Fitness consequences of habitat variability,trophic position,and energy allocation across the depth distribution of a coral-reef fish

Environmental clines such as latitude and depth that limit species’ distributions may be associated with gradients in habitat suitability that can affect the fitness of an organism. With the global loss of shallow-water photosynthetic coral reefs, mesophotic coral ecosystems (~30–150 m) may be buffered from some environmental stressors, thereby serving as refuges for a range of organisms including mobile obligate reef dwellers. Yet habitat suitability may be diminished at the depth boundary of photosynthetic coral reefs. We assessed the suitability of coral-reef habitats across the majority of the depth distribution of a common demersal reef fish (Stegastes partitus) ranging from shallow shelf (SS, <10 m) and deep shelf (DS, 20–30 m) habitats in the Florida Keys to mesophotic depths (MP, 60–70 m) at Pulley Ridge on the west Florida Shelf. Diet, behavior, and potential energetic trade-offs differed across study sites, but did not always have a monotonic relationship with depth, suggesting that some drivers of habitat suitability are decoupled from depth and may be linked with geographic location or the local environment. Feeding and diet composition differed among depths with the highest consumption of annelids, lowest ingestion of appendicularians, and the lowest gut fullness in DS habitats where predator densities were highest and fish exhibited risk-averse behavior that may restrict foraging. Fish in MP environments had a broader diet niche, higher trophic position, and higher muscle C:N ratios compared to shallower environments. High C:N ratios suggest increased tissue lipid content in fish in MP habitats that coincided with higher investment in reproduction based on gonado-somatic index. These results suggest that peripheral MP reefs are suitable habitats for demersal reef fish and may be important refuges for organisms common on declining shallow coral reefs.

     

Depth-dependent indicators of algal turf herbivory throughout the Main Hawaiian Islands

Herbivorous fish are key to maintaining a balance between coral and algae on reefs, where reefs with greater herbivore biomass often show lower algal cover. For reefs worldwide, algal turf cover is expanding and is increasingly used as an indicator of disturbance. Water depth affects reef fish composition; thus, it may be expected that herbivory could also differ by depth. We examined relationships between algal turf cover and biomass (g m⁻²), density (# m⁻²) and size (cm) of herbivore groups (grazers, browsers and scrapers) across shallow (< 6 m), mid (6–18 m) and deep (18–30 m) coral reefs in the Main Hawaiian Islands. We find that across all depth classes, algal turf cover decreased with increasing grazer and scraper density, with steeper relationships observed at mid and deep reefs than in shallow reefs. In contrast, algal turf cover slightly increased with increasing grazer and browser biomass at deep reefs. Considering fish size, algal turf cover increased with larger grazer and scrapers at mid and deep reefs. The results indicate that herbivorous fish density, rather than biomass, is a better indicator of reductions in algal turf cover and resulting coral-algal balance on Hawaiian reefs, where smaller fish exert greater top-down control on cover than larger fish. Despite significant differences in herbivorous fish compositions, length-frequency distributions and fishing intensities across depth, algal turf cover remains similar across depths. Increases in fishing would have a disproportionately negative impact in deep than shallow reefs due to a lower overall fish density, where grazing functions in deep reefs are maintained by significantly fewer and smaller grazers and browsers, and larger scrapers, than in shallow reefs. Developing an understanding of patterns of algal turf herbivory by depth is important to understanding the spatial scale at which herbivory and regime shifts operate.

     

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

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

A deep nursery for juveniles of the zebra angelfish <Emphasis Type="Italic">Genicanthus caudovittatus</Emphasis>

Juveniles of many coral reef fish species are thought to either follow the same bathymetric distribution patterns as the adults, or to occupy shallower waters. However, our knowledge base suffers a dearth of data from the deep reefs (>40 m). In a recent survey of the deep reefs of the northern Gulf of Aqaba (<65 m), we examined the bathymetric distribution of 26 diurnal zooplanktivorous species. In sharp contrast to the general trend known from the literature and from this research, the abundance of juvenile zebra angelfish, Genicanthus caudovittatus, peaked at deeper waters (60–65 m) compared with the adults (30 m). This suggests that the deeper reefs may serve as nursery grounds for the zebra angelfish. Peak juvenile abundance coincided with relatively low predator abundances. This raises the question, which factors constrain the bathymetric distribution of the remaining species. Our findings stress the potential importance of deep coral reef research for understanding the ecological patterns and processes that govern reef community structure.     

Impact of cyclones on hard coral and metapopulation structure,connectivity and genetic diversity of coral reef fish

Cyclones have one of the greatest effects on the biodiversity of coral reefs and the associated species. But it is unknown how stochastic alterations in habitat structure influence metapopulation structure, connectivity and genetic diversity. From 1993 to 2018, the reefs of the Capricorn Bunker Reef group in the southern part of the Great Barrier Reef were impacted by three tropical cyclones including cyclone Hamish (2009, category 5). This resulted in substantial loss of live habitat-forming coral and coral reef fish communities. Within 6–8 years after cyclones had devastated, live hard corals recovered by 50–60%. We show the relationship between hard coral cover and the abundance of the neon damselfish (Pomacentrus coelestis), the first fish colonizing destroyed reefs. We present the first long-term (2008–2015 years corresponding to 16–24 generations of P. coelestis) population genetic study to understand the impact of cyclones on the meta-population structure, connectivity and genetic diversity of the neon damselfish. After the cyclone, we observed the largest change in the genetic structure at reef populations compared to other years. Simultaneously, allelic richness of genetic microsatellite markers dropped indicating a great loss of genetic diversity, which increased again in subsequent years. Over years, metapopulation dynamics were characterized by high connectivity among fish populations associated with the Capricorn Bunker reefs (2200 km²); however, despite high exchange, genetic patchiness was observed with annual strong genetic divergence between populations among reefs. Some broad similarities in the genetic structure in 2015 could be explained by dispersal from a source reef and the related expansion of local populations. This study has shown that alternating cyclone-driven changes and subsequent recovery phases of coral habitat can greatly influence patterns of reef fish connectivity. The frequency of disturbances determines abundance of fish and genetic diversity within species.

     

Are artificial reefs surrogates of natural habitats for corals and fish in Dubai,United Arab Emirates?

Artificial reefs are often promoted as mitigating human impacts in coastal ecosystems and enhancing fisheries; however, evidence supporting their benefits is equivocal. Such structures must be compared with natural reefs in order to assess their performance, but past comparisons typically examined artificial structures that were too small, or were immature, relative to the natural reefs. We compared coral and fish communities on two large (>400,000 m³) and mature (>25 year) artificial reefs with six natural coral patches. Coral cover was higher on artificial reefs (50%) than in natural habitats (31%), but natural coral patches contained higher species richness (29 vs. 20) and coral diversity (H′ = 2.3 vs. 1.8). Multivariate analyses indicated strong differences between coral communities in natural and artificial habitats. Fish communities were sampled seasonally for 1 year. Multivariate fish communities differed significantly among habitat types in the summer and fall, but converged in the winter and spring. Univariate analysis indicated that species richness and abundance were stable throughout the year on natural coral patches but increased significantly in the summer on artificial reefs compared with the winter and spring, explaining the multivariate changes in community structure. The increased summer abundance on artificial reefs was mainly due to adult immigration. Piscivores were much more abundant in the fall than in the winter or spring on artificial reefs, but had low and stable abundance throughout the year in natural habitats. It is likely that the decreased winter and spring abundance of fish on the artificial reefs resulted from both predation and emigration. These results indicate that large artificial reefs can support diverse and abundant coral and fish communities. However, these communities differ structurally and functionally from those in natural habitats, and they should not be considered as replacements for natural coral and fish communities.     

Horizontal transmission of Symbiodinium cells between adult and juvenile corals is aided by benthic sediment

Of all reef-building coral species, 80–85 % initially draw their intracellular symbionts (dinoflagellates of the genus Symbiodinium) from the environment. Although Symbiodinium cells are crucial for the growth of corals and the formation of coral reefs, little is known about how corals first encounter free-living Symbiodinium cells. We report how the supply of free-living Symbiodinium cells to the benthos by adult corals can increase the rate of horizontal symbiont acquisition for conspecific recruits. Three species of newly settled aposymbiotic (i.e., symbiont-free) corals were maintained in an open aquarium system containing: sterilized sediment and adult coral fragments combined; adult coral fragments alone; sterilized sediment alone; or seawater at Heron Island, Great Barrier Reef, Australia. In all instances, the combination of an adult coral and sediment resulted in the highest symbiont acquisition rates by juvenile corals (up to five-fold greater than seawater alone). Juvenile corals exposed to individual treatments of adult coral or sediment produced an intermediate acquisition response (<52 % of recruits), and symbiont acquisition from unfiltered seawater was comparatively low (<20 % of recruits). Additionally, benthic free-living Symbiodinium cells reached their highest densities in the adult coral + sediment treatment (up to 1.2 × 10⁴ cells mL⁻¹). Our results suggest that corals seed microhabitats with free-living Symbiodinium cells suitable for many coral species during the process of coral recruitment.

     

Coral reefs studies and threats in Malaysia: a mini review

Coral reefs in Malaysia are about 4,006 km² with over 550 species contributed to nation’s economy. Coral reefs studies and threats in Malaysia have been reviewed briefly. Perspectives are addressed as coral reefs studies, threats, gaps and future studies. Coral reefs in Malaysia are being damaged at an increasing rate where it faces natural and anthropogenic stresses. Excellent summaries are available in terms of coral reefs cover throughout Malaysia however scarce in terms of qualitative, quantitative and biogeographical data. There are also limited studies on heavy metals concentration in corals skeleton studies. Poor to fair conditions of coral reefs in Peninsular Malaysia is due to increases of sedimentation and tourism impacts. Overfishing and fish blasting were main threats of coral reefs damage in Sabah. In Sarawak, coral reefs are threatened by high sedimentation and sand mining. The 1998–1999 bleaching event also affected coral reefs in Malaysia due to climate change. Gaps in coral reefs studies can be completed by continuous collaborations between local and international researchers as well as research by local universities. Economic valuation, policy analysis and community participation are directions in future coral reefs studies in Malaysia. Future studies are to understand effects of management on coral reefs health and impact of pollution on coral reefs growth with a standard coral reefs methodology. Established legal systems to reduce threats received by coral reefs are also need to be introduced. Role of science-driven management with community participation and media mass are also gaps to be highlighted in future studies.     

Low natural repopulation of marginal coral communities under the influence of upwelling

The study of coral repopulation in marginal communities may provide a useful analog for understanding the dynamics of coral reefs subjected to deleterious environmental changes. Repopulation of scleractinian reef corals may strongly impact the community structure on tropical reefs; however, the extent of this process on coral communities influenced by upwelling is unknown, especially in the Caribbean. In this study, the potential for natural repopulation of coral communities subjected to wind-driven upwelling was evaluated at three sites on the island of Cubagua, Venezuela. Coral spawning behavior was recorded and both larval settlement and juvenile abundance were estimated. Upwelling did not appear to affect coral spawning behavior, since at this locality spawning occurred at dates and times similar to those reported for well-developed reefs in the Caribbean. Also, juveniles produced by brooding corals were six times more abundant than those of broadcasting species, similar to patterns on other Caribbean reefs that are not under the influence of upwelling. By contrast, mean larval settlement (4 settlers m⁻²) and juvenile abundance (0.1 juveniles m⁻²) in Cubagua were both lower than those elsewhere in the Caribbean and on Pacific reefs. Thus, the potential for repopulation of these marginal communities seems lower than for well-developed coral reefs in other regions. These results suggest that more fully developed coral reefs also may have reduced repopulation potential, as they become influenced by suboptimal environmental conditions. Handling editor: I. Nagelkerken     

Neritic carbonate for six submerged coral reefs from northern Australia: Implications for Holocene global carbon dioxide

Total contribution of six recently discovered submerged coral reefs in northern Australia to Holocene neritic CaCO₃ and CO₂ is assessed to address a gap in global budgets. CaCO₃ production for the reef framework and inter-reefal deposits is 0.26–0.28 Mt. Holocene CO₂ production is 0.14–0.16 Mt. Coral and coralline algae are the dominant sources of Holocene CaCO₃ although foraminifers and molluscs are the dominant constituents of inter-reefal deposits. The total amount of Holocene neritic CaCO₃ produced by the six submerged coral reefs is several orders of magnitude smaller than that calculated using accepted CaCO₃ production values because of very low production, a ‘give-up’ growth history, and presumed significant dissolution and exports. The contribution of submerged coral reefs to global Holocene neritic CaCO₃ is estimated to be 0.26–0.62 Gt, which yields 0.15–0.37 Gt of CO₂. This amount of CO₂ is 0.02–0.05% of the 780 Gt added to the atmosphere since 18 kyr BP. Contributions from Australian submerged coral reefs are estimated to be 0.05 Gt CaCO₃ and 0.03 Gt CO₂ for an emergent reef area of 47.9 × 10³ km². Based on the growth history of the submerged coral reefs in the Gulf of Carpentaria, maximum global Holocene CaCO₃ fluxes could have attained 0.3 Gt yr^− 1 between 11 and 7 ka BP. This additional CaCO₃ would have culminated in a maximum CaCO₃ production from all (emergent and submerged) coral reefs of 1.2 Gt yr^− 1 and neritic CaCO₃ production of 2.75 Gt yr^− 1. The dilemma remains that the global area and CaCO₃ mass of submerged coral reefs are currently unknown. It is inevitable that many more submerged coral reefs will be found. Our findings imply that submerged coral reefs are a small but fundamental source of Holocene neritic CaCO₃ and CO₂.     

Environmental controls on daytime net community calcification on a Red Sea reef flat

Coral growth and carbonate accumulation form the foundation of the coral reef ecosystem. Changes in environmental conditions due to coastal development, climate change, and ocean acidification may pose a threat to net carbonate production in the near future. Controlled laboratory studies demonstrate that calcification by corals and coralline algae is sensitive to changes in aragonite saturation state (Ω_a), as well as temperature, light, and nutrition. Studies also show that the dissolution rate of carbonate substrates is impacted by changes in carbonate chemistry. The sensitivity of coral reefs to these parameters must be confirmed and quantified in the natural environment in order to predict how coral reefs will respond to local and global changes, particularly ocean acidification. We estimated the daytime hourly net community metabolic rates, both net community calcification (NCC) and net community productivity (NCP), at Sheltered Reef, an offshore platform reef in the central Red Sea. Average NCC was 8 ± 3 mmol m⁻² h⁻¹ in December 2010 and 11 ± 1 mmol m⁻² h⁻¹ in May 2011, and NCP was 21 ± 7 mmol m⁻² h⁻¹ in December 2010 and 44 ± 4 mmol m⁻² h⁻¹ in May 2011. We also monitored a suite of physical and chemical properties to help relate the rates at Sheltered Reef to published rates from other sites. While previous research shows that short-term field studies investigating the NCC–Ω_a relationship have differing results due to confounding factors, it is important to continue estimating NCC in different places, seasons, and years, in order to monitor changes in NCC versus Ω in space and time, and to ultimately resolve a broader understanding of this relationship.

     

Incorporating fine-scale seascape composition in an assessment of habitat quality for the giant sea anemone Stichodactyla gigantea in a coral reef shore zone

Habitat loss due to land reclamation often occurs in sandy coral reef shore zones. The giant sea anemone Stichodactyla gigantea, which harbors the false clown anemonefish Amphiprion ocellaris, both of which are potentially flagship species, inhabit these places. To assess habitat quality for S. gigantea, we examined correlative associations between the number and the body size of S. gigantea and the amount of habitat types in fine-scale seascape composition quantified from an enlarged section of a high-resolution (1/2,500) color aerial photograph of the shallow shore zone of Shiraho Reef, Ishigaki Island, Japan. This study confirmed that anemones were most abundant at the edges of dense seagrass beds characterized by shallow sandy bottoms, rock beds, and sparse seagrass beds, while they were less abundant in coral patch reefs. However, anemones inhabiting coral patch reefs were significantly larger and their rate of disappearance over 3 years was lower than those inhabiting other habitats. This suggests that coral patch reefs may be more suitable habitats supporting larger animals and greater persistence of S. gigantea. The visual census techniques applied here, combined with aerial photography and image-analysis software, may be useful as a simple analytical tool for local assessment of suitable habitats for relatively small-bodied marine fauna in shallow-water seascapes.     

Measuring recruitment of minute larvae in a complex field environment: The corallivorous nudibranch Phestilla sibogae (Bergh)

The nudibranch Phestilla sibogae feeds only on corals of the genus Porites. The nudibranch's minute (∼ 200 μm) larvae are specifically induced to settle and metamorphose by a chemical cue released by the coral, causing the larvae to recruit to reefs composed predominantly of Porites compressa. In this study, we investigated temporal and spatial patterns of recruitment of P. sibogae into coral reefs in Kane?ohe Bay, HI. We collected heads of P. compressa at 3-week intervals for 3 years, brought them to the laboratory and maintained them in aquaria fed with filtered seawater for 2 weeks, and then examined them for the presence of juvenile P. sibogae that had grown large enough to be seen. We found that P. sibogae recruits to the Porites reefs of Kane?ohe Bay sporadically and unpredictably throughout the year. Although most coral samples contained no or very few P. sibogae, three periods of intense recruitment (90-450 juvenile P. sibogae kg^− 1 of coral) were recorded, all in different seasons. Size-frequency analysis of recruits on the coral revealed high rates of post-settlement mortality in the field, most likely due to predation. Given the short pre-competent larval period of P. sibogae, the low rate of flushing of Kane?ohe Bay and the patterns of recruitment observed, we conclude that this population of P. sibogae is essentially a self-recruiting one. Two of the sampled reefs were characterized by unidirectional flow, allowing us to test a model of transport of larvae of P. sibogae responding to dissolved coral cue in turbulent, wavy flow. The model predicts that more larvae will be transported into upstream portions of a reef than into downstream portions, a prediction confirmed by analysis of the field-recruitment data. Furthermore, field releases of larval mimic particles also showed that most mimics landed in the upstream areas of reefs and down among the bases of coral branches, rather than at their tips.     

Wave exposure shapes reef community composition and recovery trajectories at a remote coral atoll

In a time of unprecedented ecological change, understanding natural biophysical relationships between reef resilience and physical drivers is of increasing importance. This study evaluates how wave forcing structures coral reef benthic community composition and recovery trajectories after the major 2015/2016 bleaching event in the remote Chagos Archipelago, Indian Ocean. Benthic cover and substrate rugosity were quantified from digital imagery at 23 fore reef sites around a small coral atoll (Salomon) in 2020 and compared to data from a similar survey in 2006 and opportunistic surveys in intermediate years. Cluster analysis and principal component analysis show strong separation of community composition between exposed (modelled wave exposure > 1000 J m⁻³) and sheltered sites (< 1000 J m⁻³) in 2020. This difference is driven by relatively high cover of Porites sp., other massive corals, encrusting corals, soft corals, rubble and dead table corals at sheltered sites versus high cover of pavement and sponges at exposed sites. Total coral cover and rugosity were also higher at sheltered sites. Adding data from previous years shows benthic community shifts from distinct exposure-driven assemblages and high live coral cover in 2006 towards bare pavement, dead Acropora tables and rubble after the 2015/2016 bleaching event. The subsequent recovery trajectories at sheltered and exposed sites are surprisingly parallel and lead communities towards their respective pre-bleaching communities. These results demonstrate that in the absence of human stressors, community patterns on fore reefs are strongly controlled by wave exposure, even during and after widespread coral loss from bleaching events.

     

The effect of microhabitat patch size on settlement differs among co-occurring coral reef fishes

Over small spatial scales, coral reefs represent a mosaic of suitable settlement microhabitat patches of varying size for late-stage larval reef fishes. Few studies have specifically examined how variation in patch size influences density of recently settled coral reef fishes (recruits). Using standardized units of coral rubble settlement substrate deployed on sandy bottom, we monitored the concurrent settlement of three reef fish taxa onto differently sized patches (0.28–1.68 m²) at 5-d intervals during a lunar settlement peak. We found marked differences among taxa in how recruit density scaled with patch size. Recruit density of a damselfish and a parrotfish decreased and increased, respectively, with the increase in patch size, while that of a wrasse was similar among patch sizes. Our results highlight the importance of the interaction between taxon-specific settlement behaviour and patch size in establishing initial spatial differences in density within and among coral reef fish taxa in a heterogeneous landscape.

     

Into the depth of population genetics: pattern of structuring in mesophotic red coral populations

Deep-sea reef-building corals are among the most conspicuous invertebrates inhabiting the hard-bottom habitats worldwide and are particularly susceptible to human threats. The precious red coral (Corallium rubrum, L. 1758) has a wide bathymetric distribution, from shallow up to 800 m depth, and represents a key species in the Mediterranean mesophotic reefs. Several studies have investigated genetic variability in shallow-water red coral populations, while geographic patterns in mesophotic habitats are largely unknown. This study investigated genetic variability of C. rubrum populations dwelling between 55 and 120 m depth, from the Ligurian to the Ionian Sea along about 1500 km of coastline. A total of 18 deep rocky banks were sampled. Colonies were analyzed by means of a set of microsatellite loci and the putative control region of the mitochondrial DNA. Collected data were compared with previous studies. Both types of molecular markers showed high genetic similarity between populations within the northern (Ligurian Sea and Tuscan Archipelago) and the southern (Tyrrhenian and Ionian seas) study areas. Variability in habitat features between the sampling sites did not affect the genetic variability of the populations. Conversely, the patchy distribution of suitable habitats affected populations’ connectivity within and among deep coral banks. Based on these results and due to the emphasis on red coral protection in the Mediterranean Sea by international institutions, red coral could be promoted as a ‘focal species’ to develop management plans for the conservation of deep coralligenous reefs, a reservoir of marine biodiversity.     

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.     

Configuration of small patch reefs and population abundance of a resident reef fish in a complex coral reef landscape

Habitat use by the resident coral reef anemonefish, Amphiprion frenatus, was examined in the complex coral reef landscape of Shiraho Reef, Ishigaki Island, Okinawa, Japan, using an enlarged color aerial photograph processed using image analysis software as an accurate field map. The anemonefish inhabit assemblages of the host sea anemone, Entacmaea quadricolor (clonal type), which inhabit various patch reefs in the back reef moat. We located all patch reefs inhabited by the host in the map based on snorkel observations: 297 anemonefish were found in 93 host assemblages in the study site of 2.9 ha. These patch reefs could be recognized by the reef colors, including the shadows (blackish color) in the photograph. Using image analysis software, the colors of the patch reefs were chosen and pixels with the same color values were regarded as potential habitat patches for the fish (PHPs). PHPs were 3D patch reefs (>0.5 m in height). Total areas (TA) and total perimeters (TP) of PHPs were measured in nine quadrats in the digitized aerial photograph. Host abundance and anemonefish abundance in a quadrat showed stronger correlations with the product of TA and TP of PHPs than TA alone. A site with numerous large 3D patch reefs (≥0.75 m² in situ) can be a better habitat for the fish than other sites consisting of several huge 3D patch reefs of the same total area. The methodology applied here may be useful for assessing the quality of habitats for small resident animals in shallow subtidal reefs.     

Large-scale bleaching of corals on the Great Barrier Reef

 The Great Barrier Reef (GBR) experienced its most intensive and extensive coral bleaching event on record in early 1998. Mild bleaching commenced in late January and intensified by late February/early March 1998. Broad-scale aerial surveys conducted of 654 reefs (∼23% of reefs on the GBR) in March and April 1998, showed that 87% of inshore reefs were bleached at least to some extent (>1% of coral cover) compared to 28% of offshore (mid- and outer-shelf) reefs. Of inshore reefs 67% had high levels of bleaching (>10% of coral) and 25% of inshore reefs had extreme levels of bleaching (>60% of coral). Fewer offshore reefs (14%) showed high levels of bleaching while none showed extreme levels of bleaching. Ground-truth surveys of 23 reefs, which experienced bleaching in intensities ranging from none to extreme, showed that the aerial survey data are likely to be underestimates of the true extent and intensity of bleaching on the GBR. The primary cause of this bleaching event is likely to be elevated sea temperature and solar radiation, exacerbated by lowered salinity on inshore and some offshore reefs in the central GBR. Accepted: 30 July 1998     

Coral recruitment and juvenile mortality as structuring factors for reef benthic communities in Biscayne National Park, USA

Coral communities of Biscayne National Park (BNP) on offshore linear bank-barrier reefs are depauperate of reef corals and have little topographic relief, while those on lagoonal patch reefs have greater coral cover and species richness despite presumably more stressful environmental regimes closer to shore. We hypothesized that differences in rates of coral recruitment and/or of coral survivorship were responsible for these differences in community structure. These processes were investigated by measuring: (1) juvenile and adult coral densities, and (2) size-frequency distributions of smaller coral size classes, at three pairs of bank- and patch-reefs distributed along the north-south range of coral reefs within the Park. In addition, small quadrats (0.25 m²) were censused for colonies <2 cm in size on three reefs (one offshore and one patch reef in the central park, and one intermediate reef at the southern end), and re-surveyed after 1 year. Density and size frequency data confirmed that large coral colonies were virtually absent from the offshore reefs, but showed that juvenile corals were common and had similar densities to those of adjacent bank and patch reefs. Large coral colonies were more common on inshore patch reefs, suggesting lower survivorship (higher mortality) of small and intermediate sized colonies on the offshore reefs. The more limited small-quadrat data showed similar survivorship rates and initial and final juvenile densities at all three sites, but a higher influx of new recruits to the patch reef site during the single annual study period. We consider the size-frequency data to be a better indicator of juvenile coral dynamics, since it is a more time-integrated measurement and was replicated at more sites. We conclude that lack of recruitment does not appear to explain the impoverished coral communities on offshore bank reefs in BNP. Instead, higher juvenile coral mortality appears to be a dominant factor structuring these communities. Accepted: 9 September 1999