Coral bleaching, reef fish community phase shifts and the resilience of coral reefs

The 1998 global coral bleaching event was the largest recorded historical disturbance of coral reefs and resulted in extensive habitat loss. Annual censuses of reef fish community structure over a 12-year period spanning the bleaching event revealed a marked phase shift from a prebleach to postbleach assemblage. Surprisingly, we found that the bleaching event had no detectable effect on the abundance, diversity or species richness of a local cryptobenthic reef fish community. Furthermore, there is no evidence of regeneration even after 5–35 generations of these short-lived species. These results have significant implications for our understanding of the response of coral reef ecosystems to global warming and highlight the importance of selecting appropriate criteria for evaluating reef resilience.     

Sediment addition drives declines in algal turf yield to herbivorous coral reef fishes: implications for reefs and reef fisheries

Coral Reefs - Coral reefs around the world are changing rapidly, with overfishing of herbivorous fishes and increased sediment inputs being two of the major local-scale stressors. We therefore...     

Forecasting decadal changes in sea surface temperatures and coral bleaching within a Caribbean coral reef

Elevated sea surface temperature (SST) caused by global warming is one of the major threats to coral reefs. While increased SST has been shown to negatively affect the health of coral reefs by increasing rates of coral bleaching, how changes to atmospheric heating impact SST distributions, modified by local flow environments, has been less understood. This study aimed to simulate future water flow patterns and water surface heating in response to increased air temperature within a coral reef system in Bocas del Toro, Panama, located within the Caribbean Sea. Water flow and SST were modeled using the Delft3D-FLOW© computer simulation package. Locally measured physical parameters, including bathymetry, astronomic tidal forcing, and coral habitat distribution were input into the model and water flow, and SST was simulated over a four-month period under present day, as well as projected warming scenarios in 2020s, 2050s, and 2080s. Changes in SST, and hence the thermal stress to corals, were quantified by degree heating weeks. Results showed that present-day reported bleaching sites were consistent with localized regions of continuous high SST. Regions with highest SST were located within shallow coastal sites adjacent to the mainland or within the interior of the bay, and characterized by low currents with high water retention times. Under projected increases in SSTs, shallow reef areas in low flow regions were found to be hot spots for future bleaching.     

Herbivory on coral reefs: algal susceptibility to herbivorous fishes

Summary The susceptibility of several tropical algal species to fish grazing was studied on the Belizean barrier reef off the Caribbean coast of Central America. Short-term transplant experiments indicate that plant species vary markedly in their rates of biomass loss to grazing by a shallow-water guild of herbivorous fishes. Algal species transplanted from habitats with low grazing pressure are highly susceptible to grazing, while species occurring in habitats with high herbivore densities are highly resistant to grazing. Algal species show differential susceptibility to grazing by two major components of the tropical herbivore guild, Acanthurus (surgeonfishes) and Sparisoma (parrotfishes).Variability in plant susceptibility to grazing by herbivorous fishes was not clearly correlated with morphological or chemical characteristics that have been previously suggested as plant defenses against herbivory. Plants found to be highly resistant to fish grazing, such as Halimeda, exhibit both morphological characteristics and secondary chemical compounds which do appear to reduce herbivory. In contrast, species of Caulerpa, Sargassum, Turbinaria, and Padina, which also possess alleged morphological and/or chemical defenses, are nevertheless highly susceptible to fish grazing.     

Comparisons of the 1995 and 1998 coral bleaching events on the patch reefs of San Salvador Island, Bahamas

Coral patch reefs around San Salvador Island, Bahamas have been monitored with the aid of Earthwatch volunteers three times a year since 1992. During that period two significant mass bleaching events occurred: autumn 1995, and late summer 1998. Elsewhere in 1995, bleaching was caused by higher-than-normal summer sea temperatures; in San Salvador, however, temperatures were normal. In 1998 a prolonged period of higher-than-normal sea temperatures preceded bleaching on San Salvador and worldwide. During the 1995 event, one of the monitored reefs had twice the percentage of coral colonies bleached as the other two. Bleaching was more evenly distributed among the reefs during the 1998 event. In 1995 Agaricia agaricites was significantly more affected than other coral species, with almost 50% of all its colonies showing bleaching. Bleaching was more evenly spread among coral species in 1998, with five species showing bleaching on more than 40% of their colonies. Bleaching began on Millepora as early as August during the 1998 event and progressed to other species through the remainder of the autumn. In 1995 bleaching was not seen until late autumn and appeared to impact all affected species at about the same time. Recovery from the 1995 event was complete: no coral death or damage above normal background levels were seen. In the 1998 event, all Acropora cervicornis on the monitored reefs died and A. palmata was severely damaged. Millepora sp. lost almost half of their live tissue, and Montastraea sp. showed significant tissue damage following this event. Phototransect analysis suggests that more than 20% of total live tissue on affected species died during the 1998 event. A. cervicornis has demonstrated no re-growth from 1998 to 2000 on monitored reefs. Monitoring has suggested significant differences in causes and courses in these two events.     

Coral reef bleaching in the 1980s and possible connections with global warming

Scleractinian corals and their symbiotic dinoflagellate algae build massive, wave-resistant coral reefs that are pre-eminent in shallow tropical seas. This mutualism is especially sensitive to numerous environmental stresses, and has been disrupted frequently during the past decade. Increased seawater temperatures have been proposed as the most likely cause of coral reef bleaching, and it has been suggested that the recent large-scale disturbances are the first biological indication of global warming. This article describes recent bleaching events and their possible link with sea warming and other environmental stresses, and offers some speculation on the fate of coral reefs if the Earth enters a sustained period of warming.     

Coral disease following massive bleaching in 2005 causes 60% decline in coral cover on reefs in the US Virgin Islands

In the northeast Caribbean, doldrum-like conditions combined with elevated water temperatures in the summer/fall 2005 created the most severe coral bleaching event ever documented within this region. Video monitoring of 100 randomly chosen, permanent transects at five study sites in the US Virgin Islands revealed over 90% of the scleractinian coral cover showed signs of thermal stress by paling or becoming completely white. Lower water temperatures in October allowed some re-coloring of corals; however, a subsequent unprecedented regional outbreak of coral disease affected all sites. Five known diseases or syndromes were recorded; however, most lesions showed signs similar to white plague. Nineteen scleractinian species were affected by disease, with >90% of the disease-induced lesions occurring on the genus Montastraea. The disease outbreak peaked several months after the onset of bleaching at all sites but did not occur at the same time. The mean number of disease-induced lesions increased 51-fold and the mean area of disease-associated mortality increased 13-fold when compared with pre-bleaching disease levels. In the 12 months following the onset of bleaching, coral cover declined at all sites (average loss: 51.5%, range: 42.4–61.8%) reducing the five-site average from 21.4% before bleaching to 10.3% with most mortality caused by white plague disease, not bleaching. Continued losses through October 2007 reduced the average coral cover of the five sites to 8.3% (average 2-year loss: 61.1%, range: 53.0–79.3%). Mean cover by M. annularis (complex) decreased 51%, Colpophyllia natans 78% and Agaricia agaricites 87%. Isolated disease outbreaks have been documented before in the Virgin Islands, but never as widespread or devastating as the one that occurred after the 2005 Caribbean coral-bleaching event. This study provides insight into the effects of continued seawater warming and subsequent coral bleaching events in the Caribbean and highlights the need to understand links between coral bleaching and disease.     

Regenerative functions and microbial ecology of coral reefs: Labelled bacteria in a coral reef microcosm

Representative coral reef organisms and substrata assembled in a laboratory microcosm removed radioactively labelled bacteria from water circulated over them. A similar experiment with a reef clam and its algal-encrusted base gave similar results. Biochemical fractionation of selected organisms in these experiments suggested digestion and possible assimilation of bacterial proteins. In view of previous results concerning the microbial ecology of coral reefs, it is suggested that reef infaunal metazoa are adapted to utilize internal sedimentary processes and regenerative functioning through suspension- (and deposit-)feeding mechanisms. A model ecosystem is presented to suggest the possible feedback of these mechanisms as they operate within a reef.     

Facilitation in Caribbean coral reefs: high densities of staghorn coral foster greater coral condition and reef fish composition

Recovery of the threatened staghorn coral (Acropora cervicornis) is posited to play a key role in Caribbean reef resilience. At four Caribbean locations (including one restored and three extant populations), we quantified characteristics of contemporary staghorn coral across increasing conspecific densities, and investigated a hypothesis of facilitation between staghorn coral and reef fishes. High staghorn densities in the Dry Tortugas exhibited significantly less partial mortality, higher branch growth, and supported greater fish abundances compared to lower densities within the same population. In contrast, partial mortality, branch growth, and fish community composition did not vary with staghorn density at the three other study locations where staghorn densities were lower overall. This suggests that density-dependent effects between the coral and fish community may only manifest at high staghorn densities. We then evaluated one facilitative mechanism for such density-dependence, whereby abundant fishes sheltering in dense staghorn aggregations deliver nutrients back to the coral, fueling faster coral growth, thereby creating more fish habitat. Indeed, dense staghorn aggregations within the Dry Tortugas exhibited significantly higher growth rates, tissue nitrogen, and zooxanthellae densities than sparse aggregations. Similarly, higher tissue nitrogen was induced in a macroalgae bioassay outplanted into the same dense and sparse aggregations, confirming greater bioavailability of nutrients at high staghorn densities. Our findings inform staghorn restoration efforts, suggesting that the most effective targets may be higher coral densities than previously thought. These coral-dense aggregations may reap the benefits of positive facilitation between the staghorn and fish community, favoring the growth and survivorship of this threatened species.     

Elevated temperatures and bleaching on a high latitude coral reef: the 1988 Bermuda event

Sea temperatures were normal in Bermuda during 1987, when Bermuda escaped the episodes of coral bleaching which were prevalent throughout the Caribbean region. Survey transecs in 1988 on 4–6 m reefs located on the rim margin and on a lagoonal patch reef revealed bleaching only of zoanthids between May and July. Transect and tow surveys in August and September revealed bleaching of several coral species;Millepora alcicornis on rim reefs was the most extensively affected. The frequency of bleaching in this species,Montastrea annularis and perhapsDiploria labyrinthiformis was significantly higher on outer reefs than on inshore reefs. This bleaching period coincided with the longest period of elevated sea temperatures in Bermuda in 38 years (28.9–30.9°C inshore, >28° offshore). By December, when temperatures had returned to normal, bleaching of seleractinians continued, but bleaching ofM. alcicornis on the outer reefs was greatly reduced. Our observations suggest that corals which normally experience wide temperature ranges are less sensitive to thermal stress, and that high-latitude reef corals are sensitive to elevated temperatures which are within the normal thermal range of corals at lower latitudes.     

Coral reef resilience differs among islands within the Gulf of Mannar,southeast India,following successive coral bleaching events

Coral Reefs - We used a 12-yr data set of benthic cover (2005–2017), spanning two bleaching events, to assess changes in benthic cover and coral community composition along 21 islands within...     

Reassessing the trophic role of reef sharks as apex predators on coral reefs

Apex predators often have strong top-down effects on ecosystem components and are therefore a priority for conservation and management. Due to their large size and conspicuous predatory behaviour, reef sharks are typically assumed to be apex predators, but their functional role is yet to be confirmed. In this study, we used stomach contents and stable isotopes to estimate diet, trophic position and carbon sources for three common species of reef shark (Triaenodon obesus, Carcharhinus melanopterus and C. amblyrhynchos) from the Great Barrier Reef (Australia) and evaluated their assumed functional role as apex predators by qualitative and quantitative comparisons with other sharks and large predatory fishes. We found that reef sharks do not occupy the apex of coral reef food chains, but instead have functional roles similar to those of large predatory fishes such as snappers, emperors and groupers, which are typically regarded as high-level mesopredators. We hypothesise that a degree of functional redundancy exists within this guild of predators, potentially explaining why shark-induced trophic cascades are rare or subtle in coral reef ecosystems. We also found that reef sharks participate in multiple food webs (pelagic and benthic) and are sustained by multiple sources of primary production. We conclude that large conspicuous predators, be they elasmobranchs or any other taxon, should not axiomatically be regarded as apex predators without thorough analysis of their diet. In the case of reef sharks, our dietary analyses suggest they should be reassigned to an alternative trophic group such as high-level mesopredators. This change will facilitate improved understanding of how reef communities function and how removal of predators (e.g., via fishing) might affect ecosystem properties.     

Foragers versus farmers: contrasting effects of two behavioural groups of herbivores on coral reefs

Herbivorous fishes have been attributed a central role in structuring benthic communities on coral reefs. However, the relative importance of different behavioural groups of herbivores may differ and their interactions may be complex. This study focuses on an experiment that discriminates between two groups of herbivorous fish: (1) “Foragers” (relatively mobile, schooling grazers, including parrotfishes and surgeonfishes) and (2) “Farmers” (highly site-attached, territorial species, primarily damselfishes). Preliminary observations at Kimbe Bay (Papua New Guinea) showed that both groups were common, and that farmers defended areas from foragers and maintained algal communities that were distinct from the surrounding undefended substratum. An orthogonal combination of a farmer removal treatment and a forager exclusion treatment was applied to isolate their separate effects on algae and corals, and to determine whether farmer territory composition results from forager exclusion or algal cultivation. The experiment showed that foragers had quantitatively greater and qualitatively different effects on sessile benthic community structure than farmers. Where foragers were excluded, there were substantial increases in the cover and biomass of macro-algae and a decline in some corals, regardless of the presence of farmers. Where farmers were removed there was a moderate decline in the cover of some food algal species, regardless of whether foragers had access. No effect of the exclusion of foragers by farmers could be detected. Our results support prevailing views that foragers have a major impact on coral reefs and farmers cultivate selected algae, but challenge the hypothesis that damselfish influence habitat structure by moderating forager disturbance.     

Coral bleaching: one disturbance too many for near-shore reefs of the Great Barrier Reef

The dynamic nature of coral communities can make it difficult to judge whether a reef system is resilient to the current disturbance regime. To address this question of resilience for near-shore coral communities of the Great Barrier Reef (Australia) a data set consisting of 350 annual observations of benthic community change was compiled from existing monitoring data. These data spanned the period 1985–2007 and were derived from coral reefs within 20 km of the coast. During years without major disturbance events, cover increase of the Acroporidae was much faster than it was for other coral families; a median of 11% per annum compared to medians of less than 4% for other coral families. Conversely, Acroporidae were more severely affected by cyclones and bleaching events than most other families. A simulation model parameterised with these observations indicated that while recovery rates of hard corals were sufficient to compensate for impacts associated with cyclones and crown-of-thorns starfish, the advent of mass bleaching has lead to a significant change in the composition of the community and a rapid decline in hard coral cover. Furthermore, if bleaching events continue to occur with the same frequency and severity as in the recent past, the model predicts that the cover of Acroporidae will continue to decline. Although significant cover of live coral remains on near-shore reefs, and recovery is observed during inter-disturbance periods, it appears that this system will not be resilient to the recent disturbance regime over the long term. Conservation strategies for coral reefs should focus on both mitigating local factors that act synergistically to increase the susceptibility of Acroporidae to climate change while promoting initiatives that maximise the recovery potential from inevitable disturbances.     

Direct and indirect effects of high pCO2 on algal grazing by coral reef herbivores from the Gulf of Aqaba (Red Sea)

Grazing on marine macroalgae is a key structuring process for coral reef communities. However, ocean acidification from rising atmospheric CO₂ concentrations is predicted to adversely affect many marine animals, while seaweed communities may benefit and prosper. We tested how exposure to different pCO₂ (400, 1,800 and 4,000 μatm) may affect grazing on the green alga Ulva lactuca by herbivorous fish and sea urchins from the coral reefs in the northern Gulf of Aqaba (Red Sea), either directly, by changing herbivore behaviour, or indirectly via changes in algal palatability. We also determined the effects of pCO₂ on algal tissue concentrations of protein and the grazing-deterrent secondary metabolite dimethylsulfoniopropionate (DMSP). Grazing preferences and overall consumption were tested in a series of multiple-choice feeding experiments in the laboratory and in situ following exposure for 14 d (algae) and 28 d (herbivores). 4,000 μatm had a significant effect on the biochemical composition and palatability of U. lactuca. No effects were observed at 1,800 relative to 400 μatm (control). Exposure of U. lactuca to 4,000 μatm resulted in a significant decrease in protein and increase in DMSP concentration. This coincided with a reduced preference for these algae by the sea urchin Tripneustes gratilla and different herbivorous fish species in situ (Acanthuridae, Siganidae and Pomacanthidae). No feeding preferences were observed for the rabbitfish Siganus rivulatus under laboratory conditions. Exposure to elevated pCO₂ had no direct effect on the overall algal consumption by T. gratilla and S. rivulatus. Our results show that CO₂ has the potential to alter algal palatability to different herbivores which could have important implications for algal abundance and coral community structure. The fact that pCO₂ effects were observed only at a pCO₂ of 4,000 μatm, however, indicates that algal-grazer interactions may be resistant to predicted pCO₂ concentrations in the near future.     

Monoculture and mixed-species algal farms on a coral reef are maintained through intensive and extensive management by damselfishes

Algal farms maintained by different species of territorial herbivorous damselfishes vary in size, algal biomass, and species composition. To determine the factors that affect the structure of these farms, we compared farming behavior and intensity between two sympatric damselfish species, Stegastes nigricans (Lacepède) and S. obreptus (Whitley), which maintain two different types of farms in the Okinawa Islands, southern Japan. By weeding of indigestible algae and prompt exclusion of herbivorous fishes and grazing sea urchins, S. nigricans manage relatively small farms largely dominated by one algal crop species, Polysiphonia sp. In contrast, by delayed exclusion of herbivores, S. obreptus maintain larger farms invaded by diverse indigestible algae, such as Chondria sp. and Padina sp. As a result of intensive management, S. nigricans attains a higher algal biomass per area than does S. obreptus. This study provides a new perspective, as it suggests that damselfishes practise both intensive and extensive farming. The former results in small and highly productive monoculture farms of one palatable alga, while the latter leads to large, species-rich, mixed-culture farms.