channels and tunnels; the reef edge is about 135 m off shore. Such water depth, substrate, temperature, illumination and water movement were recorded, about 200 common or dominant species (plants and animals) were collected, their distribution plotted and, together with other data and structural items, charted. Indicator species characterize the biophysiographical zones. Their variation as well as that of the structural and substrate zones depend on different zones of water movement. This basic factor also controls other ecological parameters such as food and oxygen supply as well as temperature and salinity gradients between fore reef and shore. From this point of view the ecological requirements of some indicator and other species and conversely the ecological settings of different reef areas are discussed. The different shapes of both reefs are explained on the basis of a “reef development cycle” — a hypothesis applicable to fringing reefs at unchanging sea level and based on the fact that only a small surf-influenced area of “living reef” is able to compensate for reef destruction: While a young coastal fringing reef is growing outwards, its back reef is gradually altered to a reef lagoon by erosion. After stillstand of seaward expansion the reef crest, too, is cut by a channel system eroded by rip currents. This stage is represented by the lagoon-fringing reef. Isolated pinnacles remain as remnants of the former reef crest; young coastal-fringing reefs develop from the shore. This stage is examplified by the first reef studied. Extension, growth intensity, dominant frame building corals, and the number of species of the Aqaba reefs are compared with those of Eilat and with reefs of the middle Red Sea, South India, Southwest-Pacific and Jamaica.     

AUV-based bed roughness mapping over a tropical reef

Identifying fixed bed roughness scales of hydrodynamic relevance to waves and currents is challenging around coral reefs due to their highly inhomogeneous bathymetry. In order to characterize the spatial variability in reef roughness, a quantitative analysis of high-resolution sidescan sonar backscatter is performed for the identification of distinct substrates around a tropical reef and is related to echo sounder-based roughness measurements. Data were collected in the vicinity of the Kilo Nalu Observatory on the south shore of Oahu using sidescan sonar and a narrow beam echo sounder incorporated in a REMUS-100 (Remote Environmental Monitoring UnitS) autonomous underwater vehicle (AUV). With basic statistics and principal component analysis of variables derived from the backscatter data, it is possible to discriminate between areas of rough reef, bare reef, and rippled sand. Echo sounder-derived spectral analysis did not reveal dominant length scales. However, by combining the seabed classification obtained from sidescan measurements with echo sounder data, spectral root mean square (RMS) height values of approximately 3.3 cm and 7.3 cm are assigned to the bare reef and rough reef areas, respectively, for roughness with wavelengths between 0.2 and 6 m.     

Coral reef resilience to thermal stress in the Eastern Tropical Pacific

Coral reefs worldwide are threatened by thermal stress caused by climate change. Especially devastating periods of coral loss frequently occur during El Niño‐Southern Oscillation (ENSO) events originating in the Eastern Tropical Pacific (ETP). El Niño‐induced thermal stress is considered the primary threat to ETP coral reefs. An increase in the frequency and intensity of ENSO events predicted in the coming decades threatens a pan‐tropical collapse of coral reefs. During the 1982–1983 El Niño, most reefs in the Galapagos Islands collapsed, and many more in the region were decimated by massive coral bleaching and mortality. However, after repeated thermal stress disturbances, such as those caused by the 1997–1998 El Niño, ETP corals reefs have demonstrated regional persistence and resiliency. Using a 44 year dataset (1970–2014) of live coral cover from the ETP, we assess whether ETP reefs exhibit the same decline as seen globally for other reefs. Also, we compare the ETP live coral cover rate of change with data from the maximum Degree Heating Weeks experienced by these reefs to assess the role of thermal stress on coral reef survival. We find that during the period 1970–2014, ETP coral cover exhibited temporary reductions following major ENSO events, but no overall decline. Further, we find that ETP reef recovery patterns allow coral to persist under these El Niño‐stressed conditions, often recovering from these events in 10–15 years. Accumulative heat stress explains 31% of the overall annual rate of change of living coral cover in the ETP. This suggests that ETP coral reefs have adapted to thermal extremes to date, and may have the ability to adapt to near‐term future climate‐change thermal anomalies. These findings for ETP reef resilience may provide general insights for the future of coral reef survival and recovery elsewhere under intensifying El Niño scenarios.     

Abundance and reproductive patterns of the excavating sponge Cliona vermifera: a threat to Pacific coral reefs?

Cliona vermifera is a common excavating sponge in coral reefs from the East Pacific. Abundance and reproductive patterns of the sponge in a Mexican Pacific coral reef over a 4-year period are herein described. Sponge abundance was estimated along three transects 50 m long which were randomly placed on the reef, and along each one, a piece of coral rubble and a branch of a live coral from the Pocillopora spp. coral colony closest to the transect were collected at random, approximately every 2 m, yielding 25 pieces of each category per transect (and 75 pieces total of each category). A 2-way ANOVA revealed that invasion was significantly higher in living coral colonies (34.8 %) than in rubble (13.7 %). It also indicated that the abundance in both coralline substrates showed a temporal variation without a clear pattern of increase over the years. It was estimated that 60–85 % of sponges in the population reproduced sexually every year. The sponge proved gonochoristic, with a sex ratio strongly departing from parity (1 male: 3 females). Over the 4-year study period, at least two cohorts of oocytes with densities of up to 3.5 oocytes per mm² tissue were observed. Spermatogenesis lasted about a month, but often producing more than a pulse from July to November, coupled with peaks of oocyte maturation. Fertilization occurred internally to produce encapsulated zygotes that were released in one or more spawning events from July to November. In the following months (December to February), which were the periods of lowest temperature (~18.5–20 °C), no gametic activity occurred in the sponges. Because anomalous temperature rises that are detrimental to corals do not appear to negatively affect the reproduction and abundance of C. vermifera, it is likely that the excavating activity of this sponge may be compromising the health of those coral reefs that are recurrently affected by episodes of thermal stress.     

Patterns of scleractinian coral recruitment at Lord Howe Island,an isolated subtropical reef off eastern Australia

Changing oceanic conditions, particularly ocean warming and altered currents, can affect the reproductive success of corals. Improving the knowledge of coral reproductive processes at the marginal range limits of coral reefs is important for understanding the ecology of subtropical coral communities and the potential for coral species to expand their ranges in higher latitudes in the future. The extent of live coral cover around subtropical Lord Howe Island (LHI; 31°33′S, 159°05′E) approximately 600 km off the east coast of Australia, has been relatively stable over the last several decades; however, shifts in dominant species in the adult coral community have been reported. To examine the potential influences of recent altered currents and shifts in dominant scleractinian taxa within this community, this study examined spatial and seasonal variation of coral larval settlement at different habitats within the LHI reef lagoon. The study also assessed whether the assemblage of scleractinian corals settling at LHI has changed between 1990–1991 and 2011–2012. Mean densities of coral settlement in 2011–2012 (230 spat m^?2 yr^?1) were consistent with those reported in 1990–1991 and in other regions. However, changes in taxonomic composition were apparent with increases in the proportion of Acroporidae spat at some sites. Settlement of all taxa was highest over summer months, whereas during winter only one coral spat (Pocilloporidae) was detected. Coral settlement was highest and most taxonomically diverse at sites closest to the reef crest, where mortality of settled spat was also greatest. Rates of settlement were high compared with juvenile densities; hence, post-settlement mortality is also likely to be high. Post-settlement processes, influenced by local environmental conditions, are likely to be very important in structuring the adult coral communities within the LHI reef lagoon.     

Coral mortality associated with thermal fluctuations in the Phoenix Islands, 2002�C2005

The Phoenix Islands (Republic of Kiribati, 172–170°W and 2.5–5°S) experience intra- and inter-annual sea surface temperature variability of ≈2°C and have few local anthropogenic impacts. From July 2002, a thermal stress event occurred, which peaked at 21 Degree Heating Weeks (DHW) in January 2003 and persisted for 4 years. Such thermal stress was greater than any thermal event reported in the coral reef literature. Reef surveys were conducted in July 2000, June 2002, and May 2005, for six of the eight islands. Sampling was stratified by exposure (windward, leeward, and lagoon) and depth (5, 10, 15, and 25 m). The thermal stress event caused mass coral mortality, and coral cover declined by approximately 60% between 2002 and 2005. However, mortality varied among sites (12–100%) and among islands (42–79%) and varied in accordance with the presence of a lagoon, island size, and windward vs. leeward exposure. Leeward reefs experienced the highest and most consistent decline in coral cover. Island size and the presence of a lagoon showed positive correlations with coral mortality, most likely because of the longer water residence time enhancing heating. Windward reefs showed cooler conditions than leeward reefs. Recently dead corals were observed at depths >35 m on windward and >45 m on leeward reefs. Between-island variation in temperature had no effect on between-island variation in coral mortality. Mortality levels reported here were comparable to those reported for the most extreme thermal stress events of 9–10 DHW in other regions. These results highlight the high degree of acclimation and/or adaptation of the corals in the Phoenix Islands to their local temperature regime, and their consequent vulnerability to anomalous events. Moreover, the results suggest the need to adjust thermal stress calculations to reflect local temperature variation.     

Tropical cyclone cooling combats region‐wide coral bleaching

Coral bleaching has become more frequent and widespread as a result of rising sea surface temperature (SST). During a regional scale SST anomaly, reef exposure to thermal stress is patchy in part due to physical factors that reduce SST to provide thermal refuge. Tropical cyclones (TCs – hurricanes, typhoons) can induce temperature drops at spatial scales comparable to that of the SST anomaly itself. Such cyclone cooling can mitigate bleaching across broad areas when well‐timed and appropriately located, yet the spatial and temporal prevalence of this phenomenon has not been quantified. Here, satellite SST and historical TC data are used to reconstruct cool wakes (n=46) across the Caribbean during two active TC seasons (2005 and 2010) where high thermal stress was widespread. Upon comparison of these datasets with thermal stress data from Coral Reef Watch and published accounts of bleaching, it is evident that TC cooling reduced thermal stress at a region‐wide scale. The results show that during a mass bleaching event, TC cooling reduced thermal stress below critical levels to potentially mitigate bleaching at some reefs, and interrupted natural warming cycles to slow the build‐up of thermal stress at others. Furthermore, reconstructed TC wave damage zones suggest that it was rare for more reef area to be damaged by waves than was cooled (only 12% of TCs). Extending the time series back to 1985 (n = 314), we estimate that for the recent period of enhanced TC activity (1995–2010), the annual probability that cooling and thermal stress co‐occur is as high as 31% at some reefs. Quantifying such probabilities across the other tropical regions where both coral reefs and TCs exist is vital for improving our understanding of how reef exposure to rising SSTs may vary, and contributes to a basis for targeting reef conservation.     

Global warming, regional trends and inshore environmental conditions influence coral bleaching in Hawaii

Hawaiian waters show a trend of increasing temperature over the past several decades that are consistent with observations in other coral reef areas of the world. The first documented large‐scale coral bleaching occurred in the Hawaii region during late summer of 1996, with a second in 2002. The bleaching events in Hawaii were triggered by a prolonged regional positive oceanic sea surface temperature (SST) anomaly greater than 1°C that developed offshore during the time of annual summer temperature maximum. High solar energy input and low winds further elevated inshore water temperature by 1–2°C in reef areas with restricted water circulation (bays, reef flats and lagoons) and in areas where mesoscale eddies often retain water masses close to shore for prolonged periods of time. Data and observations taken during these events illustrate problems in predicting the phenomena of large‐scale bleaching. Forecasts and hind‐casts of these events are based largely on offshore oceanic SST records, which are only a first approximation of inshore reef conditions. The observed oceanic warming trend is the ultimate cause of the increase in the frequency and severity of bleaching events. However, coral reefs occur in shallow inshore areas where conditions are influenced by winds, orographic cloud cover, complex bathymetry, waves and inshore currents. These factors alter local temperature, irradiance, water motion and other physical and biological variables known to influence bleaching.     

Susceptibility of central Red Sea corals during a major bleaching event

A major coral bleaching event occurred in the central Red Sea near Thuwal, Saudi Arabia, in the summer of 2010, when the region experienced up to 10–11 degree heating weeks. We documented the susceptibility of various coral taxa to bleaching at eight reefs during the peak of this thermal stress. Oculinids and agaricids were most susceptible to bleaching, with up to 100 and 80 % of colonies of these families, respectively, bleaching at some reefs. In contrast, some families, such as mussids, pocilloporids, and pectinids showed low levels of bleaching (<20 % on average). We resurveyed the reefs 7 months later to estimate subsequent mortality. Mortality was highly variable among taxa, with some taxa showing evidence of full recovery and some (e.g., acroporids) apparently suffering nearly complete mortality. The unequal mortality among families resulted in significant change in community composition following the bleaching. Significant factors in the likelihood of coral bleaching during this event were depth of the reef and distance of the reef from shore. Shallow reefs and inshore reefs had a higher prevalence of bleaching. This bleaching event shows that Red Sea reefs are subject to the same increasing pressures that reefs face worldwide. This study provides a quantitative, genus-level assessment of the vulnerability of various coral groups from within the Red Sea to bleaching and estimates subsequent mortality. As such, it can provide valuable insights into the future for reef communities in the Red Sea.     

Suitable Environmental Ranges for Potential Coral Reef Habitats in the Tropical Ocean

Coral reefs are found within a limited range of environmental conditions or tolerance limits. Estimating these limits is a critical prerequisite for understanding the impacts of climate change on the biogeography of coral reefs. Here we used the diagnostic model ReefHab to determine the current environmental tolerance limits for coral reefs and the global distribution of potential coral reef habitats as a function of six factors: temperature, salinity, nitrate, phosphate, aragonite saturation state, and light. To determine these tolerance limits, we extracted maximum and minimum values of all environmental variables in corresponding locations where coral reefs are present. We found that the global, annually averaged tolerance limits for coral reefs are 21.7—29.6 °C for temperature, 28.7—40.4 psu for salinity, 4.51 μmol L^-1 for nitrate, 0.63 μmol L^-1 for phosphate, and 2.82 for aragonite saturation state. The averaged minimum light intensity in coral reefs is 450 μmol photons m^-2 s^-1. The global area of potential reef habitats calculated by the model is 330.5 × 10³ km². Compared with previous studies, the tolerance limits for temperature, salinity, and nutrients have not changed much, whereas the minimum value of aragonite saturation in coral reef waters has decreased from 3.28 to 2.82. The potential reef habitat area calculated with ReefHab is about 121×10³ km² larger than the area estimated from the charted reefs, suggesting that the growth potential of coral reefs is higher than currently observed.     

Coral-algal competition on damaged reefs

Natural and anthropogenic catastrophes occurred at the end of the previous and in the beginning of the current centuries at the coral reefs of the World Ocean, and their consequences for the tropical shelf ecosystems have been described based on published data and our own investigations. It has been shown that in recent decades coral populations on reefs of tropical and subtropical regions of the World Ocean have been reduced by 80%, and in some areas have completely vanished. The biodiversity of reef ecosystems has been considerably reduced. The main reason for such changes is a 1-2°C increase in the temperature of surface waters in comparison with the monthly mean temperature in the hot season. The fate of the damaged coral reefs is under discussion. It is thought that in clean waters partially damaged coral reefs can recover, whereas in waters polluted as the result of human activity they collapse. The rate of coral reef restoration depends on the hydrological and hydrochemical conditions, frequency of natural calamities and competitive interrelation of algae and corals on the damaged sites of coral reefs. The nature of competitive interrelation between algae and corals is considered, viz., the dynamics of obliteration of damaged and dead coral colonies by various algal species, mechanisms of competitive interrelation, effects of the environment on the competitive ability of corals and algae, the internal and external conditions for victory in competitive activity. It has been suggested that coral reefs can be restored through temporary transformation into a vegetable reef. In the absence of natural calamities damaged reefs can be clearly restored to their original or altered state over several decades, but only in clean waters.