Recurrent disturbances, recovery trajectories, and resilience of coral assemblages on a South Central Pacific reef

Coral reefs are increasingly threatened by various disturbances, and a critical challenge is to determine their ability for resistance and resilience. Coral assemblages in Moorea, French Polynesia, have been impacted by multiple disturbances (one cyclone and four bleaching events between 1991 and 2006). The 1991 disturbances caused large declines in coral cover (~51% to ~22%), and subsequent colonization by turf algae (~16% to ~49%), but this phase-shift from coral to algal dominance has not persisted. Instead, the composition of the coral community changed following the disturbances, notably favoring an increased cover of Porites, reduced cover of Montipora and Pocillopora, and a full return of Acropora; in this form, the reef returned to pre-disturbance coral cover within a decade. Thus, this coral assemblage is characterized by resilience in terms of coral cover, but plasticity in terms of community composition.     

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.

     

Influence of the coral reef assemblages on the spatial distribution of echinoderms in a gradient of human impacts along the tropical Mexican Pacific

Fourteen species of echinoderms and their relationships to the benthic structure of the coral reefs were assessed at 27 sites—with different levels of human disturbances—along the coast of the Mexican Central Pacific. Diadema mexicanum and Phataria unifascialis were the most abundant species. The spatial variation of the echinoderm assemblages showed that D. mexicanum, Eucidaris thouarsii, P. unifascialis, Centrostephanus coronatus, Toxopneustes roseus, Holothuria fuscocinerea, Cucumaria flamma, and Echinometra vanbrunti accounted for the dissimilarities among the sites. The spatial variation among the sites was mainly explained by the cover of the hard corals (Porites, Pocillopora, Pavona, Psammocora), different macroalgae species (turf, encrusting calcareous algae, articulated calcareous algae, fleshy macroalgae), sponges, bryozoans, rocky, coral rubble, sand, soft corals (hydrocorals and octocorals), Tubastrea coccinea coral, Balanus spp., and water depth. The coverage of Porites, Pavona, and Pocillopora corals, soft coral, rock, and Balanos shows a positive relationship with the sampling sites included within the natural protected area with low human disturbances. Contrary, fleshy macroalgae, sponges, and soft coral show a positive relationship with higher disturbance sites. The results presented here show the importance of protecting the structural heterogeneity of coral reef habitats because it is a significant factor for the distribution of echinoderm species and can contribute to the design of conservation programs for the coral reef ecosystem.     

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.     

Contrasting rates of coral recovery and reassembly in coral communities on the Great Barrier Reef

Changes in the relative abundances of coral taxa during recovery from disturbance may cause shifts in essential ecological processes on coral reefs. Coral cover can return to pre-disturbance levels (coral recovery) without the assemblage returning to its previous composition (i.e., without reassembly). The processes underlying such changes are not well understood due to a scarcity of long-term studies with sufficient taxonomic resolution. We assessed the trajectories and time frames for coral recovery and reassembly of coral communities following disturbances, using modeled trajectories based on data from a broad spatial and temporal monitoring program. We studied coral communities at six reefs that suffered substantial coral loss and subsequently regained at least 50 % of their pre-disturbance coral cover. Five of the six communities regained their coral cover and the rates were remarkably consistent, taking 7–10 years. Four of the six communities reassembled to their pre-disturbance composition in 8–13 years. The coral communities at three of the reefs both regained coral cover and reassembled ten years. The trajectories of two communities suggested that they were unlikely to reassemble and the remaining community did not regain pre-disturbance coral cover. The communities that regained coral cover and reassembled had high relative abundance of tabulate Acropora spp. Coral communities of this composition appear likely to persist in a regime of pulse disturbances at intervals of ten years or more. Communities that failed to either regain coral cover or reassemble were in near-shore locations and had high relative abundance of Porites spp. and soft corals. Under current disturbance regimes, these communities are unlikely to re-establish their pre-disturbance community composition.     

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.     

Differential thermal tolerance between algae and corals may trigger the proliferation of algae in coral reefs

Marine heatwaves can lead to rapid changes in entire communities, including in the case of shallow coral reefs the potential overgrowth of algae. Here we tested experimentally the differential thermal tolerance between algae and coral species from the Red Sea through the measurement of thermal performance curves and the assessment of thermal limits. Differences across functional groups (algae vs. corals) were apparent for two key thermal performance metrics. First, two reef‐associated algae species (Halimeda tuna and Turbinaria ornata) had higher lethal thermal limits than two coral species (Pocillopora verrucosa and Stylophora pistillata) conferring those species of algae with a clear advantage during heatwaves by surpassing the thermal threshold of coral survival. Second, the coral species had generally greater deactivation energies for net and gross primary production rates compared to the algae species, indicating greater thermal sensitivity in corals once the optimum temperature is exceeded. Our field surveys in the Red Sea reefs before and after the marine heatwave of 2015 show a change in benthic cover mainly in the southern reefs, where there was a decrease in coral cover and a concomitant increase in algae abundance, mainly turf algae. Our laboratory and field observations indicate that a proliferation of algae might be expected on Red Sea coral reefs with future ocean warming.     

Vermetid gastropods reduce foraging by herbivorous fishes on algae on coral reefs

Vermetid gastropods have the potential to reduce foraging by herbivorous fishes on algae on coral reefs because they produce mucous nets that cover the surfaces of coral skeletons, potentially inhibiting foraging by fishes. We assessed this possibility using both observational and experimental approaches in Moorea, French Polynesia. Foraging rates of herbivorous fishes (total number of bites by all species per minute) were recorded in plots that varied naturally in the cover of vermetid mucous nets. This study, done at six sites, revealed that foraging on algal turf declined with increasing cover of vermetid mucous nets, ranging from ~2 to 22 bites m^?2 min^?1 at 0 % coverage to 0–5 bites m^?2 min^?1 at 100 % coverage. The magnitude of this effect of vermetid nets varied among microhabitats (high, mid, and low bommies) and sites, presumably due to variation in the intensity of herbivory. Experimental removal of vermetid mucous nets from plots more than doubled the foraging intensity on turf algae relative to when vermetid nets were present at high (≥70 %) cover. Our results indicate that algal turf on coral reefs may benefit from associational refuge from grazing provided by vermetid gastropods, which might in turn harm corals via increased competition with algal turf.     

Benthic Reef Primary Production in Response to Large Amplitude Internal Waves at the Similan Islands (Andaman Sea,Thailand)

Coral reefs are facing rapidly changing environments, but implications for reef ecosystem functioning and important services, such as productivity, are difficult to predict. Comparative investigations on coral reefs that are naturally exposed to differing environmental settings can provide essential information in this context. One prevalent phenomenon regularly introducing alterations in water chemistry into coral reefs are internal waves. This study therefore investigates the effect of large amplitude internal waves (LAIW) on primary productivity in coral reefs at the Similan Islands (Andaman Sea, Thailand). The LAIW-exposed west sides of the islands are subjected to sudden drops in water temperature accompanied by enhanced inorganic nutrient concentrations compared to the sheltered east. At the central island, Ko Miang, east and west reefs are only few hundred meters apart, but feature pronounced differences. On the west lower live coral cover (-38 %) coincides with higher turf algae cover (+64 %) and growth (+54 %) compared to the east side. Turf algae and the reef sand-associated microphytobenthos displayed similar chlorophyll a contents on both island sides, but under LAIW exposure, turf algae exhibited higher net photosynthesis (+23 %), whereas the microphytobenthos displayed reduced net and gross photosynthesis (-19 % and -26 %, respectively) accompanied by lower respiration (-42 %). In contrast, the predominant coral Porites lutea showed higher chlorophyll a tissues contents (+42 %) on the LAIW-exposed west in response to lower light availability and higher inorganic nutrient concentrations, but net photosynthesis was comparable for both sides. Turf algae were the major primary producers on the west side, whereas microphytobenthos dominated on the east. The overall primary production rate (comprising all main benthic primary producers) was similar on both island sides, which indicates high primary production variability under different environmental conditions.     

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.     

Coral reef resilience to thermal stress in the Eastern Tropical Pacific

Coral reefs worldwide are threatened by thermal stress caused by climate change. Especially devastating periods of coral loss frequently occur during El Niño‐Southern Oscillation (ENSO) events originating in the Eastern Tropical Pacific (ETP). El Niño‐induced thermal stress is considered the primary threat to ETP coral reefs. An increase in the frequency and intensity of ENSO events predicted in the coming decades threatens a pan‐tropical collapse of coral reefs. During the 1982–1983 El Niño, most reefs in the Galapagos Islands collapsed, and many more in the region were decimated by massive coral bleaching and mortality. However, after repeated thermal stress disturbances, such as those caused by the 1997–1998 El Niño, ETP corals reefs have demonstrated regional persistence and resiliency. Using a 44 year dataset (1970–2014) of live coral cover from the ETP, we assess whether ETP reefs exhibit the same decline as seen globally for other reefs. Also, we compare the ETP live coral cover rate of change with data from the maximum Degree Heating Weeks experienced by these reefs to assess the role of thermal stress on coral reef survival. We find that during the period 1970–2014, ETP coral cover exhibited temporary reductions following major ENSO events, but no overall decline. Further, we find that ETP reef recovery patterns allow coral to persist under these El Niño‐stressed conditions, often recovering from these events in 10–15 years. Accumulative heat stress explains 31% of the overall annual rate of change of living coral cover in the ETP. This suggests that ETP coral reefs have adapted to thermal extremes to date, and may have the ability to adapt to near‐term future climate‐change thermal anomalies. These findings for ETP reef resilience may provide general insights for the future of coral reef survival and recovery elsewhere under intensifying El Niño scenarios.     

Cyclone effects on coral reef habitats in New Caledonia (South Pacific)

The impacts of the unusually strong Cyclone Erica (March 2003) on coral reef habitats at a site located on the northwest coast of New Caledonia (South Pacific) were assessed using a 6-year data set (2002–2007). We examined the interannual variations of key variables describing reef habitats (live hard and soft corals, dead corals in place, coral debris, algae and relative proportion of mechanically vulnerable and resistant live hard corals). The cyclone-induced disturbances of habitats differed according to three reef types: patch reefs, barrier reefs far from passes (more than 3 km from the nearest pass) and barrier reefs near passes (less than 3 km from the nearest pass). Short-term mechanical damage was detected on the three-dimensional structure of reef habitats with a notable shift from a community dominated by mechanically vulnerable corals to one dominated by resistant corals on barrier reefs far from passes. The history of habitats and their pre-disturbance characteristics, in link with local hydrodynamics, was found to influence their short-term susceptibility to extreme events such as cyclones. However, the most significant effects appeared in the midterm (within 2 years after the cyclone) as the cover of live hard corals significantly decreased by approximately 45% between 2002 and 2004 on all reef types. The short- and midterm disturbances of coral reef habitats are discussed with regard to published temporal variations in reef fish assemblages, underlining the delayed effects of this cyclonic event on fish as well as benthic habitats. Coral reef habitats and live corals had shown significant patterns of recovery 4 years after the cyclone, followed by similar recovery in fish community, suggesting good resilience in a face of this major natural disturbance in an area under moderate anthropogenic pressure.     

Effects of the 2000 southern Mozambique floods on a marginal coral community: the case at Xai-Xai

In early 2000 the southern part of Mozambique suffered the worst flooding in 50 years, causing fatalities and considerable material loss. This study aimed to investigate the impact of this flood on the coral communities in Xai-Xai lagoon. Benthic cover was assessed in January 2000 (before the floods) and September 2000 (after the floods) using the line intercept transect technique. A decrease in hard coral cover of the order of 58.5% was observed. The soft coral community was significantly affected, with a decrease in percentage cover of 90.4%. Coralline algae also decreased by 85.1%. All other categories increased in percentage cover: turf algae (164.4%), other invertebrates (e.g. sponges, sea urchins — 111.1%), fleshy algae (80.4%), rubble (34.4%) and dead coral (379.0%). The main causes of this degradation were the reduced water salinity and the large amount of sediment discharged by the Limpopo River. Some massive (e.g. Porites, Favia, Favites and Goniopora) and encrusting (e.g. Echinopora) hard coral genera seemed less affected, suggesting an elevated capacity to cope with this kind of stress through mucus-sheet formation. The extent of the flood effect on other reefs on the southern Mozambique coast is discussed and a monitoring programme is proposed.     

Interaction of herbivory and seasonality on the dynamics of Caribbean macroalgae

Many Caribbean coral reefs are undergoing a phase shift from coral to macroalgal dominance. Understanding the processes driving changes in algal abundance and community structure requires clarification of the relative effects of top-down (e.g., herbivory) and bottom-up processes (e.g., light, temperature, and nutrients). To date, a number of studies have examined the relative effects of grazing versus nutrification but interactions between herbivory and natural, seasonal fluctuations in temperature and light have not been investigated. This study considered the dynamics of three Caribbean macroalgal species [Lobophora variegata (Lamouroux), Dictyota pulchella (H?rnig and Schnetter), and Halimeda opuntia (Linnaeus)] and algal turf. A field experiment was established to measure species-specific algal dynamics (changes in abundance) over 13?months in the presence and absence of herbivory. Both herbivory and seasonal changes were important processes controlling macroalgal and turf abundance. Water temperature and light had a key role on D. pulchella; this species?? abundance significantly increased in the summer, when water temperature and light were the highest, and decreased during winter. Surprisingly, herbivory did not seem to control D. pulchella directly. However, herbivory was the most important process controlling the abundance of L. variegata, H. opuntia, and turf. The abundance of both algal species was correlated with seasonal changes in the environment, but was depleted outside cages throughout the year. The abundance of H. opuntia was positively correlated with temperature and light, but there was no statistical interaction between drivers. The statistical interaction between temperature and light was significant for the abundance of L. variegata and turf, but algal abundance declined as both factors increased. Overall, macroalgal and turf cover were mainly controlled by herbivory, while community structure (which species contributed to the overall cover) was largely influenced by seasonal changes in temperature and light.     

Detriments to post-bleaching recovery of corals

Predicting the response of coral reefs to large-scale mortality induced by climate change will depend greatly on the factors that influence recovery after bleaching events. We experimentally transplanted hard corals from a shallow reef with highly variable seawater temperature (23–36°C) to three unfished marine parks and three fished reefs with variable coral predator abundance and benthic cover. The transplanted corals were fragmented colonies collected from a reef that was relatively undisturbed by the 1997–1998 warm-water temperature anomaly, one of the most extreme thermal events of the past century, and it was assumed that they would represent corals likely to succeed in the future temperature environment. We examined the effects of four taxa, two fragment sizes, an acclimation period, benthic cover components, predators and tourists on the survival of the coral fragments. We found the lowest survival of transplants occurred in the unfished marine parks and this could be attributed to predation and not tourist damage. The density of small coral recruits approximately 6 months after the spawning season was generally moderate (~40–60/m²), and not different on fished and unfished reefs. Coral recovery between 1998 and 2002 was variable (0–25%), low (mean of 6.5%), and not different between fished and unfished reefs. There was high variability in coral mortality among the three unfished areas despite low variation in estimates of predator biomass, with the highest predation occurring in the Malindi MNP, a site with high coralline algal cover. Stepwise multiple regression analysis with 14 variables of coral predators and substratum showed that coralline algae was positively, and turf algae negatively associated with mortality of the transplants, with all other variables being statistically insignificant. This suggests that alternate food resources and predator choices are more important than predator biomass in determining coral survival. Nonetheless, large predatory fish in areas dominated by coralline algae may considerably retard recovery of eurythermal corals. This will not necessarily retard total hard coral recovery, as other more predator-tolerant taxa can recover. Based on the results, global climate change will not necessarily favor eurythermal over stenothermal coral taxa in remote or unfished reefs, where predation is a major cause of coral mortality.     

Black reefs: iron-induced phase shifts on coral reefs

The Line Islands are calcium carbonate coral reef platforms located in iron-poor regions of the central Pacific. Natural terrestrial run-off of iron is non-existent and aerial deposition is extremely low. However, a number of ship groundings have occurred on these atolls. The reefs surrounding the shipwreck debris are characterized by high benthic cover of turf algae, macroalgae, cyanobacterial mats and corallimorphs, as well as particulate-laden, cloudy water. These sites also have very low coral and crustose coralline algal cover and are call black reefs because of the dark-colored benthic community and reduced clarity of the overlying water column. Here we use a combination of benthic surveys, chemistry, metagenomics and microcosms to investigate if and how shipwrecks initiate and maintain black reefs. Comparative surveys show that the live coral cover was reduced from 40 to 60% to <10% on black reefs on Millennium, Tabuaeran and Kingman. These three sites are relatively large (>0.75 km²). The phase shift occurs rapidly; the Kingman black reef formed within 3 years of the ship grounding. Iron concentrations in algae tissue from the Millennium black reef site were six times higher than in algae collected from reference sites. Metagenomic sequencing of the Millennium Atoll black reef-associated microbial community was enriched in iron-associated virulence genes and known pathogens. Microcosm experiments showed that corals were killed by black reef rubble through microbial activity. Together these results demonstrate that shipwrecks and their associated iron pose significant threats to coral reefs in iron-limited regions.     

“Choice” and destiny: the substrate composition and mechanical stability of settlement structures can mediate coral recruit fate in post-bleached reefs

Increasingly frequent and intense ocean warming events seriously test the buffer and recovery capacities of tropical coral reefs. Post-disturbance, available settlement structures on a reef (often dead coral skeletons) vary considerably in their mechanical stability and substrate composition, critically influencing coral recruit settlement choice and fate. In the wake of a coral mass mortality in the Lakshadweep archipelago, we examine (1) the relative availability of recruit settlement structures (from stable to unstable: reef platform, dead massive coral, consolidated rubble, dead corymbose coral, dead tabular coral, and unconsolidated rubble) in 12 recovering reefs across three atolls in the archipelago, (2) the substrate composition [crustose coralline algae (CCA), mixed turf, macroalgae] of these structural forms, and (3) whether the choice and fate of young coral are mediated by the substrate and stability of different structural forms. For this, we measured the abundance and distribution of recruit (<1 cm), juvenile (1–5 cm), and young adult (5–10) corals of 24 common coral genera. Four years after the mass mortality, reefs differed considerably in composition of settlement structures. The structures themselves varied significantly in substrate cover with dead tables largely covered in CCA [60 ± 6.05 % (SE)] and dead corymbose coral dominated by mixed turf (61.83 ± 3.8 %). The youngest visible recruits (<1 cm) clearly preferred CCA-rich structures such as dead massives and tables. However, older size classes were rarely found on unstable structures (strongly “avoiding” tables, Ivlev’s electivity index, E = ?0.5). Our results indicate that while substrate cover might mediate coral choice, the mechanical stability of settlement structures is critical in determining post-settlement coral survival. The composition and availability of settlement structures on a reef may serve as a characteristic signature of its recovery potential, aiding in assessments of reef resilience.     

Decadal coral community reassembly on an African fringing reef

Changes in the cover of the dominant hard coral taxa were studied on seven Kenyan back reefs over 20 yr. All factors of time, taxa, site, and their interactions were statistically significant and the 1998 temperature anomaly caused the greatest community changes. The 1998 disturbance changes reflected a classic coral succession, which included partial or little mortality and persistence of stress tolerant (massive and submassive growth forms) and early colonization by weedy taxa (pocilloporids). Nevertheless, competitive taxa had high and full mortality and the expected dominance of acroporids was inhibited even ~13 yr after the disturbance. So, while total hard coral cover displayed the expected logistic recovery where maximum cover was reached <10 yr after the disturbance, the poor recovery of competitive dominants resulted in less than expected coral cover. A number of stress-resistant and weedy taxa (poritids, agaricidae, faviids, and pocilloporids) are expected to dominate the composition of these reefs in the future. Nevertheless, three submassive faviids and branching Porites began to decline toward the end of the time series, indicating further stress after 1998. Increased algal cover and other unstudied factors, including milder warming, may explain these changes. The patterns of change on this continental fringing reef differ from recovery of more remote, offshore islands. This probably reflects low acroporid dominance and recruitment limitations associated with greater anthropogenic influences of high sea urchin grazing and terrestrial runoff.     

Microbial to reef scale interactions between the reef-building coral Montastraea annularis and benthic algae

Competition between reef-building corals and benthic algae is of key importance for reef dynamics. These interactions occur on many spatial scales, ranging from chemical to regional. Using microprobes, 16S rDNA pyrosequencing and underwater surveys, we examined the interactions between the reef-building coral Montastraea annularis and four types of benthic algae. The macroalgae Dictyota bartayresiana and Halimeda opuntia, as well as a mixed consortium of turf algae, caused hypoxia on the adjacent coral tissue. Turf algae were also associated with major shifts in the bacterial communities at the interaction zones, including more pathogens and virulence genes. In contrast to turf algae, interactions with crustose coralline algae (CCA) and M. annularis did not appear to be antagonistic at any scale. These zones were not hypoxic, the microbes were not pathogen-like and the abundance of coral-CCA interactions was positively correlated with per cent coral cover. We propose a model in which fleshy algae (i.e. some species of turf and fleshy macroalgae) alter benthic competition dynamics by stimulating bacterial respiration and promoting invasion of virulent bacteria on corals. This gives fleshy algae a competitive advantage over corals when human activities, such as overfishing and eutrophication, remove controls on algal abundance. Together, these results demonstrate the intricate connections and mechanisms that structure coral reefs.     

Disturbance and the dynamics of coral cover on the Great Barrier Reef (1995-2009)

Coral reef ecosystems worldwide are under pressure from chronic and acute stressors that threaten their continued existence. Most obvious among changes to reefs is loss of hard coral cover, but a precise multi-scale estimate of coral cover dynamics for the Great Barrier Reef (GBR) is currently lacking. Monitoring data collected annually from fixed sites at 47 reefs across 1300 km of the GBR indicate that overall regional coral cover was stable (averaging 29% and ranging from 23% to 33% cover across years) with no net decline between 1995 and 2009. Subregional trends (10-100 km) in hard coral were diverse with some being very dynamic and others changing little. Coral cover increased in six subregions and decreased in seven subregions. Persistent decline of corals occurred in one subregion for hard coral and Acroporidae and in four subregions in non-Acroporidae families. Change in Acroporidae accounted for 68% of change in hard coral. Crown-of-thorns starfish (Acanthaster planci) outbreaks and storm damage were responsible for more coral loss during this period than either bleaching or disease despite two mass bleaching events and an increase in the incidence of coral disease. While the limited data for the GBR prior to the 1980's suggests that coral cover was higher than in our survey, we found no evidence of consistent, system-wide decline in coral cover since 1995. Instead, fluctuations in coral cover at subregional scales (10-100 km), driven mostly by changes in fast-growing Acroporidae, occurred as a result of localized disturbance events and subsequent recovery.