Hurricanes and coral reefs: The intermediate disturbance hypothesis revisited

A review of research on the effects of hurricanes on coral reefs suggests that the intermediate disturbance hypothesis may be applicable to shallow reef zones dominated by branching or foliaceous coral species that are especially susceptible to mechanical damage from storms. Diversity (H') increases because of an increase in evenness following destruction or removal of the species that was monopolizing the space. The intermediate disturbance hypothesis as presented by Connell focuses on changes in number of species, but should be expanded to include diversity (H') and evenness. It should also be modified to incorporate changes in living cover and the time elapsed since disturbances of varying intensities. This hypothesis predicts that when cover is high, diversity will be low. However, research on coral reefs does not consistently demonstrate an inverse correlation of coral diversity, and coral cover. An increase in cover and decrease in diversity with depth would also be expected because deeper reef zones generally experience less disturbance. However, higher diversity (both H' and species richness) is often associated with deeper zones. The effects of hurricanes on coral reefs will depend on the temporal and spatial scales under consideration, the life history characteristics and morphology of the dominant species, the depth of the reef zone, the ecological history of the site, and the influence of any additional natural or human stresses.     

Fluorescent and skeletal density banding in Porites lutea from Papua New Guinea and Indonesia

Shallow water Porites lutea corals were collected along two transects normal to mainland shorelines, parallel to gradients in water quality: one, 7 km long, near Motupore Island in South Papua New Guinea, the other, 70 km long, from Jakarta Bay along the Pulau Seribu chain in the Java Sea. The corals were slabbed and studies were made of skeletal density bands as revealed by X-ray photography and fluorescent bands as revealed by ultraviolet light. Water quality measurements and rain-fall data were assembled for the two areas and related to skeletal banding patterns. For both areas, with increasing distance form mainland there is a decrease in overall brightness of fluorescence in corals and an increase in the contrast between bright and dull fluorescent bands. Fluorescence is bright, but seasonal banding is obscure in corals within about 2 km of stream mouths at Motopure and about 5 km of the coast in Jakarta Bay; this suggests that, despite low freshwater run-off during dry seasons, there are sufficient organic compounds which cause fluorescence in coral skeletons, to swamp seasonal effects. During the wet seasons, deluges of freshwater consequent on mainland rainfall of greater than about 150 mm/ month extend at least 7 km offshore in the Motupore area and perhaps tens of kilometres into Java Sea, giving distinctive bright and dull fluorescent banding in off-shore corals. The fluorescent banding pattern within corals on the Motupore reefs is similar in most corals along the transect and it correlates well with the Port Moresby monthly rainfall data. This relationship suggests that the same body (or bodies) of freshwater affect all reefs of the area during the wet season. The fluorescent banding in Java Sea corals does not show a precise correlation with either mainland or island monthly rainfall data; indeed the pattern of fluorescent banding on Pulau Seribu can only be matched in corals from reefs less than about 25 km apart. This suggests that in this area discrete water bodies carrying the relevant organic acids for coral fluorescence affect the fringing reefs on the chain of islands. Comparisons of fluorescent and density banding have revealed that for these areas, in general, periods of high freshwater run-off are times of deposition of less dense skeleton in Porites lutea corals.     

Coral Reef Community Composition in the Context of Disturbance History on the Great Barrier Reef,Australia

Much research on coral reefs has documented differential declines in coral and associated organisms. In order to contextualise this general degradation, research on community composition is necessary in the context of varied disturbance histories and the biological processes and physical features thought to retard or promote recovery. We conducted a spatial assessment of coral reef communities across five reefs of the central Great Barrier Reef, Australia, with known disturbance histories, and assessed patterns of coral cover and community composition related to a range of other variables thought to be important for reef dynamics. Two of the reefs had not been extensively disturbed for at least 15 years prior to the surveys. Three of the reefs had been severely impacted by crown-of-thorns starfish outbreaks and coral bleaching approximately a decade before the surveys, from which only one of them was showing signs of recovery based on independent surveys. We incorporated wave exposure (sheltered and exposed) and reef zone (slope, crest and flat) into our design, providing a comprehensive assessment of the spatial patterns in community composition on these reefs. Categorising corals into life history groupings, we document major coral community differences in the unrecovered reefs, compared to the composition and covers found on the undisturbed reefs. The recovered reef, despite having similar coral cover, had a different community composition from the undisturbed reefs, which may indicate slow successional processes, or a different natural community dominance pattern due to hydrology and other oceanographic factors. The variables that best correlated with patterns in the coral community among sites included the density of juvenile corals, herbivore fish biomass, fish species richness and the cover of macroalgae. Given increasing impacts to the Great Barrier Reef, efforts to mitigate local stressors will be imperative to encouraging coral communities to persist into the future.     

Maintenance of fish diversity on disturbed coral reefs

Habitat perturbations play a major role in shaping community structure; however, the elements of disturbance-related habitat change that affect diversity are not always apparent. This study examined the effects of habitat disturbances on species richness of coral reef fish assemblages using annual surveys of habitat and 210 fish species from 10 reefs on the Great Barrier Reef (GBR). Over a period of 11 years, major disturbances, including localised outbreaks of crown-of-thorns sea star (Acanthaster planci), severe storms or coral bleaching, resulted in coral decline of 46–96% in all the 10 reefs. Despite declines in coral cover, structural complexity of the reef framework was retained on five and species richness of coral reef fishes maintained on nine of the disturbed reefs. Extensive loss of coral resulted in localised declines of highly specialised coral-dependent species, but this loss of diversity was more than compensated for by increases in the number of species that feed on the epilithic algal matrix (EAM). A unimodal relationship between areal coral cover and species richness indicated species richness was greatest at approximately 20% coral cover declining by 3–4 species (6–8% of average richness) at higher and lower coral cover. Results revealed that declines in coral cover on reefs may have limited short-term impact on the diversity of coral reef fishes, though there may be fundamental changes in the community structure of fishes.     

The recovery of coral genetic diversity in the Sunda Strait following the 1883 eruption of Krakatau

Surveys of microsatellite variation show that genetic diversity has largely recovered in two reef-building corals, Pocillopora damicornis and Seriatopora hystrix (Scleractinia: Pocilloporidae), on reefs which were decimated by the eruption of the volcano Krakatau in 1883. Assignment methods and gene flow estimates indicate that the recolonization of Krakatau occurred mainly from the closest upstream reef system, Pulau Seribu, but that larval input from other regions has also occurred. This pattern is clearer in S. hystrix, which is traditionally the more dispersal-limited species. Despite these observed patterns of larval dispersal, self-recruitment appears to now be the most important factor in supplying larvae to coral populations in Krakatau. This suggests that the colonization of devastated reefs can occur quickly through larval dispersal; however, their survival requires local sources of larvae for self-recruitment. This research supports the observation that the recovery of genetic diversity in coral reef animals can occur on the order of decades and centuries rather than millennia. Conservation measures aimed at sustaining coral reef populations in Krakatau and elsewhere should include both the protection of upstream source populations for larval replenishment should disaster occur as well as the protection of large adult colonies to serve as local larval sources.     

Coral and sea urchin assemblage structure and interrelationships in Kenyan reef lagoons

Patterns of hard coral and sea urchin assemblage structure (species richness, diversity, and abundance) were studied in Kenyan coral reef lagoons which experienced different types of human resource use. Two protected reefs (Malindi and Watamu Marine National Parks) were protected from fishing and coral collection, but exposed to heavy tourist use. One reef (Mombasa MNP) received protection from fishermen for one year and was exploited for fish and corals prior to protection and was defined as a transitional reef. Three reefs (Vipingo, Kanamai, and Diani) were unprotected and experienced heavy fishing and some coral collection. Protected and unprotected reefs were distinct in terms of their assemblage structure with the transitional reef grouping with unprotected reefs based on relative and absolute abundance of coral genera. Protected reefs had slightly higher (p<0.01) coral cover (23.6 ± 8.3 % ± S.D.) than unprotected reefs (16.7 ± 8.5), but the transitional reef had the highest coral cover (30.8 ± 6.4) which increased by 250% since measured in 1987: largely attributable to a large increase inPorites nigrescens cover. Protected reefs had higher coral species richness and diversity and a greater relative abundance ofAcropora, Montipora andGalaxea than unprotected reefs. The transitional reef had high species richness, but lower diversity due to the high dominance ofPorites. Sea urchins showed the opposite pattern with highest diversity in most unprotected reefs. Coral cover, species richness, and diversity were negatively associated with sea urchin abundance, but the relative abundance ofPorites increased with sea urchin abundance to the point wherePorites composed >90% of the coral cover at sites with the highest sea urchin abundance. Effects of coral overcollection was only likely for the genusAcropora (staghorn corals). A combination of direct and indirect effects of human resource use may reduce diversity, species richness, and abundance of corals while increasing the absolute abundance of sea urchins and the relative cover ofPorites.     

Coral Reefs at the Northernmost Tip of Borneo: An Assessment of Scleractinian Species Richness Patterns and Benthic Reef Assemblages

The coral reefs at the northernmost tip of Sabah, Borneo will be established under a marine protected area: the Tun Mustapha Park (TMP) by the end of 2015. This area is a passage where the Sulu Sea meets the South China Sea and it is situated at the border of the area of maximum marine biodiversity, the Coral Triangle. The TMP includes fringing and patch reefs established on a relatively shallow sea floor. Surveys were carried out to examine features of the coral reefs in terms of scleractinian species richness, and benthic reef assemblages following the Reef Check substrate categories, with emphasis on hard coral cover. Variation in scleractinian diversity was based on the species composition of coral families Fungiidae (n = 39), Agariciidae (n = 30) and Euphylliidae (n = 15). The number of coral species was highest at reefs with a larger depth gradient i.e. at the periphery of the study area and in the deep South Banggi Channel. Average live hard coral cover across the sites was 49%. Only 7% of the examined reefs had > 75% hard coral cover, while the majority of the reef sites were rated fair (51%) and good (38%). Sites with low coral cover and high rubble fragments are evidence of blast fishing, although the observed damage appeared old. Depth was a dominant factor in influencing the coral species composition and benthic reef communities in the TMP. Besides filling in the information gaps regarding species richness and benthic cover for reef areas that were previously without any data, the results of this study together with information that is already available on the coral reefs of TMP will be used to make informed decisions on zoning plans for conservation priorities in the proposed park.     

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.     

Coral and sandstone reef-habitats in north-western Sri Lanka: patterns in the distribution of coral communities

Coral and sandstone reefs cover a significant part ofSri Lanka‘s continental shelf and contain incomparison unique reef structures. Despite this, reefsin this region of the northern Indian Ocean havereceived little research attention. In an attempt tobetter understand these ecosystems and their innatecharacter, this study describes the variety of reeftypes and habitats that are found in this area. Thestudy concentrated on four major reef areas: the BarReef Marine Sanctuary (BRMS), Kandakuliya Reefs,Talawila Reef, and Mampuri Reef. These reefs showedapparent differences in habitat structure in terms ofthe proportion coral cover, coral species compositionand structural complexity. Two reef types were presentwithin the continental shelf of BRMS: coral andsandstone patch-reefs. Acropora was the mostdominant coral genera however in total 118madreporarian species and 50 coral genera wererecorded in the sanctuary. Distinct habitats wereidentified within reef types including shallow reefflat, shallow patch reef, deep reef flat and Porites dome habitats for the coral-reef patcheswhile the sandstone-reef patches were divided intostructured and flat sandstone reef habitats.Kandakuliya Reef south of BRMS was by large dominatedby coral rubble. Talawila Reef and Mampuri Reef showedunique structures with the former being dominated bymassive corals and the latter mainly containingsandstone structures. In addition to factors such asbio-erosion, sedimentation, hydrodynamics, andrecruitment or colonisation processes, some reefs wereclearly under significant direct human impact whichappeared to play a dominant role in habitatstructuring. However, type and degree of humandisturbance varied among the reefs. Habitat alterationat Kandakuliya Reef was the result of intense fishingusing destructive fishing methods. Talawila Reef andMampuri Reef was also influenced by fishing activitiesthough reef structure seemed less affected. This revised version was published online in July 2006 with corrections to the Cover Date.     

Local and Regional Impacts of Pollution on Coral Reefs along the Thousand Islands North of the Megacity Jakarta,Indonesia

Worldwide, coral reefs are challenged by multiple stressors due to growing urbanization, industrialization and coastal development. Coral reefs along the Thousand Islands off Jakarta, one of the largest megacities worldwide, have degraded dramatically over recent decades. The shift and decline in coral cover and composition has been extensively studied with a focus on large-scale gradients (i.e. regional drivers), however special focus on local drivers in shaping spatial community composition is still lacking. Here, the spatial impact of anthropogenic stressors on local and regional scales on coral reefs north of Jakarta was investigated. Results indicate that the direct impact of Jakarta is mainly restricted to inshore reefs, separating reefs in Jakarta Bay from reefs along the Thousand Islands further north. A spatial patchwork of differentially degraded reefs is present along the islands as a result of localized anthropogenic effects rather than regional gradients. Pollution is the main anthropogenic stressor, with over 80% of variation in benthic community composition driven by sedimentation rate, NO₂, PO₄ and Chlorophyll a. Thus, the spatial structure of reefs is directly related to intense anthropogenic pressure from local as well as regional sources. Therefore, improved spatial management that accounts for both local and regional stressors is needed for effective marine conservation.     

Depth distributions of coral reef fishes: the influence of microhabitat structure,settlement, and post-settlement processes

Many coral reef fishes have restricted depth ranges that are established at settlement or soon after, but the factors limiting these distributions are largely unknown. This study examines whether the availability of microhabitats (reef substrata) explains depth limits, and evaluates whether juvenile growth and survival are lower beyond these limits. Depth-stratified surveys of reef fishes at Kimbe Bay (Papua New Guinea) showed that the abundance of new settlers and the cover of coral substrata differed significantly among depths. A field experiment investigated whether settling coral reef fishes preferred particular depths, and whether these depth preferences were dependent on microhabitat. Small patch reefs composed of identical coral substrata were set up at five depths (3, 6, 10, 15 and 20 m), and settlement patterns were compared to those on unmanipulated reef habitat at the same five depths. For all species, settlement on patch reefs differed significantly among depths despite uniform substratum composition. For four of the six species tested, depth-related settlement patterns on unmanipulated habitat and on patch reefs did not differ, while for the other two, depth ranges were greater on the patch reefs than on unmanipulated habitat. A second experiment examined whether depth preferences reflected variation in growth and survival when microhabitat was similar. Newly settled individuals of Chrysiptera parasema and Dascyllus melanurus were placed, separately, on patch reefs at five depths (as above) and their survival and growth monitored. D. melanurus, which is restricted to shallow depths, had highest survival and growth at the shallowest depth. Depth did not affect either survival or growth of C. parasema, which has a broader depth range than D. melanurus (between 6 and 15 m). This suggests that the fitness costs potentially incurred by settling outside a preferred depth range may depend on the strength of the depth preference.     

Otolith growth of Springer's demoiselle, Chrysiptera springeri (Pomacentridae, Allen & Lubbock), on a protected and non-protected coral reef

The structural complexity of coral reefs is important for their function as shelter and feeding habitats for coral reef fishes, but physical disturbance by human activities often reduce complexity of the reefs by selectively destroying fragile and more complex coral species. The damselfish Springer's demoiselle Chrysiptera springeri primarily utilize complex coral heads for shelter and are hence vulnerable to human disturbance. In order to evaluate the potential effect of habitat degradation on juvenile fish growth, coral reef cover, fish age at settling and otolith growth, juvenile Springer's demoiselle was investigated on a protected and non‐protected coral reef in Darvel Bay, Borneo. The protected reef had higher coverage of complex branching corals and exhibited a more complex 3‐dimensional structure than the non‐protected reef. Springer's demoiselle settled at the same age on non‐protected and protected reefs. The growth rates of the otoliths from Springer's demoiselle were similar during the pre‐settlement period on the two reefs (manova , P > 0.05), but from age 20 to 48 days (post‐settlement period) the otolith growth rate of juveniles on the non‐protected reef was reduced compared to those from the protected reef (manova , P = 0.017). However, the differences in the otolith size, and by inference, fish size, after 48 days were small. The small effect of habitat degradation on growth is likely related to the fact that the Springer's demoiselles collected on the non‐protected reef were associated with the few remaining complex coral heads. Increased foraging‐predation tradeoffs on the non‐protected reef may decrease food intake and growth of juvenile Springer's demoiselle, but the main effect of habitat degradation on their abundance is likely to be related to lack of suitable shelter, and consequently reduced carrying capacity, on disturbed reefs.     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Spatial resilience of the Great Barrier Reef under cumulative disturbance impacts

In the face of increasing cumulative effects from human and natural disturbances, sustaining coral reefs will require a deeper understanding of the drivers of coral resilience in space and time. Here we develop a high‐resolution, spatially explicit model of coral dynamics on Australia's Great Barrier Reef (GBR). Our model accounts for biological, ecological and environmental processes, as well as spatial variation in water quality and the cumulative effects of coral diseases, bleaching, outbreaks of crown‐of‐thorns starfish (Acanthaster cf. solaris), and tropical cyclones. Our projections reconstruct coral cover trajectories between 1996 and 2017 over a total reef area of 14,780 km², predicting a mean annual coral loss of ?0.67%/year mostly due to the impact of cyclones, followed by starfish outbreaks and coral bleaching. Coral growth rate was the highest for outer shelf coral communities characterized by digitate and tabulate Acropora spp. and exposed to low seasonal variations in salinity and sea surface temperature, and the lowest for inner‐shelf communities exposed to reduced water quality. We show that coral resilience (defined as the net effect of resistance and recovery following disturbance) was negatively related to the frequency of river plume conditions, and to reef accessibility to a lesser extent. Surprisingly, reef resilience was substantially lower within no‐take marine protected areas, however this difference was mostly driven by the effect of water quality. Our model provides a new validated, spatially explicit platform for identifying the reefs that face the greatest risk of biodiversity loss, and those that have the highest chances to persist under increasing disturbance regimes.     

Environmental factors associated with the spatial distribution of crustose coralline algae on the Great Barrier Reef

Crustose coralline algae (CCA) fulfill two key functional roles in coral reef ecosystems: they contribute significantly to reef calcification, and they induce larval settlement of many benthic organisms. Percentage cover of CCA, and environmental conditions, were visually estimated on 144 reefs of the Great Barrier Reef between 10 and 24° latitude S. Reefs were located across the shelf and ranged from turbid near-shore reefs close to rivers to clean-water reefs hundreds of kilometers from coastal influences. On each reef, two sites were surveyed between 0.5 and 18 m depth. Strong cross-shelf trends occurred in cover of CCA, amount of sediment deposited, water clarity, and slope angle. Relative distance across the shelf and sedimentation jointly explained 84% of variation in CCA cover. Three regions running parallel to the shore were identified, with a mean CCA cover of <1% on the inner third of the shelf, and >20% cover on the outer half of the shelf, with a narrow transition region between the two. Within each region, the cover of CCA was unrelated to distance across the shelf, but was related to the sedimentary environment, being relatively higher on reefs with low sediment deposits. On the inner third of the shelf, the most sediment-exposed reefs were unsuitable habitats for CCA. The inverse relationship between CCA and sediment has implications for the recruitment of CCA-specialised organisms, and for rates of reef calcification.     

Effect of Phase Shift from Corals to Zoantharia on Reef Fish Assemblages

Consequences of reef phase shifts on fish communities remain poorly understood. Studies on the causes, effects and consequences of phase shifts on reef fish communities have only been considered for coral-to-macroalgae shifts. Therefore, there is a large information gap regarding the consequences of novel phase shifts and how these kinds of phase shifts impact on fish assemblages. This study aimed to compare the fish assemblages on reefs under normal conditions (relatively high cover of corals) to those which have shifted to a dominance of the zoantharian Palythoa cf. variabilis on coral reefs in Todos os Santos Bay (TSB), Brazilian eastern coast. We examined eight reefs, where we estimated cover of corals and P. cf. variabilis and coral reef fish richness, abundance and body size. Fish richness differed significantly between normal reefs (48 species) and phase-shift reefs (38 species), a 20% reduction in species. However there was no difference in fish abundance between normal and phase shift reefs. One fish species, Chaetodon striatus, was significantly less abundant on normal reefs. The differences in fish assemblages between different reef phases was due to differences in trophic groups of fish; on normal reefs carnivorous fishes were more abundant, while on phase shift reefs mobile invertivores dominated.     

Seasonal trends and fish-habitat associations around Pari Island, Indonesia: setting a baseline for environmental monitoring

Indonesia is the world??s richest country regarding reef fish diversity. Nevertheless, the reef ichthyofauna of the Indonesian Archipelago remains poorly known, primarily due to a lack of sampling. Coral reefs in the Kepulauan Seribu Marine National Park close to the Indonesian capital Jakarta are under threat by many destructive activities that trigger a loss of habitat and species diversity. This communication: (1) describes the reef fish community structure from three distinct reef habitats in the Pari Island group dominated by Acropora branching corals (ACB), foliose corals (CF) and massive corals (CM), using a number of community properties such as numerical abundance, species richness, diversity, and multivariate similarity; (2) examines the temporal variation of the fish community from the three habitats; and (3) discusses possible implications for the monitoring of qualitative changes in coral reef systems on small islands. During this study, a total of 13 536 individual fishes were counted, representing 205 species belonging to 36 families. In terms of species richness, Pomacentridae was the dominant fish family in ACB and CF sites (40?% and 48.6?%, respectively), and Labridae (27.4?%) was the dominant family in the CM plots. The most species-rich habitat was ACB with 125 species (with Amblyglyphidodon curacao as the most characteristic species), followed by CM and CF with 117 (Thalassoma lunare) and 79 species (Pomacentrus alexanderae), respectively. Average Shannon-Wiener diversity (ln basis) ranged from 2.0?C2.9 (ACB), 2.4?C3.1 (CF), and 2.1?C3.0 (CM), with no significant difference between growth forms. Abundance, species richness and diversity showed significant seasonal variability, but the effects differed between habitats. Multivariate analysis of the reef fish community was able to detect significant differences between species composition and diversity of the reef fish community between sites with different coral growth forms at Pari Island, both when based on species abundances and when aggregated according to trophic categories. It thus constitutes a useful tool to detect qualitative differences of the species-rich Indonesian coral reef ecosystems.     

Mesophotic depths as refuge areas for fishery-targeted species on coral reefs

Coral reefs are subjected to unprecedented levels of disturbance with population growth and climate change combining to reduce standing coral cover and stocks of reef fishes. Most of the damage is concentrated in shallow waters (<30 m deep) where humans can comfortably operate and where physical disturbances are most disruptive to marine organisms. Yet coral reefs can extend to depths exceeding 100 m, potentially offering refuge from the threats facing shallower reefs. We deployed baited remote underwater stereo-video systems (stereo-BRUVs) at depths of 10–90 m around the southern Mariana Islands to investigate whether fish species targeted by fishing in the shallows may be accruing benefits from being at depth. We show that biomass, abundance and species richness of fishery-targeted species increased from shallow reef areas to a depth of 60 m, whereas at greater depths, a lack of live coral habitat corresponded to lower numbers of fish. The majority of targeted species were found to have distributions that ranged from shallow depths (10 m) to depths of at least 70 m, emphasising that habitat, not depth, is the limiting factor in their vertical distribution. While the gradient of abundance and biomass versus depth was steepest for predatory species, the first species usually targeted by fishing, we also found that fishery-targeted herbivores prevailed in similar biomass and species richness to 60 m. Compared to shallow marine protected areas, there was clearly greater biomass of fishery-targeted species accrued in mesophotic depths. Particularly some species typically harvested by depth-limited fishing methods (e.g., spearfishing), such as the endangered humphead wrasse Cheilinus undulatus, were found in greater abundance on deeper reefs. We conclude that mesophotic depths provide essential fish habitat and refuge for fishery-targeted species, representing crucial zones for fishery management and research into the resilience of disturbed coral reef ecosystems.     

Multiple disturbances and the global degradation of coral reefs: are reef fishes at risk or resilient?

Increased frequency of disturbances and anthropogenic activities are predicted to have a devastating impact on coral reefs that will ultimately change the composition of reef associated fish communities. We reviewed and analysed studies that document the effects of disturbance‐mediated coral loss on coral reef fishes. Meta‐analysis of 17 independent studies revealed that 62% of fish species declined in abundance within 3 years of disturbances that resulted in >10% decline in coral cover. Abundances of species reliant on live coral for food and shelter consistently declined during this time frame, while abundance of some species that feed on invertebrates, algae and/or detritus increased. The response of species, particularly those expected to benefit from the immediate loss of coral, is, however, variable and is attributed to erratic replenishment of stocks, ecological versatility of species and sublethal responses, such as changes in growth, body condition and feeding rates. The diversity of fish communities was found to be negatively and linearly correlated to disturbance‐mediated coral loss. Coral loss >20% typically resulted in a decline in species richness of fish communities, although diversity may initially increase following small declines in coral cover from high coverage. Disturbances that result in an immediate loss of habitat complexity (e.g. severe tropical storms), have a greater impact on fishes from all trophic levels, compared with disturbances that kill corals, but leave the reef framework intact (e.g. coral bleaching and outbreaks of Acanthaster planci). This is most evident among small bodied species and suggests the long‐term consequences of coral loss through coral bleaching and crown‐of‐thorn starfish outbreaks may be much more substantial than the short‐term effects currently documented.     

Soft coral abundance on the central Great Barrier Reef: effects of Acanthaster planci, space availability, and aspects of the physical environment

 The distribution and abundance of soft coral genera on reefs of the central Great Barrier Reef was investigated in relation to reef position, recent history of disturbance, wave exposure, substratum slope and depth. Eighty-five 25 m long transects were surveyed at 10 m depth on windward sides of 14 mid- and outer-shelf reefs. A further 75 transects in different zones on one mid-shelf reef (Davies Reef) between 5 and 30 m depth were investigated. The crown-of-thorns starfish Acanthaster planci had caused large-scale mortality of scleractinians on eight of these reefs five to ten years prior to the study, and as a result, scleractinian cover was only 35–55% of that on the six unimpacted reefs. On the impacted reefs, stony corals with massive and encrusting growths form had smaller average colony diameters but similar or slightly lower numerical abundance. In contrast, mean colony size, cover and abundance of branching stony corals showed no difference between impacted and unimpacted reefs. Twenty-four genera of soft corals (in eight families) were recorded, and none showed different abundance or cover in areas of former A. planci impact, compared to unaffected sites. Similarly, no difference was detected among locations in the numbers or area cover of sponges, tunicates, zoanthids, Halimeda or other macro-algae. Mean soft coral cover was 2 to 5% at 10 m on sheltered mid-shelf reefs, and 12 to 17% on more current-exposed reefs. Highest cover and abundances generally occurred on platforms of outer-shelf reefs exposed to relatively strong currents but low wave energy. On Davies Reef, cover and colony numbers of the families Nephtheidae and Xeniidae were low within the zone of wave impact, in flow-protected bays and lagoons, on shaded steep slopes, and at depths above 10 and below 25 m. In contrast, distributions of genera of the family Alcyoniidae were not related to these physical parameters. The physical conditions of a large proportion of habitats appear “sub-optimal” for the fastest growing taxa, possibly preventing an invasion of the cleared space. Thus, in the absence of additional stress these shallow-water fore-reef zones appear sufficiently resilient to return to their pre-outbreak state of scleractinian dominance. Accepted: 20 August 1996