High macroalgal cover and low coral recruitment undermines the potential resilience of the world's southernmost coral reef assemblages

Coral reefs are under increasing pressure from anthropogenic and climate-induced stressors. The ability of reefs to reassemble and regenerate after disturbances (i.e., resilience) is largely dependent on the capacity of herbivores to prevent macroalgal expansion, and the replenishment of coral populations through larval recruitment. Currently there is a paucity of this information for higher latitude, subtropical reefs. To assess the potential resilience of the benthic reef assemblages of Lord Howe Island (31°32'S, 159°04'E), the worlds' southernmost coral reef, we quantified the benthic composition, densities of juvenile corals (as a proxy for coral recruitment), and herbivorous fish communities. Despite some variation among habitats and sites, benthic communities were dominated by live scleractinian corals (mean cover 37.4%) and fleshy macroalgae (20.9%). Live coral cover was higher than in most other subtropical reefs and directly comparable to lower latitude tropical reefs. Juvenile coral densities (0.8 ind.m(-2)), however, were 5-200 times lower than those reported for tropical reefs. Overall, macroalgal cover was negatively related to the cover of live coral and the density of juvenile corals, but displayed no relationship with herbivorous fish biomass. The biomass of herbivorous fishes was relatively low (204 kg.ha(-1)), and in marked contrast to tropical reefs was dominated by macroalgal browsing species (84.1%) with relatively few grazing species. Despite their extremely low biomass, grazing fishes were positively related to both the density of juvenile corals and the cover of bare substrata, suggesting that they may enhance the recruitment of corals through the provision of suitable settlement sites. Although Lord Howe Islands' reefs are currently coral-dominated, the high macroalgal cover, coupled with limited coral recruitment and low coral growth rates suggest these reefs may be extremely susceptible to future disturbances.     

High resilience masks underlying sensitivity to algal phase shifts of Pacific coral reefs

A single ecosystem can exhibit great biogeographic and environmental variability. While a given ecological driver might have a strong impact in one region, it does not necessarily hold that its importance will extend elsewhere. Coral reefs provide a striking example in that coral communities have low resilience in the Atlantic and remarkable resilience in parts of the species‐rich Pacific. Recent experimental evidence from the Atlantic finds that fishing of large herbivorous fish can be a strong driver of coral resilience. Here, we repeat the Atlantic experiment in the highly resilient forereef of Moorea (French Polynesia), which has repeatedly recovered from disturbances. A combination of cages, fish deterrents (FDs), and controls allowed us to simulate the consequences of fishing large herbivores on algal assemblages, coral recruitment, and the demographic rates of coral juveniles. We find that the impacts of removing large herbivorous reef fish vary with early coral ontogeny. Reduced herbivore access led to a modest macroalgal bloom and reduction in coral recruitment. However, larger juvenile corals (> 1 cm diameter) survived better and grew faster under these conditions because of a reduction in corallivory. To determine the net impact of losing larger herbivorous fish, we combined experimental results with estimated demographic parameters in an individual‐based model. Simulating coral recovery trajectories for five years, we find that protecting larger reef fish led to better recovery in 66–99% of simulations, depending on underlying assumptions (with the more credible assumptions being associated with greater likelihood of net positive impacts). While we find that fishing effects are detrimental to corals in both the Atlantic and Pacific systems studied, the nature of the interactions varied markedly. In the identical previously‐published study in the Atlantic, macroalgae exhibited a rapid bloom and caused a sufficiently large reduction in coral recruitment to force a predicted ecosystem shift to an alternative attractor. The commensurate macroalgal bloom in Moorea was weak yet the corals were two orders of magnitude more sensitive to its presence. We do not suggest that a reduction in recruitment in Moorea will lead to alternative attractors but the long‐term risks of a reduction in recovery rate are cause for concern as rates of coral mortality are projected to increase. The emerging picture is that Pacific reefs are less likely to experience macroalgal blooms but are surprisingly sensitive to such blooms if they occur.     

Global assessment of the status of coral reef herbivorous fishes: evidence for fishing effects

On coral reefs, herbivorous fishes consume benthic primary producers and regulate competition between fleshy algae and reef-building corals. Many of these species are also important fishery targets, yet little is known about their global status. Using a large-scale synthesis of peer-reviewed and unpublished data, we examine variability in abundance and biomass of herbivorous reef fishes and explore evidence for fishing impacts globally and within regions. We show that biomass is more than twice as high in locations not accessible to fisheries relative to fisheries-accessible locations. Although there are large biogeographic differences in total biomass, the effects of fishing are consistent in nearly all regions. We also show that exposure to fishing alters the structure of the herbivore community by disproportionately reducing biomass of large-bodied functional groups (scraper/excavators, browsers, grazer/detritivores), while increasing biomass and abundance of territorial algal-farming damselfishes (Pomacentridae). The browser functional group that consumes macroalgae and can help to prevent coral–macroalgal phase shifts appears to be most susceptible to fishing. This fishing down the herbivore guild probably alters the effectiveness of these fishes in regulating algal abundance on reefs. Finally, data from remote and unfished locations provide important baselines for setting management and conservation targets for this important group of fishes.     

Interactions between herbivorous fish guilds and their influence on algal succession on a coastal coral reef

Herbivory is an important mechanism affecting algal succession, particularly on coral reefs where the relationship between algae and corals is largely controlled by herbivores. However, different functional groups of herbivores may have contrasting effects on succession, which may explain different trajectories of coral reef recovery after disturbance. Here, the effects of different herbivore groups (roving herbivores = foragers and territorial damselfish = farmers) were isolated by a multi-factorial experiment carried out on a coastal coral reef with high macroalgal cover, high farmer densities and relatively low forager abundance. The effects of foragers and farmers were distinguished by monitoring algal succession on settlement tiles placed inside and outside exclusion cages, with orthogonal treatments established inside and outside damselfish territories (with appropriate cage controls). Within 12 months, algal assemblages on ungrazed tiles inside exclusion cages proceeded rapidly from fine filamentous turfs, to corticated algae, to tough erect (e.g. Amphiroa spp.) and foliose (e.g. Peyssonnellidae) calcified algae. Farmers had a dramatic impact on succession, essentially arresting the development of the algal community at a point where it was dominated by palatable filamentous algae of the genus Polysiphonia. Fleshy macroalgae such as Sargassum spp. were excluded from farmer territories. In contrast, foragers did not suppress fleshy macroalgae, but rather, appeared to decelerate succession and promote a relatively diverse assemblage. In contrast to forager-dominated reefs, farmer territories did not appear to function solely as forager exclusion areas or promote algal diversity as a result of intermediate grazing pressure. The relatively strong effects of farmers observed here may represent a future scenario for coral reefs that are increasingly subject to overfishing of large grazing fishes.     

Effects of herbivory,nutrients, and reef protection on algal proliferation and coral growth on a tropical reef

Maintaining coral reef resilience against increasing anthropogenic disturbance is critical for effective reef management. Resilience is partially determined by how processes, such as herbivory and nutrient supply, affect coral recovery versus macroalgal proliferation following disturbances. However, the relative effects of herbivory versus nutrient enrichment on algal proliferation remain debated. Here, we manipulated herbivory and nutrients on a coral-dominated reef protected from fishing, and on an adjacent macroalgal-dominated reef subject to fishing and riverine discharge, over 152 days. On both reefs, herbivore exclusion increased total and upright macroalgal cover by 9-46 times, upright macroalgal biomass by 23-84 times, and cyanobacteria cover by 0-27 times, but decreased cover of encrusting coralline algae by 46-100% and short turf algae by 14-39%. In contrast, nutrient enrichment had no effect on algal proliferation, but suppressed cover of total macroalgae (by 33-42%) and cyanobacteria (by 71% on the protected reef) when herbivores were excluded. Herbivore exclusion, but not nutrient enrichment, also increased sediment accumulation, suggesting a strong link between herbivory, macroalgal growth, and sediment retention. Growth rates of the corals Porites cylindrica and Acropora millepora were 30-35% greater on the protected versus fished reef, but nutrient and herbivore manipulations within a site did not affect coral growth. Cumulatively, these data suggest that herbivory rather than eutrophication plays the dominant role in mediating macroalgal proliferation, that macroalgae trap sediments that may further suppress herbivory and enhance macroalgal dominance, and that corals are relatively resistant to damage from some macroalgae but are significantly impacted by ambient reef condition.     

Herbivory versus corallivory: are parrotfish good or bad for Caribbean coral reefs?

With coral cover in decline on many Caribbean reefs, any process of coral mortality is of potential concern. While sparisomid parrotfishes are major grazers of Caribbean reefs and help control algal blooms, the fact that they also undertake corallivory has prompted some to question the rationale for their conservation. Here the weight of evidence for beneficial effects of parrotfishes, in terms of reducing algal cover and facilitating demographic processes in corals, and the deleterious effects of parrotfishes in terms of causing coral mortality and chronic stress, are reviewed. While elevated parrotfish density will likely increase the predation rate upon juvenile corals, the net effect appears to be positive in enhancing coral recruitment through removal of macroalgal competitors. Parrotfish corallivory can cause modest partial colony mortality in the most intensively grazed species of Montastraea but the generation and healing of bite scars appear to be in near equilibrium, even when coral cover is low. Whole colony mortality in adult corals can lead to complete exclusion of some delicate, lagoonal species of Porites from forereef environments but is only reported for one reef species (Porites astreoides), for one habitat (backreef), and with uncertain incidence (though likely <<10%). No deleterious effects of predation on coral growth or fecundity have been reported, though recovery of zooxanthellae after bleaching events may be retarded. The balance of evidence to date finds strong support for the herbivory role of parrotfishes in facilitating coral recruitment, growth, and fecundity. In contrast, no net deleterious effects of corallivory have been reported for reef corals. Corallivory is unlikely to constrain overall coral cover but contraints upon dwindling populations of the Montastraea annularis species complex are feasible and the role of parrotfishes as a vector of coral disease requires evaluation. However, any assertion that conservation practices should guard against protecting corallivorous parrotfishes appears to be unwarranted at this stage.     

Monitoring Herbivorous Fishes as Indicators of Coral Reef Resilience in American Samoa

Resilience-based management aims to promote or protect processes and species that underpin an ecosystem''s capacity to withstand and recover from disturbance. The management of ecological processes is a developing field that requires reliable indicators that can be monitored over time. Herbivory is a key ecological process on coral reefs, and pooling herbivorous fishes into functional groups based on their feeding mode is increasingly used as it may quantify herbivory in ways that indicate resilience. Here we evaluate whether the biomass estimates of these herbivore functional groups are good predictors of reef benthic assemblages, using data from 240 sites from five island groups in American Samoa. Using an information theoretic approach, we assembled a candidate set of linear and nonlinear models to identify the relations between benthic cover and total herbivore and non-herbivore biomass and the biomass of the aforementioned functional groups. For each benthic substrate type considered (encrusting algae, fleshy macroalgae, hard coral and turf algae), the biomass of herbivorous fishes were important explanatory variables in predicting benthic cover, whereas biomass of all fishes combined generally was not. Also, in all four cases, variation in cover was best explained by the biomass of specific functional groups rather than by all herbivores combined. Specifically: 1) macroalgal and turf algal cover decreased with increasing biomass of ‘grazers/detritivores’; and 2) cover of encrusting algae increased with increasing biomass of ‘grazers/detritivores’ and browsers. Furthermore, hard coral cover increased with the biomass of large excavators/bio-eroders (made up of large-bodied parrotfishes). Collectively, these findings emphasize the link between herbivorous fishes and the benthic community and demonstrate support for the use of functional groups of herbivores as indicators for resilience-based monitoring.     

Large-scale associations between macroalgal cover and grazer biomass on mid-depth reefs in the Caribbean

Since the 1970s, macroalgae have become considerably more abundant on many Caribbean reefs and overfishing of grazing fishes has been implicated as a contributory factor. We explored relationships between algal cover and grazers (biomass of herbivorous fishes and abundance of the sea-urchin Diadema antillarum) on mid-depth reefs (12-15 m) in 19 areas at seven locations in Jamaica, Barbados, Belize, Grand Cayman and Cuba, between April 1997 and April 1998. Diadema antillarum density was never >0.01 m^-2, while herbivorous fish biomass (acanthurids and scarids ⁸ cm total length) varied from 2-5 g m^-2 in Jamaica to 17.1 g m^-2 in Barbados, and was strongly correlated, negatively with macroalgal cover and positively with 'cropped' substratum (sum of 'bare', turf and crustose-coralline substrata) cover. However, overfishing of herbivorous fishes alone cannot explain the widespread abundance of macroalgae, as even on lightly fished reefs, macroalgal cover was mostly >20%. Herbivorous fish populations on those reefs were apparently only able to maintain approximately 40-60% of reef substratum in cropped states, but due to low space-occupation by coral and other invertebrates, 70-90% of substratum was available to algae. The abundance of macroalgae on lightly fished reefs may therefore be a symptom of low coral cover in combination with the continuing absence of Diadema antillarum.     

Spatial patterns of parrotfish corallivory in the Caribbean: the importance of coral taxa, density and size

The past few decades have seen an increase in the frequency and intensity of disturbance on coral reefs, resulting in shifts in size and composition of coral populations. These changes have lead to a renewed focus on processes that influence demographic rates in corals, such as corallivory. While previous research indicates selective corallivory among coral taxa, the importance of coral size and the density of coral colonies in influencing corallivory are unknown. We surveyed the size, taxonomy and number of bites by parrotfish per colony of corals and the abundance of three main corallivorous parrotfish (Sparisoma viride, Sparisoma aurofrenatum, Scarus vetula) at multiple spatial scales (reefs within islands: 1-100 km, and between islands: >100 km) within the Bahamas Archipelago. We used a linear mixed model to determine the influence of coral taxa, colony size, colony density, and parrotfish abundance on the intensity of corallivory (bites per m(2) of coral tissue). While the effect of colony density was significant in determining the intensity of corallivory, we found no significant influence of colony size or parrotfish abundance (density, biomass or community structure). Parrotfish bites were most frequently observed on the dominant species of reef building corals (Montastraea annularis, Montastraea faveolata and Porites astreoides), yet our results indicate that when the confounding effects of colony density and size were removed, selective corallivory existed only for the less dominant Porites porites. As changes in disturbance regimes result in the decline of dominant frame-work building corals such as Montastraea spp., the projected success of P. porites on Caribbean reefs through high reproductive output, resistance to disease and rapid growth rates may be attenuated through selective corallivory by parrotfish.     

Suppression of herbivory by macroalgal density: a critical feedback on coral reefs?

Coral reefs globally are in decline, with some reefs undergoing phase shifts from coral-dominance to degraded states dominated by large fleshy macroalgae. These shifts have been underpinned by the overharvesting of herbivorous fishes and represent a fundamental change in the physical structure of these reefs. Although the physical structure provided by corals is regarded as a key feature that facilitates herbivore activity, the influence of the physical structure of macroalgal stands is largely unknown. Using transplanted Sargassum, the largest coral reef macroalga, we created habitat patches of predetermined macroalgal density (0.25-6.23 kg m(-2)). Remote video cameras revealed both grazing and browsing fishes avoided high density patches, preferring relatively open areas with low macroalgal cover. This behaviour may provide a positive feedback leading to the growth and persistence of macroalgal stands; increasing the stability of phase shifts to macroalgae.     

Global disparity in the resilience of coral reefs

The great sensitivity of coral reefs to climate change has raised concern over their resilience. An emerging body of resilience theory stems largely from research carried out in a single biogeographic region; the Caribbean. Such geographic bias raises the question of transferability of concepts among regions. In this article, we identify factors that might predispose the Caribbean to its low resilience, including faster rates of macroalgal growth, higher rates of algal recruitment, basin-wide iron-enrichment of algal growth from aeolian dust, a lack of acroporid corals, lower herbivore biomass and missing groups of herbivores. Although mechanisms of resilience are likely to be ubiquitous, our analysis suggests that Indo-Pacific reefs would have to be heavily degraded to exhibit bistability or undergo coral-macroalgal phase shifts.     

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.

     

Marine Reserves Enhance the Recovery of Corals on Caribbean Reefs

The fisheries and biodiversity benefits of marine reserves are widely recognised but there is mounting interest in exploiting the importance of herbivorous fishes as a tool to help ecosystems recover from climate change impacts. This approach might be particularly suitable for coral reefs, which are acutely threatened by climate change, yet the trophic cascades generated by reserves are strong enough that they might theoretically enhance the rate of coral recovery after disturbance. However, evidence for reserves facilitating coral recovery has been lacking. Here we investigate whether reductions in macroalgal cover, caused by recovery of herbivorous parrotfishes within a reserve, have resulted in a faster rate of coral recovery than in areas subject to fishing. Surveys of ten sites inside and outside a Bahamian marine reserve over a 2.5-year period demonstrated that increases in coral cover, including adjustments for the initial size-distribution of corals, were significantly higher at reserve sites than those in non-reserve sites. Furthermore, macroalgal cover was significantly negatively correlated with the change in total coral cover over time. Recovery rates of individual species were generally consistent with small-scale manipulations on coral-macroalgal interactions, but also revealed differences that demonstrate the difficulties of translating experiments across spatial scales. Size-frequency data indicated that species which were particularly affected by high abundances of macroalgae outside the reserve had a population bottleneck restricting the supply of smaller corals to larger size classes. Importantly, because coral cover increased from a heavily degraded state, and recovery from such states has not previously been described, similar or better outcomes should be expected for many reefs in the region. Reducing herbivore exploitation as part of an ecosystem-based management strategy for coral reefs appears to be justified.     

Context-dependent corallivory by parrotfishes in a Caribbean reef ecosystem

This study assesses the patterns of corallivory by parrotfishes across reefs of the Florida Keys, USA. These reefs represent a relatively unique combination within the wider Caribbean of low coral cover and high parrotfish abundance suggesting that predation pressure could be intense. Surveys across eight shallow forereefs documented the abundance of corals, corallivorous parrotfishes, and predation scars on corals. The corals Porites porites and Porites astreoides were preyed on most frequently with the rates of predation an order of magnitude greater than has been documented for other areas of the Caribbean. In fact, parrotfish bite density on these preferred corals was up to 34 times greater than reported for corals on other reefs worldwide. On reefs where coral cover was low and corals such as Montastraea faveolata, often preferred prey for parrotfishes, were rare, predation rates on P. porites and P. astreoides, and other less common corals, intensified further. The intensity of parrotfish predation increased significantly as coral cover decreased. However, parrotfish abundance showed only a marginal positive relationship with predation pressure on corals, likely because corallivorous parrotfish were abundant across all reefs. Parrotfishes often have significant positive impacts on coral cover by facilitating coral recruitment, survival, and growth via their grazing of algae. However, abundant corallivorous parrotfishes combined with low coral cover may result in higher predation on corals and intensify the negative impact that parrotfishes have on remaining corals.     

Phase shifts and the role of herbivory in the resilience of coral reefs

Cousin Island marine reserve (Seychelles) has been an effectively protected no-take marine protected area (MPA) since 1968 and was shown in 1994 to support a healthy herbivorous fish assemblage. In 1998 Cousin Island reefs suffered extensive coral mortality following a coral bleaching event, and a phase shift from coral to algal dominance ensued. By 2005 mean coral cover was <1%, structural complexity had fallen and there had been a substantial increase in macroalgal cover, up to 40% in some areas. No clear trends were apparent in the overall numerical abundance and biomass of herbivorous fishes between 1994 and 2005, although smaller individuals became relatively scarce, most likely due to the loss of reef structure. Analysis of the feeding habits of six abundant and representative herbivorous fish species around Cousin Island in 2006 demonstrated that epilithic algae were the preferred food resource of all species and that macroalgae were avoided. Given the current dominance of macroalgae and the apparent absence of macroalgal consumers, it is suggested that the increasing abundance of macroalgae is reducing the probability of the system reverting to a coral dominated state.     

Does Herbivorous Fish Protection Really Improve Coral Reef Resilience? A Case Study from New Caledonia (South Pacific)

Parts of coral reefs from New Caledonia (South Pacific) were registered at the UNESCO World Heritage list in 2008. Management strategies aiming at preserving the exceptional ecological value of these reefs in the context of climate change are currently being considered. This study evaluates the appropriateness of an exclusive fishing ban of herbivorous fish as a strategy to enhance coral reef resilience to hurricanes and bleaching in the UNESCO-registered areas of New Caledonia. A two-phase approach was developed: 1) coral, macroalgal, and herbivorous fish communities were examined in four biotopes from 14 reefs submitted to different fishing pressures in New Caledonia, and 2) results from these analyses were challenged in the context of a global synthesis of the relationship between herbivorous fish protection, coral recovery and relative macroalgal development after hurricanes and bleaching. Analyses of New Caledonia data indicated that 1) current fishing pressure only slightly affected herbivorous fish communities in the country, and 2) coral and macroalgal covers remained unrelated, and macroalgal cover was not related to the biomass, density or diversity of macroalgae feeders, whatever the biotope or level of fishing pressure considered. At a global scale, we found no relationship between reef protection status, coral recovery and relative macroalgal development after major climatic events. These results suggest that an exclusive protection of herbivorous fish in New Caledonia is unlikely to improve coral reef resilience to large-scale climatic disturbances, especially in the lightly fished UNESCO-registered areas. More efforts towards the survey and regulation of major chronic stress factors such as mining are rather recommended. In the most heavily fished areas of the country, carnivorous fish and large targeted herbivores may however be monitored as part of a precautionary approach.     

Spatial variation in the effects of grazing on epilithic algal turfs on the Great Barrier Reef, Australia

Of all benthic components on tropical reefs, algal turfs are the most widespread and the main source of primary productivity. We compared the importance of grazing by herbivores on algal turfs on two zones with marked differences in terms of benthic composition, herbivore biomass and grazing pressure, the inner flat and crest, of an inshore reef on the Great Barrier Reef, Australia. A combination of herbivore exclusion cages and transplants of coral rubble covered by algal turfs between reef zones was used to examine changes in algal turfs over a 4-day experimental period. In situ crest turfs had lower algal height, sediment loads and particulate content than reef flat turfs. Caged samples on the crest exhibited an increase in all three variables. In contrast, in situ and caged treatments on the flat presented algal turfs with similar values for the three analysed variables, with high algal height and heavy particulate and sediment loads. In the absence of cages, reef flat turfs transplanted to the crest had decreased algal height, total particulate material and particulate inorganic content, while the opposite was found in crest turf samples transplanted to the flat. Our results highlight the dynamic nature of algal turfs and the clear differences in the relative importance of herbivory in shaping turf length and sediment load between the reef crest and inner flat.     

Overgrowth and killing of corals by the brown alga Lobophora hederacea (Dictyotales,Phaeophyceae) on healthy reefs in New Caledonia: A new case of the epizoism syndrome

Coral reef degradation is often associated with regime shifts from coral‐ to macroalgal‐dominated reefs. These shifts demonstrate that under certain conditions (e.g. coral mortality, decrease in herbivory, increased nutrients supply) some macroalgae may overgrow corals. The outcome of the competition is dependent on algal aggressiveness and the coral susceptibility. In undisturbed reefs, herbivore grazing is regulating macroalgal cover, thus preventing the latter from overgrowing corals. However, some macroalgae have evolved strategies not only to outcompete corals but also to escape herbivory to some extent, allowing overgrowth of some coral species in undisturbed reefs. Epizoism represents one of those successful strategies, and has been previously documented with red algae, cyanobacteria and Lobophora variegata (Dictyotales, Phaeophyceae). Here we report a new case of epizoism leading to coral mortality, involving a recently described species of Lobophora, L. hederacea, overgrowing the coral Seriatopora caliendrum (Pocilloporidae) in undisturbed reefs in New Caledonia.     

The influence of coral reef benthic condition on associated fish assemblages

Accumulative disturbances can erode a coral reef's resilience, often leading to replacement of scleractinian corals by macroalgae or other non-coral organisms. These degraded reef systems have been mostly described based on changes in the composition of the reef benthos, and there is little understanding of how such changes are influenced by, and in turn influence, other components of the reef ecosystem. This study investigated the spatial variation in benthic communities on fringing reefs around the inner Seychelles islands. Specifically, relationships between benthic composition and the underlying substrata, as well as the associated fish assemblages were assessed. High variability in benthic composition was found among reefs, with a gradient from high coral cover (up to 58%) and high structural complexity to high macroalgae cover (up to 95%) and low structural complexity at the extremes. This gradient was associated with declining species richness of fishes, reduced diversity of fish functional groups, and lower abundance of corallivorous fishes. There were no reciprocal increases in herbivorous fish abundances, and relationships with other fish functional groups and total fish abundance were weak. Reefs grouping at the extremes of complex coral habitats or low-complexity macroalgal habitats displayed markedly different fish communities, with only two species of benthic invertebrate feeding fishes in greater abundance in the macroalgal habitat. These results have negative implications for the continuation of many coral reef ecosystem processes and services if more reefs shift to extreme degraded conditions dominated by macroalgae.     

Competition between macroalgae and corals: effects of herbivore exclusion and increased algal biomass on coral survivorship and growth

Recent declines in coral abundance accompanied by increases in macroalgal cover on Florida reefs highlight the importance of competition for space between these groups. This paper documents the frequency of coral-algal interactions on the Northern Florida Reef Tract and evaluates the effects of grazer exclusions and experimental algal addition on growth and tissue mortality of three coral species, Siderastrea siderea, Porites astreoides, and Montastraea faveolata. The frequency of interactions between corals and macroalgae was high as more than 50% of the basal perimeter of colonies was in contact with macroalgae; turf forms, Halimeda spp., and Dictyota spp. were the most common groups in contact with corals. Decreased grazing pressure resulted in significant increases in algal biomass within cages, and caged corals showed species-specific susceptibility to increased algal biomass. While no effects were detected for S. siderea, significant decreases in growth rates were documented for caged P. astreoides which had growth rates three to four times lower than uncaged colonies. When an algal addition treatment was included to duplicate maximum algal biomass levels documented for reefs in the area, colonies of P. astreoides in the algal addition treatment had growth rates up to ten times lower than uncaged colonies. High susceptibility to algal overgrowth was also found for the reef-building coral M. faveolata, which experienced significant tissue mortality under both uncaged (5.2% decrease in live tissue area per month) and caged (10.2% per month) conditions. The documented effects of increased algal biomass on coral growth and tissue mortality suggest a potential threat for the long-term survivorship and growth of corals in the Florida Reef Tract if present rates of algal growth and space utilization are maintained.