Impacts of water quality on Acropora coral settlement: The relative importance of substrate quality and light.

Coral larval settlement patterns are influenced by a vast array of factors; however, the relative roles of individual factors are rarely tested in isolation, leading to confusion about which are most crucial for settlement. For example, direct effects of the light environment are often cited as a major factor influencing settlement patterns, yet this has not been demonstrated under environmentally realistic lighting regimes in the absence of confounding factors. Here we apply programmable multispectral lights to create realistic light spectra, while removing correlating (but not obvious) factors that are common in laboratory settlement experiments. Using two common species of Acropora - key framework builders of the Great Barrier Reef - we find little evidence that light intensity or changes in the spectral profile play a substantial role in larval settlement under most environmentally realistic settings but can under more extreme or artificial settings. We alternatively hypothesise and provide evidence that chronic light conditions and recent sediment exposures that impact benthic substrates (e.g., crustose coralline algae) have a greater impact on settlement success. Under these conditions, there was a decrease of up to 74% settlement success. Management of water quality conditions that impact the quality of benthic-settlement substrates therefore should present a priority area of focus for improving coral recruitment.

The effects of ultraviolet radiation and climate on oil toxicity to coral reef organisms - A review.

Oil pollution remains a significant local threat to shallow tropical coral reef environments, but the environmental conditions typical of coral reefs are rarely considered in oil toxicity testing and risk assessments. Here we review the effects of three environmental co-factors on petroleum oil toxicity towards coral reef organisms, and show that the impacts of oil pollution on coral reef taxa can be exacerbated by environmental conditions commonly encountered in tropical reef environments. Shallow reefs are routinely exposed to high levels of ultraviolet radiation (UVR), which can substantially increase the toxicity of some oil components through phototoxicity. Exposure to UVR represents the most likely and harmful environmental co-factor reviewed here, leading to an average toxicity increase of 7.2-fold across all tests reviewed. The clear relevance of UVR co-exposure and its strong influence on tropical reef oil toxicity highlights the need to account for UVR as a standard practice in future oil toxicity studies. Indeed, quantifying the influence of UVR on toxic thresholds of oil to coral reef species is essential to develop credible oil spill risk models required for oil extraction developments, shipping management and spill responses in the tropics. The few studies available indicate that co-exposure to elevated temperature and low pH, both within the range of current daily and seasonal fluctuations and/or projected under continued climate change, can increase oil toxicity on average by 3.0- and 1.3-fold, respectively. While all three of the reviewed environmental co-factors have the potential to substantially increase the impacts of oil pollution in shallow reef environments, their simultaneous effects have not been investigated. Assessments of the combined effects of oil pollution, UVR, temperature and low pH will become increasingly important to identify realistic hazard thresholds suitable for future risk assessments over the coming century.

Sediment characteristics influence the fertilisation success of the corals Acropora tenuis and Acropora millepora.

Elevated suspended sediment concentrations (SSCs) often impact coral fertilisation success, but sediment composition can influence effect thresholds, which is problematic for accurately predicting risk. Here, we derived concentration-response thresholds and cause-effect pathways for SSCs comprising a range of realistic mineral and organic compositions on coral fertilisation success. Effect concentration thresholds (EC10: 10% fertilisation inhibition) varied markedly, with fertilisation highly sensitive to inshore organic-clay rich sediments and bentonite clay at <5 mg L-1. Mineral clays and organic matter within these sediments likely promoted flocculation of the coral sperm, which in turn reduced fertilisation. In contrast, sediments lacking these properties bound less sperm, leading to higher SSC thresholds for coral fertilisation (EC10 > 40 mg L-1). The effect thresholds for relevant sediment types were combined with in situ turbidity data from locations near dredging operations to assess the risks posed by dredging to coral fertilisation at these locations.

Settlement patterns of the coral Acropora millepora on sediment-laden surfaces.

Successful recruitment in corals is important for the sustenance of coral reefs, and is considered a demographic bottleneck in the recovery of reef populations following disturbance events. Yet several factors influence larval settlement behaviour, and here we quantified thresholds associated with light attenuation and accumulated sediments on settlement substrates. Sediments deposited on calcareous red algae (CRA) directly and indirectly impacted coral settlement patterns. Although not avoiding direct contact, Acropora millepora larvae were very reluctant to settle on surfaces layered with sediments, progressively shifting their settlement preference from upward to downward facing (sediment-free) surfaces under increasing levels of deposited sediment. When only upward-facing surfaces were presented, 10% of settlement was inhibited at thresholds from 0.9 to 16mgcm-2 (EC10), regardless of sediment type (carbonate and siliciclastic) or particle size (fine and coarse silt). These levels equate to a very thin (<150µm) veneer of sediment that occurs within background levels on reefs. Grooves within settlement surfaces slightly improved options for settlement on sediment-coated surfaces (EC10: 29mgcm-2), but were quickly infilled at higher deposited sediment levels. CRA that was temporarily smothered by sediment for 6d became bleached (53% surface area), and inhibited settlement at ~7mgcm-2 (EC10). A minor decrease in settlement was observed at high and very low light intensities when using suboptimal concentrations of a settlement inducer (CRA extract); however, no inhibition was observed when natural CRA surfaces along with more realistic diel-light patterns were applied. The low deposited sediment thresholds indicate that even a thin veneer of sediment can have consequences for larval settlement due to a reduction of optimal substrate. And while grooves and overhangs provide more settlement options in high deposition areas, recruits settling at these locations may be subject to ongoing stress from shading, competition, and sediment infilling.

Mucous Secretion and Cilia Beating Defend Developing Coral Larvae from Suspended Sediments.

Suspended sediments produced from dredging activities, or added to the sediment budget via river runoff, are a concern for marine resource managers. Understanding the impact of suspended sediments on critical life history stages of keystone species like corals is fundamental to effective management of coastlines and reefs. Coral embryos (Acropora tenuis and A. millepora) and larvae (A. tenuis, A. millepora and Pocillopora acuta) were subjected to a range of suspended sediment concentrations of different sediment types (siliciclastic and carbonate) to assess concentration-response relationships on ecologically relevant endpoints, including survivorship and ability to metamorphose. Embryos were subjected to short (12 h) suspended sediment exposures from ages of 3-12 hours old or a long (30 h) exposure at 6 hours old. Neither the survivorship nor metamorphosis function of embryos were significantly affected by realistic sediment exposures to ~1000 mg L-1. However, some embryos exhibited a previously undescribed response to dynamically suspended sediments, which saw 10% of the embryos form negatively buoyant cocoons at siliciclastic suspended sediment concentrations ≥35 mg L-1. Scanning electron and optical microscopy confirmed the presence of a coating on these embryos, possibly mucus with incorporated sediment particles. Cocoon formation was common in embryos but not in larvae, and occurred more often after exposure to siliciclastic rather than carbonate sediments. Once transferred into sediment-free seawater, functional ~36-h-old embryos began emerging from the cocoons, coinciding with cilia development. Ciliated (> 36-h-old) larvae exposed to suspended sediments for 60 h were also observed to secrete mucus and were similarly unaffected by suspended sediment concentrations to ~800 mg L-1. This study provides evidence that mucous secretion and cilia beating effectively protect coral embryos and larvae from suspended sediment and that these mechanisms may enhance their chances of successful recruitment.

Long distance dispersal and vertical gene flow in the Caribbean brooding coral Porites astreoides.

To date, most assessments of coral connectivity have emphasized long-distance horizontal dispersal of propagules from one shallow reef to another. The extent of vertical connectivity, however, remains largely understudied. Here, we used newly-developed and existing DNA microsatellite loci for the brooding coral Porites astreoides to assess patterns of horizontal and vertical connectivity in 590 colonies collected from three depth zones (≤10 m, 15-20 m and ≥25 m) at sites in Florida, Bermuda and the U.S. Virgin Islands (USVI). We also tested whether maternal transmission of algal symbionts (Symbiodinium spp.) might limit effective vertical connectivity. Overall, shallow P. astreoides exhibited high gene flow between Florida and USVI, but limited gene flow between these locations and Bermuda. In contrast, there was significant genetic differentiation by depth in Florida (Upper Keys, Lower Keys and Dry Tortugas), but not in Bermuda or USVI, despite strong patterns of depth zonation in algal symbionts at two of these locations. Together, these findings suggest that P. astreoides is effective at dispersing both horizontally and vertically despite its brooding reproductive mode and maternal transmission of algal symbionts. In addition, these findings might help explain the ecological success reported for P. astreoides in the Caribbean in recent decades.

Acute ecotoxicology of natural oil and gas condensate to coral reef larvae.

Risks posed by oil spills to coral reefs are difficult to evaluate, partially due to the absence of studies that adequately assess toxicity to relevant coral reef species. Here we experimentally tested the acute toxicity of condensate, representing a fraction of light crude oil, to coral (Acropora tenuis) and sponge (Rhopaloeides odorabile) larvae. The metamorphosis of coral larvae was inhibited at total petroleum aromatic hydrocarbon (TPAH) concentrations of water accommodated fractions (WAF) as low as 103 µg l(-1), similar to concentrations detected in seawater following large spills. The sensitivity of coral larvae increased by 40% when co-exposed to UV light that they might encounter in shallow reefal systems. Condensate WAF was more toxic to coral larvae than predicted by summing the toxicity of its main components (benzene, toluene, p-xylene and napthalene). In contrast, the sensitivity of sponge larvae to condensate WAF (>10,000 µg l(-1) TPAH) was far less than coral in the presence and absence of UV, but similar to that of other marine invertebrates. While these results highlight the relative sensitivity of coral larvae to oil, further research is needed to better understand and predict the impacts and risks posed by hydrocarbons to tropical reef systems.

That sinking feeling: Suspended sediments can prevent the ascent of coral egg bundles.

Spawning synchrony represents a common reproductive strategy in sessile marine organisms and for broadcast spawning corals, buoyancy of egg-sperm bundles is critical to maximise fertilisation at the ocean surface. Here we demonstrate a novel threat to coral reproduction whereby buoyant egg-sperm bundles intercept and are "ballasted" by sediment grains on their journey to the ocean surface, preventing them from reaching the ocean surface and greatly reducing egg-sperm encounter rates. Empirical observations of this mechanism are successfully captured by a mathematical model that predicts the reduction in ascent probability and egg-sperm encounters as a function of sediment load. When applied to 15 m deep reefs, the model predicts that 10% and 50% reductions in egg-sperm encounters occur at 35 mg L(-1) and 87 mg L(-1) suspended sediment concentrations, respectively, and for a 5 m deep reef a 10% reduction occurs at 106 mg L(-1). These concentrations are commonly associated with sediment plumes from dredging or natural resuspension events. The potential for sediments to sink coral gametes highlights the need to carefully manage the timing of turbidity-generating human activities near reefs during spawning periods.

Suspended sediments limit coral sperm availability.

Suspended sediment from dredging activities and natural resuspension events represent a risk to the reproductive processes of coral, and therefore the ongoing maintenance of reefal populations. To investigate the underlying mechanisms that could reduce the fertilisation success in turbid water, we conducted several experiments exposing gametes of the corals Acropora tenuis and A. millepora to two sediment types. Sperm limitation was identified in the presence of siliciclastic sediment (230 and ~700 mg L(-1)), with 2-37 fold more sperm required to achieve maximum fertilisation rates, when compared with sediment-free treatments. This effect was more pronounced at sub-optimum sperm concentrations. Considerable (>45%) decreases in sperm concentration at the water's surface was recorded in the presence of siliciclastic sediment and a >20% decrease for carbonate sediment. Electron microscopy then confirmed sediment entangled sperm and we propose entrapment and sinking is the primary mechanism reducing sperm available to the egg. Longer exposure to suspended sediments and gamete aging further decreased fertilisation success when compared with a shorter exposure. Collectively, these findings demonstrate that high concentrations of suspended sediments effectively remove sperm from the water's surface during coral spawning events, reducing the window for fertilisation with potential subsequent flow-on effects for recruitment.

Spatial patterns of chemical contamination (metals, PAHs, PCBs, PCDDs/PCDFS) in sediments of a non-industrialized but densely populated coral atoll/small island state (Bermuda).

There is a recognized dearth of standard environmental quality data in the wider Caribbean area, especially on coral atolls/small island states. Extensive surveys of sediment contamination (n=109 samples) in Bermuda revealed a wide spectrum of environmental quality. Zinc and especially copper levels were elevated at some locations, associated with boating (antifouling paints and boatyard discharges). Mercury contamination was surprisingly prevalent, with total levels as high as 12mg kg(-1)DW, although methyl mercury levels were quite low. PAH, PCB and PCDD/PCDF contamination was detected a several hotspots associated with road run-off, a marine landfill, and a former US Naval annexe. NOAA sediment quality guidelines were exceeded at several locations, indicating biological effects are possible, or at some locations probable. Overall, and despite lack of industrialization, anthropogenic chemicals in sediments of the atoll presented a risk to benthic biodiversity at a number of hotspots suggesting a need for sediment management strategies.

Caribbean corals in crisis: record thermal stress, bleaching, and mortality in 2005.

BACKGROUND: The rising temperature of the world's oceans has become a major threat to coral reefs globally as the severity and frequency of mass coral bleaching and mortality events increase. In 2005, high ocean temperatures in the tropical Atlantic and Caribbean resulted in the most severe bleaching event ever recorded in the basin. METHODOLOGY/PRINCIPAL FINDINGS: Satellite-based tools provided warnings for coral reef managers and scientists, guiding both the timing and location of researchers' field observations as anomalously warm conditions developed and spread across the greater Caribbean region from June to October 2005. Field surveys of bleaching and mortality exceeded prior efforts in detail and extent, and provided a new standard for documenting the effects of bleaching and for testing nowcast and forecast products. Collaborators from 22 countries undertook the most comprehensive documentation of basin-scale bleaching to date and found that over 80% of corals bleached and over 40% died at many sites. The most severe bleaching coincided with waters nearest a western Atlantic warm pool that was centered off the northern end of the Lesser Antilles. CONCLUSIONS/SIGNIFICANCE: Thermal stress during the 2005 event exceeded any observed from the Caribbean in the prior 20 years, and regionally-averaged temperatures were the warmest in over 150 years. Comparison of satellite data against field surveys demonstrated a significant predictive relationship between accumulated heat stress (measured using NOAA Coral Reef Watch's Degree Heating Weeks) and bleaching intensity. This severe, widespread bleaching and mortality will undoubtedly have long-term consequences for reef ecosystems and suggests a troubled future for tropical marine ecosystems under a warming climate.

Alterations in dimethylsulfoniopropionate (DMSP) levels in the coral Montastraea franksi in response to copper exposure.

Symbiotic corals routinely experience hyperoxic conditions within their tissues due to the photosynthesis of the endosymbiotic dinoflagellate microalgae (Symbiodinium spp.). Symbiodinium spp. produce high intracellular levels of the osmolyte dimethylsulfoniopropionate (DMSP). It has recently been discovered in marine algae that DMSP and its enzymatic breakdown products also play a significant role in the scavenging of cellular reactive oxygen species (ROS). To examine this potential for DMSP in corals, we exposed the hard coral Montastraea franksi to 1, 10 and 50 microg L(-1) (ppb) concentrations of the oxidative stressor, copper. Levels of total (DMSP(t), all coral tissue) were higher than particulate DMSP(p) (algal component only), demonstrating partitioning of DMSP between algal symbionts and coral host. Significant changes in levels of DMSP(t) and DMSP(p) occurred in M. franksi after 48 h, demonstrating a response to copper and indicating a potential antioxidant role for DMSP. DMSP(t) and DMSP(p) levels decreased with copper dose, although at the highest copper dose DMSP(p) levels increased, whereas DMSP(t) levels did not. This observed differential response to copper between DMSP(t) and DMSP(p) demonstrates that physiological changes may be overlooked if conclusions are based upon DMSP(t) levels alone, which is a common measure used in coral studies. Decreases in chlorophyll a and algal cell numbers in response to elevated copper were also observed. These indices are important physiological indicators and are often used as indices to normalize DMSP levels. Our data suggest that the use of these common indices for normalizing DMSP may not always be appropriate.

Chemical contamination of a coral reef by the grounding of a cruise ship in Bermuda.

Bulk metal analyses of surficial sediments collected around the Norwegian Crown cruise ship grounding site in Bermuda indicated significant but localized contamination of reef sediments by copper and zinc, caused by the stripping of the tri-butyltin (TBT)-free antifouling (AF) paint (Intersmooth 460) from the underside of the hull. Highest copper and zinc values were found in heavily compacted and red-pigmented sediments inside the impact scar and were comparable to levels found close to slip ways of local boat yards where AF paints from hull stripping and cleaning processes are washed into the sea. The re-distribution of AF contaminated sediments by storms and deposition on nearby reefs constitutes a significant ecological risk that could delay recovery processes and reduce the effectiveness of remediation efforts. Whilst the ecotoxicological effects of AF paint particles interspersed with sediment is unknown, and in need of further study, it is argued that the significance of AF paint contamination of grounding sites has been overlooked.

Communication arising. Is coral bleaching really adaptive?

From an experiment in which corals are transplanted between two depths on a Panamanian coral reef, Baker infers that bleaching may sometimes help reef corals to survive environmental change. Although Baker's results hint at further mechanisms by which reef-building corals may acclimatize to changing light conditions, we do not consider that the evidence supports his inference.