High-frequency winter cooling and reef coral mortality during the Holocene climatic optimum

A detailed ecological, micro-structural and skeletal Sr/Ca study of a 3.42 m thick Goniopora reef profile from an emerged Holocene reef terrace at the northern South China Sea reveals at least nine abrupt massive Goniopora stress and mortality events occurred in winter during the 7.0-7.5 thousand calendar years before present (cal. ka BP) (within the Holocene climatic optimum). Whilst calculated Sr/Ca-SST (sea surface temperature) maxima during this period are comparable to those in the 1990s, Sr/Ca-SST minima are significantly lower, probably due to stronger winter monsoons. Such generally cooler winters, superimposed by further exceptional winter cooling on inter-annual to decadal scales, may have caused stress and mortality of the corals about every 50 years. Sea level rose by similar to 3.42 m during this period, with present sea-level reached at similar to 7.3 ka BP and a sea-level highstand of at least similar to 1.8 m occurred at similar to 7.0 ka. The results show that it took about 20-25 years for a killed Goniopora coral reef to recover. (C) 2004 Elsevier B.V. All rights reserved.

Coral mortality increases wave energy reaching shores protected by reef flats: Examples from the Seychelles

In the granitic Seychelles, many shores and beaches are fringed by coral reef flats which provide protection to shores from erosion by waves. The surfaces of these reef flats support a complex ecology. About 10 years ago their seaward zones were extensively covered by a rich coral growth, which reached approximately to mean low water level, but in 1998 this was largely killed by seawater warming. The resulting large expanses of dead coral skeletons in these locations are now disintegrating, and much of the subsequent modest recovery by new coral recruitment was set back by further mortalities. A mathematical model of wave energy reaching shorelines protected by coral reef flats has been applied to 14 Seychelles reefs. It is derived from equations which predict: (1) the raised water level, or wave set-up, on reef flats resulting from wave breaking, which depends upon offshore wave height and period, depth of still water over the reef flat and the reef crest profile, and (2) the decay of energy from reef edge to shoreline that is affected by width of reef flat, surface roughness, sea level rise and 'pseudo-sea level rise' created by increased depth resulting from disintegration of coral colonies. The model treats each reef as one entity, but because biota and zonation on reef flats are not homogenous, all reefs are divided into four zones. In each, cover by both living and dead biota was estimated for calculation of parameters, and then averaged to obtain input data for the model. All possible biological factors were taken into account, such as the ability of seagrass beds to grow upwards to match expected sea level rise, reduction in height of the reef flat in relation to sea level as zones of dead corals decay, and the observed 'rounding' of reef crests as erosion removes corals from those areas. Estimates were also made of all these factors for a time approximately a decade ago, representing a time before the mass coral mortality, and for approximately a decade in the future when the observed rapid state of dead coral colony disintegration is assumed to have reached an end point. Results of increased energy over the past decade explain observations of erosion in some sites in the Seychelles. Most importantly, it is estimated that the rise in energy reaching shores protected by fringing reefs will now accelerate more rapidly, such that the increase expected over the next decade will be approximately double than that seen over the past decade. (c) 2005 Elsevier Ltd. All rights reserved.

ENCORE: The effect of nutrient enrichment on coral reefs. Synthesis of results and conclusions

Coral reef degradation resulting from nutrient enrichment of coastal waters is of increasing global concern. Although effects of nutrients on coral reef organisms have been demonstrated in the laboratory, there is little direct evidence of nutrient effects on coral reef biota in situ. The ENCORE experiment investigated responses of coral reef organisms and processes to controlled additions of dissolved inorganic nitrogen (N) and/or phosphorus (P) on an offshore reef(One Tree Island) at the southern end of the Great Barrier Reef, Australia. A multi-disciplinary team assessed a variety of factors focusing on nutrient dynamics and biotic responses. A controlled and replicated experiment was conducted over two years using twelve small patch reefs ponded at low tide by a coral rim. Treatments included three control reefs (no nutrient addition) and three + N reefs (NH4Cl added), three + P reefs (KH2PO4 added), and three + N + P reefs. Nutrients were added as pulses at each low tide (ca twice per day) by remotely operated units. There were two phases of nutrient additions. During the initial, low-loading phase of the experiment nutrient pulses (mean dose = 11.5 muM NH4+; 2.3 muM PO4-3) rapidly declined, reaching near-background levels (mean = 0.9 muM NH4+; 0.5 muM PO4-3) within 2-3 h. A variety of biotic processes, assessed over a year during this initial nutrient loading phase, were not significantly affected, with the exception of coral reproduction, which was affected in all nutrient treatments. In Acropora longicyathus and A. aspera, fewer successfully developed embryos were formed, and in A. longicyathus fertilization rates and lipid levels decreased. In the second, high-loading, phase of ENCORE an increased nutrient dosage (mean dose = 36.2 muM NH4+; 5.1 muM PO4-3 declining to means of 11.3 muM NH4+ and 2.4 muM PO4-3 at the end of low tide) was used for a further year, and a variety of significant biotic responses occurred. Encrusting algae incorporated virtually none of the added nutrients. Organisms containing endosymbiotic zooxanthellae (corals and giant clams) assimilated dissolved nutrients rapidly and were responsive to added nutrients. Coral mortality, not detected during the initial low-loading phase, became evident with increased nutrient dosage, particularly in Pocillopora damicornis. Nitrogen additions stunted coral growth, and phosphorus additions had a variable effect. Coral calcification rate and linear extension increased in the presence of added phosphorus but skeletal density was reduced, making corals more susceptible to breakage. Settlement of all coral larvae was reduced in nitrogen treatments, yet settlement of larvae from brooded species was enhanced in phosphorus treatments. Recruitment of stomatopods, benthic crustaceans living in coral rubble, was reduced in nitrogen and nitrogen plus phosphorus treatments. Grazing rates and reproductive effort of various fish species were not affected by the nutrient treatments. Microbial nitrogen transformations in sediments,were responsive to nutrient loading with nitrogen fixation significantly increased in phosphorus treatments and denitrification increased in all treatments to which nitrogen had been added. Rates of bioerosion and grazing showed no significant effects of added nutrients, ENCORE has shown that reef organisms and processes investigated ill situ were impacted by elevated nutrients. Impacts mere dependent on dose level, whether nitrogen and/or phosphorus mere elevated and were often species-specific. The impacts were generally sub-lethal and subtle and the treated reefs at the end of the experiment mere visually similar to control reefs. Rapid nutrient uptake indicates that nutrient concentrations alone are not adequate to assess nutrient condition of reefs. Sensitive and quantifiable biological indicators need to be developed for coral reef ecosystems. The potential bioindicators identified in ENCORE should be tested in future research on coral reef/nutrient interactions. Synergistic and cumulative effects of elevated nutrients and other environmental parameters, comparative studies of intact vs. disturbed reefs, offshore vs, inshore reefs, or the ability of a nutrient-stressed reef to respond to natural disturbances require elucidation. An expanded understanding of coral reef responses to anthropogenic impacts is necessary, particularly regarding the subtle, sub-lethal effects detected in the ENCORE studies. (C) 2001 Published by Elsevier Science Ltd.

Seawater carbonate chemistry, nutrients and growth rate of coral (Acropora intermedia) and coral-reef seaweed (Lobophora papenfussii) during experiments, 2011

Space competition between corals and seaweeds is an important ecological process underlying coral-reef dynamics. Processes promoting seaweed growth and survival, such as herbivore overfishing and eutrophication, can lead to local reef degradation. Here, we present the case that increasing concentrations of atmospheric CO2 may be an additional process driving a shift from corals to seaweeds on reefs. Coral (Acropora intermedia) mortality in contact with a common coral-reef seaweed (Lobophora papenfussii) increased two- to threefold between background CO2 (400 ppm) and highest level projected for late 21st century (1140 ppm). The strong interaction between CO2 and seaweeds on coral mortality was most likely attributable to a chemical competitive mechanism, as control corals with algal mimics showed no mortality. Our results suggest that coral (Acropora) reefs may become increasingly susceptible to seaweed proliferation under ocean acidification, and processes regulating algal abundance (e.g. herbivory) will play an increasingly important role in maintaining coral abundance.

Cell death and degeneration in the symbiotic dinoflagellates of the coral Stylophora pistillata during bleaching

Rising sea temperatures are increasing the incidences of mass coral bleaching (the dissociation of the coral-algal symbiosis) and coral mortality. In this study, the effects of bleaching (induced by elevated light and temperature) on the condition of symbiotic dinoflagellates (Symbiodinium sp.) within the tissue of the hard coral Stylophora pistillata (Esper) were assessed using a suite of techniques. Bleaching of S. pistillata was accompanied by declines in the maximum potential quantum yield of photosynthesis (F-v/F-m, measured using pulse amplitude modulated [PAM] fluorometry), an increase in the number of Sytox-green-stained algae (indicating compromised algal membrane integrity and cell death), an increase in 2',7'-dichlorodihydrofluroscein diacetate (H(2)DCFDA)stained algae (indicating increased oxidative stress), as well as ultrastructural changes (vacuolisation, losses of chlorophyll, and an increase in accumulation bodies). Algae expelled from S. pistillata exhibited a complete disorganisation of cellular contents; expelled cells contained only amorphous material. In situ samples taken during a natural mass coral bleaching event on the Great Barrier Reef in February 2002 also revealed a high number of Sytox-labelled algae cells in symbio. Dinoflagellate degeneration during bleaching seems to be similar to the changes resulting from senescence-phase cell death in cultured algae. These data support a role for oxidative stress in the mechanism of coral bleaching and highlight the importance of algal degeneration during the bleaching of a reef coral.

Coral disease dynamics and environmental drivers in the northern Florida Keys and Lee Stocking Island, Bahamas

Coral reefs are experiencing declines worldwide and recently coral diseases have been identified as significant contributors to coral mortality. However, little is known regarding the factors that drive coral disease distributions and dynamics. Current knowledge of the organisms that cause coral diseases is also limited, with pathogens having been identified for only 5 of the 21 described coral diseases. The study presented here describes coral disease dynamics in terms of occurrence, prevalence, spatial distribution, and host species susceptibility from 2002--2004 on reefs of the Northern Florida Keys (NFK) and Lee Stocking Island (LSI) in the Bahamas' Exuma chain. In addition, this research investigated the influence of temperature, sediment, and nutrient availability on coral disease prevalence and severity. Finally, microbial communities associated with a polymicrobial disease, black band, were examined to address spatial and temporal variability. ^ Four scleractinian diseases were observed in repeated surveys conducted during June-August of each year: black band disease (BBD), white plague type 2 (WP), dark spots syndrome (DSS), and yellow band disease-(YBD). Coral disease prevalence was generally low in both the NFK and LSI as compared to epizootic levels reported previously in the NFK and other regions of the Caribbean. Disease prevalence and species susceptibility varied spatially and temporally. Massive framework species, including Siderastrea siderea, Colpophyllia natans, and Montastraea annularis, along with relatively smaller colonies of Meandrina meandrites and Dichocoenia stokesi, were most susceptible to disease. Temperature, sedimentation, and dissolved inorganic nitrogen were positively correlated with BBD infections. Furthermore, experimental nutrient enrichment exacerbated coral tissue loss to BBD both in situ and in vivo. Profiling of BBD microbial communities using length heterogeneity PCR revealed variation over space and time, with significantly distinct bacterial assemblages in the NFK, LSI, and US Virgin Islands. ^ This study contributes to knowledge of the relationship between coral diseases and the environment, and facilitates predictions regarding potential changes in coral reef communities under differing environmental conditions. Additionally, this research provides further understanding of coral disease dynamics at both the host and microbial pathogen levels.^

Diurnal and seasonal measurements of seawater chemistry, temperature and PAR in Hurricane Hole, U.S. Virgin Islands - 2010-2012

Risk analyses indicate that more than 90% of the world's reefs will be threatened by climate change and local anthropogenic impacts by the year 2030 under "business-as-usual" climate scenarios. Increasing temperatures and solar radiation cause coral bleaching that has resulted in extensive coral mortality. Increasing carbon dioxide reduces seawater pH, slows coral growth, and may cause loss of reef structure. Management strategies include establishment of marine protected areas with environmental conditions that promote reef resiliency. However, few resilient reefs have been identified, and resiliency factors are poorly defined. Here we characterize the first natural, non-reef coral refuge from thermal stress and ocean acidification and identify resiliency factors for mangrove-coral habitats. We measured diurnal and seasonal variations in temperature, salinity, photosynthetically active radiation (PAR), and seawater chemistry; characterized substrate parameters; and examined water circulation patterns in mangrove communities where scleractinian corals are growing attached to and under mangrove prop roots in Hurricane Hole, St. John, US Virgin Islands. Additionally, we inventoried the coral species and quantified incidences of coral bleaching, mortality, and recovery for two major reef-building corals, Colpophyllia natans and Diploria labyrinthiformis, growing in mangrove-shaded and exposed (unshaded) areas. Over 30 species of scleractinian corals were growing in association with mangroves. Corals were thriving in low-light (more than 70% attenuation of incident PAR) from mangrove shading and at higher temperatures than nearby reef tract corals. A higher percentage of C. natans colonies were living shaded by mangroves, and no shaded colonies were bleached. Fewer D. labyrinthiformis colonies were shaded by mangroves, however more unshaded colonies were bleached. A combination of substrate and habitat heterogeneity, proximity of different habitat types, hydrographic conditions, and biological influences on seawater chemistry generate chemical conditions that buffer against ocean acidification. This previously undocumented refuge for corals provides evidence for adaptation of coastal organisms and ecosystem transition due to recent climate change. Identifying and protecting other natural, non-reef coral refuges is critical for sustaining corals and other reef species into the future.

Increased mortality and photoinhibition in the symbiotic dinoflagellates of the Indo-Pacific coral Stylophora pistillata (Esper) after summer bleaching

Coral bleaching (the loss of symbiotic dinoflagellates from reef-building corals) is most frequently caused by high-light and temperature conditions. We exposed the explants of the hermatypic coral Stylophora pistillata to four combinations of light and temperature in late spring and also in late summer. During mid-summer, two NOAA bleaching warnings were issued for Heron Island reef (Southern Great Barrier Reef, Australia) when sea temperature exceeded the NOAA bleaching threshold, and a 'mild' (in terms of the whole coral community) bleaching event occurred, resulting in widespread S. pistillata bleaching and mortality. Symbiotic dinoflagellate biomass decreased by more than half from late spring to late summer (from 2.5x10(6) to 0.8x10(6) dinoflagellates cm(2) coral tissue), and those dinoflagellates that remained after summer became photoinhibited more readily (dark-adapted F (V) : F (M) decreased to (0.3 compared with 0.4 in spring), and died in greater numbers (up to 17% dinoflagellate mortality compared with 5% in the spring) when exposed to artificially elevated light and temperature. Adding exogenous antioxidants (D-mannitol and L-ascorbic acid) to the water surrounding the coral had no clear effect on either photoinhibition or symbiont mortality. These data show that light and temperature stress cause mortality of the dinoflagellate symbionts within the coral, and that susceptibility to light and temperature stress is strongly related to coral condition. Photoinhibitory mechanisms are clearly involved, and will increase through a positive feedback mechanism: symbiont loss promotes further symbiont loss as the light microenvironment becomes progressively harsher.

Mass mortality following disturbance in Holocene coral reefs from Papua New Guinea

The frequency and intensity of disturbance on living coral reefs have been accelerating for the past few decades, resulting in a changed seascape. What is unclear but vital for management is whether this acceleration is natural or coincident only with recent human impact. We surveyed nine uplifted early to mid-Holocene (11,000-3700 calendar [cal] yr B.P.) fringing and barrier reefs along similar to 27 km at the Huon Peninsula, Papua New Guinea. We found evidence for several episodes of coral mass mortality, but frequency was < 1 in 1500 yr. The most striking mortality event extends > 16 km along the ancient coastline, occurred ca. 9100-9400 cal yr B.P., and is associated with a volcanic ash horizon. Recolonization of the reef surface and resumption of vertical reef accretion was rapid (< 100 yr), but the post-disturbance reef communities contrasted with their pre-disturbance counterparts. Assessing the frequency, nature, and long-term ecological consequences of mass-mortality events in fossil coral reefs may provide important insights to guide management of modern reefs in this time of environmental degradation and change.

Colony-specific calcification and mortality under ocean acidification in the branching coral Montipora digitata

Ocean acidification (OA) threatens calcifying marine organisms including reef-building corals. In this study, we examined the OA responses of individual colonies of the branching scleractinian coral Montipora digitata. We exposed nubbins of unique colonies (n = 15) to ambient or elevated pCO2 under natural light and temperature regimes for 110 days. Although elevated pCO2 exposure on average reduced calcification, individual colonies showed unique responses ranging from declines in positive calcification to negative calcification (decalcification) to no change. Similarly, mortality was greater on average in elevated pCO2, but also showed colony-specific patterns. High variation in colony responses suggests the possibility that ongoing OA may lead to natural selection of OA-tolerant colonies within a coral population.

The effects of copper on the microbial community of a coral reef sponge

Marine sponges often harbour communities of symbiotic microorganisms that fulfil necessary functions for the well-being of their hosts. Microbial communities associated with the sponge Rhopaloeides odorabile were used as bioindicators far sublethal cupric ion (Cu2+) stress. A combined strategy incorporating molecular, cultivation and electron microscopy techniques was adopted to monitor changes in microbial diversity. The total density of sponge-associated bacteria and counts of the predominant cultivated symbiont (alpha -proteobacterium strain NW001) were significantly reduced in response to Cu2+ concentrations of 1.7 mug l(-1) and above after 14 days of exposure. The number of operational taxonomic units (OTUs) detected by restriction fragment length polymorphism (RFLP) decreased by 64% in sponges exposed to 223 mug l(-1) Cu2+ for 48 h and by 46% in sponges exposed to 19.4 mug l(-1) Cu2+ for 14 days. Electron microscopy was used to identify 17 predominant bacterial morphotypes, composing 47% of the total observed cells in control sponges. A reduction In the proportion of these morphotypes to 25% of observed cells was evident in sponges exposed to a Cu2+ concentration of 19.4 mug l(-1). Although the abundance of most morphotypes decreased under Cu2+ stress, three morphotypes were not reduced in numbers and a single morphotype actually increased in abundance. Bacterial numbers, as detected using fluorescence in situ hybridization (FISH), decreased significantly after 48 h exposure to 19.4 mug l(-1) Cu2+. Archaea, which are normally prolific in R. odorabile, were not detected after exposure to a Cu2+ concentration of 19.4 mug l(-1) for 14 days, indicating that many of the microorganisms associated with R. odorabile are sensitive to free copper. Sponges exposed to a Cu2+ concentration of 223 mug l(-1) became highly necrosed after 48 h and accumulated 142 +/- 18 mg kg(-1) copper, whereas sponges exposed to 19.4 mug l(-1) Cu2+ accumulated 306 +/- 15 mg kg(-1) copper after 14 days without apoptosis or mortality. Not only do sponges have potential for monitoring elevated concentrations of heavy metals but also examining changes in their microbial symbionts is a novel and sensitive bioindicator for the assessment of pollution on important microbial communities.

High-frequency climatic oscillations recorded in a Holocene coral reef at Leizhou Peninsula, South China sea

A detailed study of the Goniopora reef profile at Dengloujiao, Xuwen County, Leizhou Peninsula, the northern coast of the South China Sea suggests that a series of high-frequency, large-amplitude and abrupt cold events occurred during the Holocene Hypsithermal, an unusual phenomenon termed Leizhou Events in this paper. This period (corresponding to C-14 age of 6.2 -6.7 kaBP or calendar age of 6.7-7.2 kaBP), when the climatic conditions were ideal for coral. reefs to develop, can be divided into at least nine stages. Each stage (or called a climate optimum), lasting about 20 to 50 a, was terminated by an abrupt cold nap and (or) a sea-level lowering event in winter, leading to widespread emergence and death of the Goniopora corals, and growth discontinuities on the coral surface. Such a cyclic process resulted in the creation of a > 4m thick Goniopora reef flat. During this period, the crust subsided periodically but the sea level was rising. The reef profile provides valuable archives for the study of decadal-scale mid-Holocene climatic oscillations in the tropical area of South China. Our results provide new evidence for high-frequency climate instability in the Holocene Hypsithermal, and challenge the traditional understanding of Holocene climate.

The influence of increased line-fishing mortality on the sex ratio and age of sex reversal of the venus tusk fish

The age of sex reversal of the venus tusk fish Choerodon venustus, caught by line fishing at various locations on the southern Great Barrier Reef, indicated that C. venustus is capable of modifying its life cycle in response to increased mortality. The evidence suggests Masthead Reef fish, which experience the highest mortality, underwent sex reversal at a smaller size and younger age than at the other sites. The largest female fish, sexually transitional fish and males were smaller at Masthead Reef than at the Swains Reefs or One Tree Reef at Masthead Reef. There was also considerable overlap in the size of males and females within the exploited populations indicating that sex reversal is not initiated at a particular length but may have a social cause. The sex ratio of fish was essentially the same for fish fully susceptible to line fishing in the Swains and Masthead samples. Circumstantial evidence suggested that the absence of large males in a population may initiate sex reversal, indicating the maintenance of a constant sex ratio may have a social basis. (C) 2002 The Fisheries Society of the British Isles.