Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

A geological perspective on the degradation and conservation of western Atlantic coral reefs

Continuing coral‐reef degradation in the western Atlantic is resulting in loss of ecological and geologic functions of reefs. With the goal of assisting resource managers and stewards of reefs in setting and measuring progress toward realistic goals for coral‐reef conservation and restoration, we examined reef degradation in this region from a geological perspective. The importance of ecosystem services provided by coral reefs—as breakwaters that dissipate wave energy and protect shorelines and as providers of habitat for innumerable species—cannot be overstated. However, the few coral species responsible for reef building in the western Atlantic during the last approximately 1.5 million years are not thriving in the 21st century. These species are highly sensitive to abrupt temperature extremes, prone to disease infection, and have low sexual reproductive potential. Their vulnerability and the low functional redundancy of branching corals have led to the low resilience of western Atlantic reef ecosystems. The decrease in live coral cover over the last 50 years highlights the need for study of relict (senescent) reefs, which, from the perspective of coastline protection and habitat structure, may be just as important to conserve as the living coral veneer. Research is needed to characterize the geological processes of bioerosion, reef cementation, and sediment transport as they relate to modern‐day changes in reef elevation. For example, although parrotfish remove nuisance macroalgae, possibly promoting coral recruitment, they will not save Atlantic reefs from geological degradation. In fact, these fish are quickly nibbling away significant quantities of Holocene reef framework. The question of how different biota covering dead reefs affect framework resistance to biological and physical erosion needs to be addressed. Monitoring and managing reefs with respect to physical resilience, in addition to ecological resilience, could optimize the expenditure of resources in conserving Atlantic reefs and the services they provide.     

A new apparatus for force measurement in marine bioadhesion

In an experimental study on the effect of parrotfish (probably Scarus taeniurus) grazing on the structure of benthic reef communities, fishes in densities of 0.6 to 1.5 parrotfish per m² or 9 to 17 g wet weight of fish per m² of feeding surface were found to have an optimum effect, resulting in the greatest benthic species richness and biomass on 2-dimensional surfaces. The presence of refuges (3-dimensional habitats), however, has a greater impact on bemthic community structure (number of species and biomass) than does just the density of parrotfish in such an experimental system. Coral recruitment is enhanced by the presence of refuges and, like coralline algae, is more successful under increased grazing pressure. These optimum densities of parrotfishes relate well to observed field densities where, in a collection from a Hawaiian patch reef, there were 1.1 fish or 10.8 g wet weight of parrotfish per square meter of collection area. The success of coralline algae and corals under high grazing pressure may have important consequences for the stability and structure of modern coral reefs.     

Influence of dead coral substrate morphology on patterns of juvenile coral distribution

This study examines the abundances of three morphological categories of juvenile corals (massive, branching and encrusting) on two different types of natural substratum, dead massive and dead branching corals. The overall results show that the morphological characteristics of dead coral substratum have a significant influence on the coral recruitment patterns with respect to the morphology of the recruits: juvenile corals of massive and branching types were more abundant on substrates of corresponding morphology. The results obtained from this study suggest that dead coral might attract coral larvae that are morphologically similar. On the other hand, it may be the result of post-settlement mortality. Whatever the mechanism shaping the patterns is, it seems that the physical morphology of the dead coral substrate has a significant influence on the coral recruit assemblage. Hence, we suggest that substrate morphology can be an important qualitative factor for coral settlement and a possible determinant of community structure.     

Predator biomass,prey density,and species composition effects on group size in recruit coral reef fishes

Group incidence and size are described for recruit parrotfishes, wrasses, and damselfishes on Hawaiian reefs over 3 years (2006–2008) at sites spanning the archipelago (20–28°N, 155–177°W). Coral-poor and coral-rich areas were surveyed at sites with both low (Hawaii Island) and high (Midway Atoll) predator densities, facilitating examination of relations among predator and recruit densities, habitat, and group metrics. Predator and recruit densities varied spatially and temporally, with a sixfold range in total recruit densities among years. Group (≥2 recruits) metrics varied with time and tracked predator and recruit densities and the proportion of schooling species. Groups often included heterospecifics whose proportion increased with group size. A non-saturating relationship between group size and recruit density suggests that the anti-predator benefits of aggregation exceeded competitive costs. Grouping behavior may have overarching importance for recruit survival–even at high recruit densities–and merits further study on Hawaiian reefs and elsewhere.     

Positive and Negative Effects of a Threatened Parrotfish on Reef Ecosystems

Species that are strong interactors play disproportionately important roles in the dynamics of natural ecosystems. It has been proposed that their presence is necessary for positively shaping the structure and functioning of ecosystems. We evaluated this hypothesis using the case of the world's largest parrotfish (Bolbometopon muricatum), a globally imperiled species. We used direct observation, animal tracking, and computer simulations to examine the diverse routes through which B. muricatum affects the diversity, dispersal, relative abundance, and survival of the corals that comprise the foundation of reef ecosystems. Our results suggest that this species can influence reef building corals in both positive and negative ways. Field observation and simulation outputs indicated that B. muricatum reduced the abundance of macroalgae that can outcompete corals, but they also feed directly on corals, decreasing coral abundance, diversity, and colony size. B. muricatum appeared to facilitate coral advancement by mechanically dispersing coral fragments and opening up bare space for coral settlement, but they also damaged adult corals and remobilized a large volume of potentially stressful carbonate sediment. The impacts this species has on reefs appears to be regulated in part by its abundance—the effects of B. muricatum were more intense in simulation scenarios populated with high densities of these fish. Observations conducted in regions with high and low predator (e.g., sharks) abundance generated results that are consistent with the hypothesis that these predators of B. muricatum may play a role in governing their abundance; thus, predation may modulate the intensity of the effects they have on reef dynamics. Overall our results illustrate that functionally unique and threatened species may not have universally positive impacts on ecosystems and that it may be necessary for environmental managers to consider the diverse effects of such species and the forces that mediate the strength of their influence. Efectos Positivos y Negativos de un Pez Loro Amenazado Sobre Ecosistemas Arrecifales     

Importance of macroalgal fields as coral reef fish nursery habitat in north-west Australia

Macroalgal fields are a feature of the shallow tropical benthos, yet their importance for coral reef fish population dynamics remains poorly understood. The abundance of fish recruits was recorded using underwater visual census at six macroalgal and 11 coral reef sites in the Montebello and Barrow Islands. Surveys identified 6,935 individual recruit fish from 105 species, 54 genera and 20 families. Of these, 1,401 recruits from 48 species, 31 genera and 14 families were observed in macroalgal sites. Sixteen of the 105 recruit species (15.2 %) were observed exclusively at macroalgal sites. Forty-two (87.5 %) of these species have been observed as adults on adjacent coral reefs. Species composition of fish recruits differed significantly between the two habitats. Corallivore, small omnivore and zooplanktivore recruits had significantly higher numbers in the coral sites, while the results clearly demonstrate that juveniles, within the genera Lethrinus and Choerodon, as well as large algal croppers, are predominantly found at macroalgal (74–100 %) rather than coral-dominated sites. High-canopy macroalgae cover was positively correlated with abundance of these taxa, particularly Lethrinids (r ² = 0.40). This study is the first to highlight the important attributes of tropical macroalgal fields and suggests that they have a similar role to seagrass meadows as essential juvenile habitat, thus warranting greater attention in conservation planning and ecological studies.     

Marine reserves,fisheries ban,and 20 years of positive change in a coral reef ecosystem

By 2004, Belize was exhibiting classic fishing down of the food web. Groupers (Serranidae) and snappers (Lutjanidae) were scarce and fisheries turned to parrotfishes (Scarinae), leading to a 41% decline in their biomass. Several policies were enacted in 2009–2010, including a moratorium on fishing parrotfish and a new marine park with no-take areas. Using a 20-year time series on reef fish and benthos, we evaluated the impact of these policies approximately 10 years after their implementation. Establishment of the Southwater Caye Marine Reserve led to a recovery of snapper at 2 out of 3 sites, but there was no evidence of recovery outside the reserve. Snapper populations in an older reserve continued to increase, implying that at least 9 years is required for their recovery. Despite concerns over the feasibility of banning parrotfish harvest once it has become a dominant fin fishery, parrotfishes returned and exceeded biomass levels prior to the fishery. The majority of these changes involved an increase in parrotfish density; species composition and adult body size generally exhibited little change. Recovery occurred equally well in reserves and areas open to other forms of fishing, implying strong compliance. Temporal trends in parrotfish grazing intensity were strongly negatively associated with the cover of macroalgae, which by 2018 had fallen to the lowest levels observed since measurements began in 1998. Coral populations remained resilient and continued to exhibit periods of net recovery after disturbance. We found that a moratorium on parrotfish harvesting is feasible and appears to help constrain macroalgae, which can otherwise impede coral resilience.     

The impact of exploiting grazers (Scaridae) on the dynamics of Caribbean coral reefs.

Coral reefs provide a number of ecosystem services including coastal defense from storms, the generation of building materials, and fisheries. It is increasingly clear that the management of reef resources requires an ecosystem approach in which extractive activities are weighed against the needs of the ecosystem and its functions rather than solely those of the fishery. Here, I use a spatially explicit simulation model of a Caribbean coral reef to examine the ecosystem requirements for grazing which is primarily conducted by parrotfishes (Scaridae). The model allows the impact of fishing grazers to be assessed in the wider context of other ecosystem processes including coral-algal competition, hurricanes, and mass extinction of the herbivorous urchin Diadema antillarum. Using a new analytical model of scarid grazing, it is estimated that parrotfishes can only maintain between 10% and 30% of a structurally complex forereef in a grazed state. Predictions from this grazing model were then incorporated into a broader simulation model of the ecosystem. Simulations predict that scarid grazing is unable to maintain high levels of coral cover (> or = 30%) when severe hurricanes occur on a decadal basis, such as occurs in parts of the northern Caribbean. However, reefs can withstand such intense disturbance when grazing is undertaken by both scarids and the urchin Diadema. Scarid grazing is predicted to allow recovery from hurricanes when their incidence falls to 20 years or less (e.g., most of Central and South America). Sensitivity analyses revealed that scarid grazing had the most acute impact on model behavior, and depletion led to the emergence of a stable, algal-dominated community state. Under conditions of heavy grazer depletion, coral cover was predicted to decline rapidly from an initial level of 30% to less than 1% within 40 years, even when hurricane frequency was low at 60 years. Depleted grazers caused a population bottleneck in juvenile corals in which algal overgrowth caused elevated levels of postsettlement mortality and resulted in a bimodal distribution of coral sizes. Several new hypotheses were generated including a region-wide change in the spatial heterogeneity of coral reefs following extinction of Diadema. The management of parrotfishes on Caribbean reefs is usually approached implicitly through no-take marine reserves. The model predicts that depletion of grazers in nonreserve areas can severely limit coral accretion. Other studies have shown that low coral accretion can reduce the structural complexity and therefore quality of the reef habitat for many organisms. A speculative yet rational inference from the model is that failure to manage scarid populations outside reserves will have a profoundly negative impact on the functioning of the reserve system and status of non-reserve reefs.     

Ecological versatility and the decline of coral feeding fishes following climate driven coral mortality

Coral reefs are under threat due to climate-mediated coral mortality, which affects some reef coral genera more severely than others. The impact this has on coral reef fish is receiving increasing attention, with one focal area assessing impacts on fish that feed directly on live coral. It appears that the more specialised a species of corallivore, the more susceptible it is to coral declines. However data are sparse for the Indian Ocean, and little is known about why some corals are preferentially fed upon over others. Here I assess feeding specialisation in three species of coral feeding butterflyfish in the Chagos Archipelago, central Indian Ocean, assess the food quality of the coral genera they target and document patterns of decline in the Seychelles following a severe coral mortality event. Cheatodon trifascialis was the most specialised coral feeder, preferentially selecting for Acropora corals, however, when Acropora was scarce, individuals showed considerable feeding plasticity, particularly for the dominant Pocillopora corals. C. trifasciatus also preferentially fed on Acropora corals, but fed on a much more diverse suite of corals and also displayed some selectivity for Porites. C. auriga is a facultative corallivore and consumed ∼55% live coral, which lies within the wide range of coral dependence reported for this species. C:N ratio analysis indicated Lobophyllia and Acropora have the highest food quality, with Pocillopora having the lowest, which conforms with diet selection of corallivores and helps explain preferential feeding. Obligate specialist feeders displayed the greatest declines through coral mortality in the Seychelles with obligate generalists also declining substantially, but facultative feeders showing little change. Clearly a greater understanding of the species most vulnerable to disturbance, their habitat requirements and the functional roles they play will greatly assist biodiversity conservation in a changing climate.     

Acute effects of removing large fish from a near-pristine coral reef

Large animals are severely depleted in many ecosystems, yet we are only beginning to understand the ecological implications of their loss. To empirically measure the short-term effects of removing large animals from an ocean ecosystem, we used exclosures to remove large fish from a near-pristine coral reef at Palmyra Atoll, Central Pacific Ocean. We identified a range of effects that followed from the removal of these large fish. These effects were revealed within weeks of their removal. Removing large fish (1) altered the behavior of prey fish; (2) reduced rates of herbivory on certain species of reef algae; (3) had both direct positive (reduced mortality of coral recruits) and indirect negative (through reduced grazing pressure on competitive algae) impacts on recruiting corals; and (4) tended to decrease abundances of small mobile benthic invertebrates. Results of this kind help advance our understanding of the ecological importance of large animals in ecosystems.     

Sabaki River sediment load and coral stress: correlation between sediments and condition of the Malindi-Watamu reefs in Kenya (Indian Ocean)

Sediment discharges from rivers have a negative impact on coral reef ecosystems. Indicators of coral decline measured in the present study were: (1) injury to living stony corals; (2) soft coral cover; and (3) bare rocky substrate suitable for colonization by corals. The relationship between these indicators and the distribution of terrigenous sediment was studied for the Malindi-Watamu fringing reef complex along the Kenyan coast off East Africa during 1982 and 1983. Decline of this reef had been repeatedly noted during the preceding decade. The influence of terrigenous sediment from the Sabaki River appears to be strongest in the Watamu area in the south and in the northern-most part of the Malindi reef area. Correlations, between each of the above three coral stress response indicators, on the one hand, and quantitative indicators of sediment loading, on the other hand, were not clear. However, a combined coral stress indicator involving all three factors was shown to have a clear relationship with terrigenous sediment loading and provided a rapid means of field evaluation of the effects of sediment stress on stony corals. Values for the combined coral stress indicator were found to increase in proportion to increasing values of terrigenous sediment loads in both study areas. A higher coral stress indicator value means a high proportion of injured or algae infested corals, and/or a high soft coral cover, and/or a high proportion of rocky substrate suitable for, but unoccupied by, living corals.     

Motile cryptofauna associated with live and dead coral substrates: implications for coral mortality and framework erosion

Coral reef cryptofauna are a diverse group of metazoan taxa that live within intra- and inter-skeletal voids formed by framework structures. Despite a hypothesized high biomass and numerous trophic roles, they remain uncharacterized relative to exposed reef communities. Motile cryptofauna were sampled from live coral colonies and dead frameworks typifying four successive levels of degradation on an eastern Pacific pocilloporid reef. Abundances and biomass were higher on live versus dead corals habitats. The density of cryptofauna per volume substrate was highest on dead coral frameworks of intermediate degradation, where complex eroded substrates provide abundant shelters. These data have important and far-reaching ramifications for how the diverse multispecies assemblages that are reef ecosystems will respond to anthropogenic stressors such as those associated with climate change. Extreme levels of coral mortality, bioerosion, and habitat destruction will lead to impairment and eventually loss of ecosystem functions.     

Direct versus indirect methods of quantifying herbivore grazing impact on a coral reef

Two methods were used to assess the grazing impact of roving herbivorous fishes across a coral reef depth gradient within Pioneer Bay, Orpheus Island, Great Barrier Reef. The first technique employed was a method traditionally used to quantify herbivory on coral reefs via the (indirect) inference of herbivore impact from biomass estimates and reported feeding rates. The second method (one of a range of direct approaches) used remote underwater video cameras to film the daily feeding activity of roving herbivores in the absence of divers. Both techniques recorded similar patterns and relative levels of herbivore biomass across five reef zones at the study site. Indirect estimates of the grazing impact across the reef depth gradient of the three predominant species of herbivore broadly coincided with levels quantified directly by remote underwater video, indicating that, to a large extent, presence does correspond to function. However, the video data suggested that, for individual species in particular reef zones, the absolute level of impact may be less than that inferred from presence. In the case of the parrotfish Scarus rivulatus, the video recordings suggested that, at the reef crest, an average of 52% (±18 SE) of each m² area of reef would be grazed each month, compared with an area of 109% (±41 SE) suggested by inferring grazing activity from presence alone. Potential biases associated with remote video recorders may explain some of the discrepancy between values. Overall, the results suggest that, for some fish groups, the indirect method of inferring function from presence can provide a good indication of relative levels of herbivore impact across a coral reef. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Dynamics of a coral reef community after mass mortality of branching <Emphasis Type="Italic">Acropora</Emphasis> corals and an outbreak of anemones

Dynamics of a coral reef community at Tiao-Shi Reef, southern Taiwan were studied using permanent transects to examine coral recovery and successive cascades to collapse stage resulting from chronic anthropogenic impacts and typhoons. Three distinct zones were recognized within a relatively small study area (250 m across) formerly dominated by large stands of branching Acropora corals. The first zone still retains the dominance of branching Acropora corals, although they show a significant decreasing tendency. The second zone exhibits recovery with a significant increase in branching Montipora stellata, which is recruited and grows faster than branching Acropora corals. The third zone is occupied by anemone, Condylactis sp., and demonstrates a stable phase of coral deterioration without recovery. Such differences in coral reef community dynamics within a small spatial scale illustrate mosaic dynamics which have resulted from degradation of the water quality, patchy mortality of large branching Acropora thickets caused by typhoons, the rapid asexual fragmentation and growth of M. stellata making it a successful colonizer, and occupation by anemone, Condylactis sp., together with unstable remnants of dead Acropora rubbles have not allowed coral recruits to survive.     

Estimating the footprint of pollution on coral reefs with models of species turnover

Ecological communities typically change along gradients of human impact, although it is difficult to estimate the footprint of impacts for diffuse threats such as pollution. We developed a joint model (i.e., one that includes multiple species and their interactions with each other and environmental covariates) of benthic habitats on lagoonal coral reefs and used it to infer change in benthic composition along a gradient of distance from logging operations. The model estimated both changes in abundances of benthic groups and their compositional turnover, a type of beta diversity. We used the model to predict the footprint of turbidity impacts from past and recent logging. Benthic communities far from logging were dominated by branching corals, whereas communities close to logging had higher cover of dead coral, massive corals, and soft sediment. Recent impacts were predicted to be small relative to the extensive impacts of past logging because recent logging has occurred far from lagoonal reefs. Our model can be used more generally to estimate the footprint of human impacts on ecosystems and evaluate the benefits of conservation actions for ecosystems.     

Resource use by five sympatric parrotfishes in the San Blas Archipelago,Panama

Resource use by five sympatric species of parrotfish was quantified in the San Blas Archipelago of the Republic of Panama from March to August 1987. Detailed observations of parrotfishes on patch reefs and surrounding seagrass beds showed that they partition resources with respect to habitat, food and size, but not time. Although parrotfishes shared resources, the proportions of each resource used differed significantly among species. Scarus iserti (Bloch) scraped filamentous microalgae that grew from eroded coral pavement on lower slopes of patch reefs and in halos, the area of sparse vegetation surrounding reefs. Sparisoma viride (Bonnaterre) foraged on upper slopes of patch reefs where they mostly took bites from dead coral and associated algae. S. aurofrenatum (Cuvier and Valenciennes) had the broadest diet, which consisted mostly of seagrasses and macro- and microalgae that were attached to dead coral on lower reef slopes and in halos. Although S. chrysopterum (Bloch and Schneider) commonly occurred on patch reefs, it primarily foraged in seagrass beds that surround them. S. rubripinne (Cuvier and Valenciennes) was distributed most widely, ranging from seagrass beds to reef crests, where it took bites from seagrasses, dead coral and macroalgae. Juveniles of all species occurred on lower slopes or in halos where they scraped filamentous microalgae from coral pavement. As they matured, parrotfishes moved into other habitats changing access to different types of food. All of these parrotfishes fed throughout the daytime, and resource use did not differ between morning and afternoon.     

Implications of Urbanization for Artisanal Parrotfish Fisheries in the Western Solomon Islands

Abstract: Increasing migration into urbanized centers in the Solomon Islands poses a great threat to adjacent coral reef fisheries because of negative effects on the fisheries and because it further erodes customary management systems. Parrotfish fisheries are of particular importance because the feeding habits of parrotfish (scrape and excavate coral) are thought to be critical to the resilience of coral reefs and to maintaining coral reef health within marine protected areas. We investigated the ecological impact of localized subsistence and artisanal fishing pressure on parrotfish fisheries in Gizo Town, Western Solomon Islands, by analyzing the density and size distribution of parrotfish with an underwater visual census (UVC), recall diary (i.e., interviews with fishers), and creel surveys to independently assess changes in abundance and catch‐per‐unit‐effort (CPUE) over 2 years. We then compared parrotfish data from Gizo Town with equivalent data from sites open to and closed to fishing in Kida and Nusa Hope villages, which have different customary management regimes. Results indicated a gradient of customary management effectiveness. Parrotfish abundance was greater in customary management areas closed to fishing, especially with regard to larger fish sizes, than in areas open to fishing. The decline in parrotfish abundance from 2004 to 2005 in Gizo was roughly the same magnitude as the difference in abundance decline between inside and outside customary management marine reserves. Our results highlight how weak forms of customary management can result in the rapid decline of vulnerable fisheries around urbanized regions, and we present examples in which working customary management systems (Kinda and Nusa Hope) can positively affect the conservation of parrotfish—and reef fisheries in general—in the highly biodiverse Coral Triangle region.     

Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.     

Spatial and temporal variation in coral recruitment on the Great Barrier Reef: Implications for dispersal hypotheses

Over 15 000 coral recruits were counted on settlement plates from three mid-shelf reefs and six fringing reefs in the northern section of the Great Barrier Reef during two summers (1986 and 1987) and one winter (1987). The density of coral recruits on some settlement plates from a fringing reef was up to 4.88 cm^–2, the highest value ever reported. Mean density of recruits was greater on fringing reefs (81.1 recruits/settlement plate) than on mid-shelf reefs (15.6 recruits/settlement plate), but there was greater spatial variation in abundance of recruits between the fringing reef sites. Other differences between the mid-shelf reefs and the fringing reefs were that different taxa were dominant, and that settlement orientation differed, with mid-shelf recruits settling preferentially on horizontally oriented surfaces and fringingreef recruits preferring vertical surfaces. Of the three midshelf reefs, Green Island reef recorded the highest recruitment rate for each of the two summers, despite having a depauperate adult coral population following predation by the asteroidAcanthaster planci. This suggests that coral larvae frequently travel between reefs. In contrast with an earlier study, there was no consistent difference in abundance of recruits between forereef and backreef locations. Overall, the results indicated great spatial variation in the availability of coral larvae, both on the scale of whole reefs and within-reef habitats.