Phase shifts and the stability of macroalgal communities on Caribbean coral reefs

Caribbean coral reefs are widely thought to exhibit two alternate stable states with one being dominated by coral and the other by macroalgae. However, the observation of linear empirical relationships among grazing, algal cover and coral recruitment has led the existence of alternate stable states to be questioned; are reefs simply exhibiting a continuous phase shift in response to grazing or are the alternate states robust to certain changes in grazing? Here, a model of a Caribbean forereef is used to reconcile the existence of two stable community states with common empirical observations. Coral-depauperate and coral-dominated reef states are predicted to be stable on equilibrial time scales of decades to centuries and their emergence depends on the presence or absence of a bottleneck in coral recruitment, which is determined by threshold levels of grazing intensity and other process variables. Under certain physical and biological conditions, corals can be persistently depleted even while increases in grazing reduce macroalgal cover and enhance coral recruitment; only once levels of recruitment becomes sufficient to overwhelm the population bottleneck will the coral-dominated state begin to emerge. Therefore, modest increases in grazing will not necessarily allow coral populations to recover, whereas large increases, such as those associated with recovery of the urchin Diadema antillarum, are likely to exceed threshold levels of grazing intensity and set a trajectory of coral recovery. The postulated existence of alternate stable states is consistent with field observations of linear relationships between grazing, algal cover and coral recruitment when coral cover is low and algal exclusion when coral cover is high. The term ‘macroalgal dominated’ is potentially misleading because the coral-depauperate state can be associated with various levels of macroalgal cover. The term ‘coral depauperate’ is preferable to ‘macroalgal dominated’ when describing alternate states of Caribbean reefs.     

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.     

Spatial dynamics of benthic competition on coral reefs

The community structure of sedentary organisms is largely controlled by the outcome of direct competition for space. Understanding factors defining competitive outcomes among neighbors is thus critical for predicting large-scale changes, such as transitions to alternate states within coral reefs. Using a spatially explicit model, we explored the importance of variation in two spatial properties in benthic dynamics on coral reefs: (1) patterns of herbivory are spatially distinct between fishes and sea urchins and (2) there is wide variation in the areal extent into which different coral species can expand. We reveal that the size-specific, competitive asymmetry of corals versus fleshy algae highlights the significance of spatial patterning of herbivory and of coral growth. Spatial dynamics that alter the demographic importance of coral recruitment and maturation have profound effects on the emergent structure of the reef benthic community. Spatially constrained herbivory (as by sea urchins) is more effective than spatially unconstrained herbivory (as by many fish) at opening space for the time needed for corals to settle and to recruit to the adult population. Further, spatially unconstrained coral growth (as by many branching coral species) reduces the number of recruitment events needed to fill a habitat with coral relative to more spatially constrained growth (as by many massive species). Our model predicts that widespread mortality of branching corals (e.g., Acropora spp) and herbivorous sea urchins (particularly Diadema antillarum) in the Caribbean has greatly reduced the potential for restoration across the region.     

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.     

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.     

Habitat complexity and consumer-mediated positive feedbacks on a Caribbean coral reef

Theoretical and empirical evidence suggest that positive feedbacks can increase resilience in ecological communities. On Caribbean coral reefs, there have been striking shifts from physically complex communities with high coral cover to relatively homogenous communities dominated by macroalgae, which have persisted for decades. However, little is known about positive feedbacks that may maintain coral reef community states. Here, I explore a potential consumer-mediated feedback on a Jamaican reef by examining how grazing by a keystone herbivore ( Diadema antillarum ) is enhanced by physical structure, which offer refugia from predation. Surveys revealed that habitat complexity and Diadema density were positively related. Increasing habitat complexity by adding physical structure significantly decreased macroalgal cover and increased the proportion of urchins in algal habitats in field manipulations. Experimental increases in urchin density also decreased macroalgal cover, but did not affect the proportion of urchins in algal habitats. These results suggest that the low habitat complexity of macroalgal-dominated reefs may inhibit an urchin-mediated shift to coral dominance and that positive feedbacks must be considered in reef restoration efforts.     

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.     

Influence of Diadema antillarum populations (Echinodermata: Diadematidae) on algal community structure in Jardines de la Reina, Cuba

The 1983-1984 mass mortality of Diadema antillarum produced severe damages on Caribbean reefs contributing to substantial changes in community structure that still persist. Despite the importance of Diadema grazing in structuring coral reefs, available information on current abundances and algal-urchin interactions in Cuba is scarce. We analyzed spatial variations in Diadema abundance and its influence on algal community structure in 22 reef sites in Jardines de la Reina, in June/2004 and April/2005. Urchins were counted in five 30 x 2m transects per site, and algal coverage was estimated in randomly located 0.25m side quadrats (15 per site). Abundances of Diadema were higher at reef crests (0.013-1.553 ind/m2), while reef slope populations showed values up to three orders of magnitude lower and were overgrown by macroalgae (up to 87%, local values). Algal community structure at reef slopes were dominated by macroalgae, especially Dictyota, Lobophora and Halimeda while the most abundant macroalgae at reef crests were Halimeda and Amphiroa. Urchin densities were negatively and positively correlated with mean coverage of macroalgae and crustose coralline algae, respectively, when analyzing data pooled across all sites, but not with data from separate habitats (specially reef crest), suggesting, along with historical fish biomass, that shallow reef community structure is being shaped by the synergistic action of other factors (e.g. fish grazing) rather than the influence of Diadema alone. However, we observed clear signs of Diadema grazing at reef crests and decreased macroalgal cover according to 2001 data, what suggest that grazing intensity at this habitat increased at the same time that Diadema recruitment began to be noticeable. Furthermore, the excessive abundance of macroalgae at reef slopes and the scarcity of crustose coralline algae seems to be due by the almost complete absence of D. antillarum at mid depth reefs, where local densities of this urchin were predominantly low.     

Allelochemicals produced by Caribbean macroalgae and cyanobacteria have species-specific effects on reef coral microorganisms

Coral populations have precipitously declined on Caribbean reefs while algal abundance has increased, leading to enhanced competitive damage to corals, which likely is mediated by the potent allelochemicals produced by both macroalgae and benthic cyanobacteria. Allelochemicals may affect the composition and abundance of coral-associated microorganisms that control host responses and adaptations to environmental change, including susceptibility to bacterial diseases. Here, we demonstrate that extracts of six Caribbean macroalgae and two benthic cyanobacteria have both inhibitory and stimulatory effects on bacterial taxa cultured from the surfaces of Caribbean corals, macroalgae, and corals exposed to macroalgal extracts. The growth of 54 bacterial isolates was monitored in the presence of lipophilic and hydrophilic crude extracts derived from Caribbean macroalgae and cyanobacteria using 96-well plate bioassays. All 54 bacterial cultures were identified by ribotyping. Lipophilic extracts from two species of Dictyota brown algae inhibited >50% of the reef coral bacteria assayed, and hydrophilic compounds from Dictyota menstrualis particularly inhibited Vibrio bacteria, a genus associated with several coral diseases. In contrast, both lipo- and hydrophilic extracts from 2 species of Lyngbya cyanobacteria strongly stimulated bacterial growth. The brown alga Lobophora variegata produced hydrophilic compounds with broad-spectrum antibacterial effects, which inhibited 93% of the bacterial cultures. Furthermore, bacteria cultured from different locations (corals vs. macroalgae vs. coral surfaces exposed to macroalgal extracts) responded differently to algal extracts. These results reveal that extracts from macroalgae and cyanobacteria have species-specific effects on the composition of coral-microbial assemblages, which in turn may increase coral host susceptibility to disease and result in coral mortality.     

Empirical relationships among resilience indicators on Micronesian reefs

A process-orientated understanding of ecosystems usually starts with an exploratory analysis of empirical relationships among potential drivers and state variables. While relationships among herbivory, algal cover, and coral recruitment, have been explored in the Caribbean, the nature of such relationships in the Pacific appears to be variable or unclear. Here, we examine potential drivers structuring the benthos and herbivorous fish assemblages of outer-shelf reefs in Micronesia (Palau, Guam and Pohnpei). Surveys were stratified by wave exposure and protection from fishing. High biomass of most herbivores was favoured by high wave exposure. High abundance of large-bodied scarids was associated with low turf abundance, high coral cover, and marine reserves. The remaining herbivores were more abundant in reefs with low coral cover, possibly because space and hence food limitation occur in high-coral-cover reefs. Rugosity had no detectable effect on herbivorous fish abundance once differences in exposure and coral cover were accounted for. At identical depths, high wave exposure was associated with greater volumes (cover × canopy height) of macroalgae and algal turfs, which most likely resulted from high primary productivity driven by flow. In exposed areas, macroalgal cover declined as the acanthurid biomass increased. The volume of algal turfs was negatively associated with coral cover and herbivore biomass. In turn, high coral cover and herbivore biomass are likely to intensify grazing. The density of juvenile corals was variable where macroalgal cover was low but was confined to lower densities where macroalgal cover was high. High coral cover and density of juvenile corals were favoured in sheltered habitats. While a weak positive relationship was found between scarid biomass and juvenile coral density, we hypothesise that high scarid densities may hinder juvenile density through increased corallivory. New hypotheses emerged that will help clarify the role of acanthurids, wave exposure, and corallivory in driving the recovery of Pacific coral communities.     

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.     

Three-dimensional species distribution modelling reveals the realized spatial niche for coral recruitment on contemporary Caribbean reefs

The three-dimensional structure of habitats is a critical component of species' niches driving coexistence in species-rich ecosystems. However, its influence on structuring and partitioning recruitment niches has not been widely addressed. We developed a new method to combine species distribution modelling and structure from motion, and characterized three-dimensional recruitment niches of two ecosystem engineers on Caribbean coral reefs, scleractinian corals and gorgonians. Fine-scale roughness was the most important predictor of suitable habitat for both taxa, and their niches largely overlapped, primarily due to scleractinians' broader niche breadth. Crevices and holes at mm scales on calcareous rock with low coral cover were more suitable for octocorals than for scleractinian recruits, suggesting that the decline in scleractinian corals is facilitating the recruitment of octocorals on contemporary Caribbean reefs. However, the relative abundances of the taxa were independent of the amount of suitable habitat on the reef, emphasizing that niche processes alone do not predict recruitment rates.     

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

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

White-band disease and the changing face of Caribbean coral reefs

In recent decades, the cover of fleshy macroalgae has increased and coral cover has decreased on most Caribbean reefs. Coral mortality precipitated this transition, and the accumulation of macroalgal biomass has been enhanced by decreased herbivory and increased nutrient input. Populations of Acropora palmata (elkhorn coral) and A. cervicornis (staghorn coral), two of the most important framework-building species, have died throughout the Caribbean, substantially reducing coral cover and providing substratum for algal growth. Hurricanes have devastated local populations of Acropora spp. over the past 20–25 years, but white-band disease, a putative bacterial syndrome specific to the genus Acropora, has been a more significant source of mortality over large areas of the Caribbean region.Paleontological data suggest that the regional Acropora kill is without precedent in the late Holocene. In Belize, A. cervicornis was the primary ecological and geological constituent of reefs in the central shelf lagoon until the mid-1980s. After constructing reef framework for thousands of years, A. cervicornis was virtually eliminated from the area over a ten-year period. Evidence from other parts of the Caribbean supports the hypothesis of continuous Holocene accumulation and recent mass mortality of Acropora spp. Prospects are poor for the rapid recovery of A. cervicornis, because its reproductive strategy emphasizes asexual fragmentation at the expense of dispersive sexual reproduction. A. palmata also relies on fragmentation, but this species has a higher rate of sexual recruitment than A. cervicornis. If the Acropora spp. do not recover, macroalgae will continue to dominate Caribbean reefs, accompanied by increased abundances of brooding corals, particularly Agaricia spp. and Porites spp. The outbreak of white-band disease has been coincident with increased human activity, and the possibility of a causal connection should be further investigated.     

Preservation of in situ reef framework in regions of low hurricane frequency: Pleistocene of Curaçao and Bonaire, southern Caribbean

Luxuriant fringing reefs along the southwestern shores of the Caribbean islands of Curaçao and Bonaire (12°N), located outside the most frequent hurricane tracks, are rarely affected by major storms. Consequently, reef growth and long-term preservation are potentially optimal and distinct from reefs experiencing greater hurricane frequency. Hurricane Lenny (November 1999) took an unusual west-to-east track, bisecting the Caribbean Basin north of these islands, but generated heavy waves (3-6  m) that severely damaged reefs along the normally leeward shores. Massive coral colonies >100 years old were toppled, but even at the most severely damaged sites, 82–85% of colonies remained in growth position. Late Pleistocene (125  ka) elevated reefs in the Lower Terrace of Curaçao record even higher proportions of corals in growth position (93%), possibly reflecting a low hurricane frequency during the Pleistocene highstand. In comparison, coeval Pleistocene reefs in regions that today experience a high hurricane frequency (Great Inagua Island and San Salvador, Bahamas) have lower proportions of corals preserved in growth position (79% and 38%, respectively). These results are consistent with the hypothesis that reefs in regions experiencing very low hurricane frequency, like the southern Caribbean, are more likely to be preserved with corals in primary growth position in comparison to regions with higher hurricane frequency.     

Flattening of Caribbean coral reefs: region-wide declines in architectural complexity

Coral reefs are rich in biodiversity, in large part because their highly complex architecture provides shelter and resources for a wide range of organisms. Recent rapid declines in hard coral cover have occurred across the Caribbean region, but the concomitant consequences for reef architecture have not been quantified on a large scale to date. We provide, to our knowledge, the first region-wide analysis of changes in reef architectural complexity, using nearly 500 surveys across 200 reefs, between 1969 and 2008. The architectural complexity of Caribbean reefs has declined nonlinearly with the near disappearance of the most complex reefs over the last 40 years. The flattening of Caribbean reefs was apparent by the early 1980s, followed by a period of stasis between 1985 and 1998 and then a resumption of the decline in complexity to the present. Rates of loss are similar on shallow (<6 m), mid-water (6–20 m) and deep (>20 m) reefs and are consistent across all five subregions. The temporal pattern of declining architecture coincides with key events in recent Caribbean ecological history: the loss of structurally complex Acropora corals, the mass mortality of the grazing urchin Diadema antillarum and the 1998 El Nino Southern Oscillation-induced worldwide coral bleaching event. The consistently low estimates of current architectural complexity suggest regional-scale degradation and homogenization of reef structure. The widespread loss of architectural complexity is likely to have serious consequences for reef biodiversity, ecosystem functioning and associated environmental services.     

Competitive interactions between corals and Trididemnum solidum on Mexican Caribbean reefs

The ascidian Trididemnum solidum competes for space on Caribbean reefs and is capable of overgrowing live scleractinian corals. From 2006 to 2009, we monitored over 30,000 coral colonies and quantified competitive interactions with this ascidian at four reef sites along the Mexican Caribbean. The total number of competitive interactions increased in time, but the mean percentage of coral colonies involved in interactions remained lower than 1% in all reefs. Bottom cover by T. solidum was also low (mean < 0.5%) in all reef sites in all sampling years. We conclude that during the temporal scope of our study, the overall potential effect of T. solidum on the dynamics of Mexican Caribbean coral populations was minimal.     

Chemical and Physical Environmental Conditions Underneath Mat- and Canopy-Forming Macroalgae,and Their Effects on Understorey Corals

Disturbed coral reefs are often dominated by dense mat- or canopy-forming assemblages of macroalgae. This study investigated how such dense macroalgal assemblages change the chemical and physical microenvironment for understorey corals, and how the altered environmental conditions affect the physiological performance of corals. Field measurements were conducted on macroalgal-dominated inshore reefs in the Great Barrier Reef in quadrats with macroalgal biomass ranging from 235 to 1029 g DW m⁻² dry weight. Underneath mat-forming assemblages, the mean concentration of dissolved oxygen was reduced by 26% and irradiance by 96% compared with conditions above the mat, while concentrations of dissolved organic carbon and soluble reactive phosphorous increased by 26% and 267%, respectively. The difference was significant but less pronounced under canopy-forming assemblages. Dissolved oxygen declined and dissolved inorganic carbon and alkalinity increased with increasing algal biomass underneath mat-forming but not under canopy-forming assemblages. The responses of corals to conditions similar to those found underneath algal assemblages were investigated in an aquarium experiment. Coral nubbins of the species Acropora millepora showed reduced photosynthetic yields and increased RNA/DNA ratios when exposed to conditions simulating those underneath assemblages (pre-incubating seawater with macroalgae, and shading). The magnitude of these stress responses increased with increasing proportion of pre-incubated algal water. Our study shows that mat-forming and, to a lesser extent, canopy-forming macroalgal assemblages alter the physical and chemical microenvironment sufficiently to directly and detrimentally affect the metabolism of corals, potentially impeding reef recovery from algal to coral-dominated states after disturbance. Macroalgal dominance on coral reefs therefore simultaneously represents a consequence and cause of coral reef degradation.     

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.     

Algal contact as a trigger for coral disease

Diseases are causing alarming declines in reef‐building coral species, the foundation blocks of coral reefs. The emergence of these diseases has occurred simultaneously with large increases in the abundance of benthic macroalgae. Here, we show that physical contact with the macroalga Halimeda opuntia can trigger a virulent disease known as white plague type II that has caused widespread mortality in most Caribbean coral species. Colonies of the dominant coral Montastraea faveolata exposed to algal transplants developed the disease whereas unexposed colonies did not. The bacterium Aurantimonas coralicida, causative agent of the disease, was present on H. opuntia sampled close to, and away from diseased corals, indicating that the alga serves as a reservoir for this pathogen. Our results suggest that the spread of macroalgae on coral reefs could account for the elevated incidence of coral diseases over past decades and that reduction of macroalgal abundance could help control coral epizootics.