The impact of short-term exposure to near shore stressors on the early life stages of the reef building coral Montipora capitata.

Successful reproduction and survival are crucial to the continuation and resilience of corals globally. As reef waters warm due to climate change, episodic largescale tropical storms are becoming more frequent, drastically altering the near shore water quality for short periods of time. Therefore, it is critical that we understand the effects warming waters, fresh water input, and run-off have on sexual reproduction of coral. To better understand the effects of these near shore stressors on Hawaiian coral, laboratory experiments were conducted at the Institute of Marine Biology to determine the independent effects of suspended sediment concentrations (100 mg l-1 and 200 mg l-1), lowered salinity (28‰), and elevated temperature (31 °C) on the successful fertilization, larval survival, and settlement of the scleractinian coral Montipora capitata. In the present study, early developmental stages of coral were exposed to one of three near shore stressors for a period of 24 h and the immediate (fertilization) and latent effects (larval survival and settlement) were observed and measured. Fertilization success and settlement were not affected by any of the treatments; however, larval survival was negatively affected by all of the treatments by 50% or greater (p > 0.05). These data show that early life stages of M. capitata may be impacted by near shore stressors associated with warming and more frequent storm events.

Patterns of bleaching and mortality following widespread warming events in 2014 and 2015 at the Hanauma Bay Nature Preserve, Hawai'i.

Drastic increases in global carbon emissions in the past century have led to elevated sea surface temperatures that negatively affect coral reef organisms. Worldwide coral bleaching-related mortality is increasing and data has shown even isolated and protected reefs are vulnerable to the effects of global climate change. In 2014 and 2015, coral reefs in the main Hawaiian Islands (MHI) suffered up to 90% bleaching, with higher than 50% subsequent mortality in some areas. The location and severity of bleaching and mortality was strongly influenced by the spatial and temporal patterns of elevated seawater temperatures. The main objective of this research was to understand the spatial extent of bleaching mortality in Hanauma Bay Nature Preserve (HBNP), O'ahu, Hawai'i to gain a baseline understanding of the physical processes that influence localized bleaching dynamics. Surveys at HBNP in October 2015 and January 2016 revealed extensive bleaching (47%) and high levels of coral mortality (9.8%). Bleaching was highly variable among the four HBNP sectors and ranged from a low of ∼31% in the central bay at Channel (CH) to a high of 57% in the area most frequented by visitors (Keyhole; KH). The highest levels of bleaching occurred in two sectors with different circulation patterns: KH experienced comparatively low circulation velocity and a low temperature increase while Witches Brew (WB) and Backdoors (BD) experienced higher circulation velocity and higher temperature increase. Cumulative mortality was highest at WB (5.0%) and at BD (2.9%) although WB circulation velocity is significantly higher. HBNP is minimally impacted by local factors that can lead to decline such as high fishing pressure or sedimentation although human use is high. Despite the lack of these influences, high coral mortality occurred. Visitor impacts are strikingly different in the two sectors that experienced the highest mortality evidenced by the differences in coral cover associated with visitor use however, coral mortality was similar. These results suggest that elevated temperature was more influential in coral bleaching and the associated mortality than high circulation or visitor use.

Effectiveness of coral relocation as a mitigation strategy in Kane'ohe Bay, Hawai'i.

Coral reef restoration and management techniques are in ever-increasing demand due to the global decline of coral reefs in the last several decades. Coral relocation has been established as an appropriate restoration technique in select cases, particularly where corals are scheduled for destruction. However, continued long-term monitoring of recovery of transplanted corals is seldom sustained. Removal of coral from a navigation channel and relocation to a similar nearby dredged site occurred in 2005. Coral recovery at the donor site and changes in fish populations at the receiving site were tracked periodically over the following decade. Coral regrowth at the donor site was rapid until a recent bleaching event reduced coral cover by more than half. The transplant of mature colonies increased spatial complexity at the receiving site, immediately increasing fish biomass, abundance, and species that was maintained throughout subsequent surveys. Our research indicates that unlike the majority of historical accounts of coral relocation in the Pacific, corals transplanted into wave-protected areas with similar conditions as the original site can have high survival rates. Data on long-term monitoring of coral transplants in diverse environments is central in developing management and mitigation strategies.

The 2014 coral bleaching and freshwater flood events in Kane'ohe Bay, Hawai'i.

Until recently, subtropical Hawai'i escaped the major bleaching events that have devastated many tropical regions, but the continued increases in global long-term mean temperatures and the apparent ending of the Pacific Decadal Oscillation (PDO) cool phase have increased the risk of bleaching events. Climate models and observations predict that bleaching in Hawai'i will occur with increasing frequency and increasing severity over future decades. A freshwater "kill" event occurred during July 2014 in the northern part of Kane'ohe Bay that reduced coral cover by 22.5% in the area directly impacted by flooding. A subsequent major bleaching event during September 2014 caused extensive coral bleaching and mortality throughout the bay and further reduced coral cover in the freshwater kill area by 60.0%. The high temperature bleaching event only caused a 1.0% reduction in live coral throughout the portion of the bay not directly impacted by the freshwater event. Thus, the combined impact of the low salinity event and the thermal bleaching event appears to be more than simply additive. The temperature regime during the September 2014 bleaching event was analogous in duration and intensity to that of the large bleaching event that occurred previously during August 1996, but resulted in a much larger area of bleaching and coral mortality. Apparently seasonal timing as well as duration and magnitude of heating is important. Coral spawning in the dominant coral species occurs early in the summer, so reservoirs of stored lipid in the corals had been depleted by spawning prior to the September 2014 event. Warm months above 27 °C result in lower coral growth and presumably could further decrease lipid reserves, leading to a bleaching event that was more severe than would have happened if the high temperatures occurred earlier in the summer. Hawaiian reef corals decrease skeletal growth at temperatures above 27 °C, so perhaps the "stress period" actually started long before the bleaching threshold of 29 °C was reached. Hawai'i is directly influenced by the PDO which may become a factor influencing bleaching events in subtropical Hawai'i in much the same manner as variations in the El Niño Southern Oscillation (ENSO) influences bleaching events at low latitudes in the tropical Pacific. Records show that offshore temperatures measured by satellite will not always predict inshore bleaching because other factors (high cloud cover, high wind and wave action, tidal exchange rate) can limit inshore heating and prevent temperatures in the bay from reaching the bleaching threshold. Low light levels due to cloud cover or high turbidity can also serve to prevent bleaching.

The unnatural history of Kane'ohe Bay: coral reef resilience in the face of centuries of anthropogenic impacts.

Kane'ohe Bay, which is located on the on the NE coast of O'ahu, Hawai'i, represents one of the most intensively studied estuarine coral reef ecosystems in the world. Despite a long history of anthropogenic disturbance, from early settlement to post European contact, the coral reef ecosystem of Kane'ohe Bay appears to be in better condition in comparison to other reefs around the world. The island of Moku o Lo'e (Coconut Island) in the southern region of the bay became home to the Hawai'i Institute of Marine Biology in 1947, where researchers have since documented the various aspects of the unique physical, chemical, and biological features of this coral reef ecosystem. The first human contact by voyaging Polynesians occurred at least 700 years ago. By A.D. 1250 Polynesians voyagers had settled inhabitable islands in the region which led to development of an intensive agricultural, fish pond and ocean resource system that supported a large human population. Anthropogenic disturbance initially involved clearing of land for agriculture, intentional or accidental introduction of alien species, modification of streams to supply water for taro culture, and construction of massive shoreline fish pond enclosures and extensive terraces in the valleys that were used for taro culture. The arrival by the first Europeans in 1778 led to further introductions of plants and animals that radically changed the landscape. Subsequent development of a plantation agricultural system led to increased human immigration, population growth and an end to traditional land and water management practices. The reefs were devastated by extensive dredge and fill operations as well as rapid growth of human population, which led to extensive urbanization of the watershed. By the 1960's the bay was severely impacted by increased sewage discharge along with increased sedimentation due to improper grading practices and stream channelization, resulting in extensive loss of coral cover. The reefs of Kane'ohe Bay developed under estuarine conditions and thus have been subjected to multiple natural stresses. These include storm floods, a more extreme temperature range than more oceanic reefs, high rates of sedimentation, and exposure at extreme low tides. Deposition and degradation of organic materials carried into the bay from the watershed results in low pH conditions such that according to some ocean acidification projections the rich coral reefs in the bay should not exist. Increased global temperature due to anthropogenic fossil fuel emmisions is now impacting these reefs with the first "bleaching event" in 1996 and a second more severe event in 2014. The reefs of Kane'ohe Bay have developed and persist under rather severe natural and anthropogenic perturbations. To date, these reefs have proved to be very resilient once the stressor has been removed. A major question remains to be answered concerning the limits of Kane'ohe Bay reef resilience in the face of global climate change.

Comparison of methods used to estimate coral cover in the Hawaiian Islands.

Nine coral survey methods were compared at ten sites in various reef habitats with different levels of coral cover in Kane'ohe Bay, O'ahu, Hawai'i. Mean estimated coverage at the different sites ranged from less than 10% cover to greater than 90% cover. The methods evaluated include line transects, various visual and photographic belt transects, video transects and visual estimates. At each site 25 m transect lines were laid out and secured. Observers skilled in each method measured coral cover at each site. The time required to run each transect, time required to process data and time to record the results were documented. Cost of hardware and software for each method was also tabulated. Results of this investigation indicate that all of the methods used provide a good first estimate of coral cover on a reef. However, there were differences between the methods in detecting the number of coral species. For example, the classic "quadrat" method allows close examination of small and cryptic coral species that are not detected by other methods such as the "towboard" surveys. The time, effort and cost involved with each method varied widely, and the suitability of each method for answering particular research questions in various environments was evaluated. Results of this study support the finding of three other comparison method studies conducted at various geographic locations throughout the world. Thus, coral cover measured by different methods can be legitimately combined or compared in many situations. The success of a recent modeling effort based on coral cover data consisting of observations taken in Hawai'i using the different methods supports this conclusion.

Coral-algae metabolism and diurnal changes in the CO2-carbonate system of bulk sea water.

Precise measurements were conducted in continuous flow seawater mesocosms located in full sunlight that compared metabolic response of coral, coral-macroalgae and macroalgae systems over a diurnal cycle. Irradiance controlled net photosynthesis (P net), which in turn drove net calcification (G net), and altered pH. P net exerted the dominant control on [CO3 (2-)] and aragonite saturation state (Ωarag) over the diel cycle. Dark calcification rate decreased after sunset, reaching zero near midnight followed by an increasing rate that peaked at 03:00 h. Changes in Ωarag and pH lagged behind G net throughout the daily cycle by two or more hours. The flux rate P net was the primary driver of calcification. Daytime coral metabolism rapidly removes dissolved inorganic carbon (DIC) from the bulk seawater and photosynthesis provides the energy that drives G net while increasing the bulk water pH. These relationships result in a correlation between G net and Ωarag, with Ωarag as the dependent variable. High rates of H(+) efflux continued for several hours following mid-day peak G net suggesting that corals have difficulty in shedding waste protons as described by the Proton Flux Hypothesis. DIC flux (uptake) followed P net and G net and dropped off rapidly following peak P net and peak G net indicating that corals can cope more effectively with the problem of limited DIC supply compared to the problem of eliminating H(+). Over a 24 h period the plot of total alkalinity (AT ) versus DIC as well as the plot of G net versus Ωarag revealed a circular hysteresis pattern over the diel cycle in the coral and coral-algae mesocosms, but not the macroalgae mesocosm. Presence of macroalgae did not change G net of the corals, but altered the relationship between Ωarag and G net. Predictive models of how future global changes will effect coral growth that are based on oceanic Ωarag must include the influence of future localized P net on G net and changes in rate of reef carbonate dissolution. The correlation between Ωarag and G net over the diel cycle is simply the response of the CO2-carbonate system to increased pH as photosynthesis shifts the equilibria and increases the [CO3 (2-)] relative to the other DIC components of [HCO3 (-)] and [CO2]. Therefore Ωarag closely tracked pH as an effect of changes in P net, which also drove changes in G net. Measurements of DIC flux and H(+) flux are far more useful than concentrations in describing coral metabolism dynamics. Coral reefs are systems that exist in constant disequilibrium with the water column.

Effects of terrigenous sediment on settlement and survival of the reef coral Pocillopora damicornis.

Survival and settlement of Pocillopora damicornis larvae on hard surfaces covered with fine (<63 µm) terrigenous red clay was measured in laboratory Petri dishes. The dishes were prepared with sediment films of various thicknesses covering the bottoms. Coral larvae were incubated in the dishes for two weeks and the percent that settled on the bottom was determined. There was a statistically significant relationship between the amount of sediment and coral recruitment on the bottom, with no recruitment on surfaces having a sediment cover above 0.9 mg cm(-2). Experimental conditions for the delicate coral larvae were favorable in these experiments. Total survival over the two week settlement tests expressed as the sum of coral recruits and live larvae at the end of the experiment did not show a significant decline, so the major impact of the sediment was on successful settlement rather than on mortality. Larval substrate selection behavior was the primary factor in the observed result.

Alkalinity to calcium flux ratios for corals and coral reef communities: variances between isolated and community conditions.

Calcification in reef corals and coral reefs is widely measured using the alkalinity depletion method which is based on the fact that two protons are produced for every mole of CaCO3 precipitated. This assumption was tested by measuring the total alkalinity (TA) flux and Ca(2+) flux of isolated components (corals, alga, sediment and plankton) in reference to that of a mixed-community. Experiments were conducted in a flume under natural conditions of sunlight, nutrients, plankton and organic matter. A realistic hydrodynamic regime was provided. Groups of corals were run separately and in conjunction with the other reef components in a mixed-community. The TA flux to Ca(2+) flux ratio (ΔTA: ΔCa(2+)) was consistently higher in the coral-only run (2.06 ± 0.19) than in the mixed-community run (1.60 ± 0.14, p-value = 0.011). The pH was higher and more stable in the mixed-community run (7.94 ± 0.03 vs. 7.52 ± 0.07, p-value = 3 × 10(-5)). Aragonite saturation state (Ω arag) was also higher in the mixed-community run (2.51 ± 0.2 vs. 1.12 ± 0.14, p-value = 2 × 10(-6)). The sediment-only run revealed that sediment is the source of TA that can account for the lower ΔTA: ΔCa(2+) ratio in the mixed-community run. The macroalgae-only run showed that algae were responsible for the increased pH in the mixed-community run. Corals growing in a mixed-community will experience an environment that is more favorable to calcification (higher daytime pH due to algae photosynthesis, additional TA and inorganic carbon from sediments, higher Ω arag). A paradox is that the alkalinity depletion method will yield a lower net calcification for a mixed-community versus a coral-only community due to TA recycling, even though the corals may be calcifying at a higher rate due to a more optimal environment.

Thirty years of coral reef change in relation to coastal construction and increased sedimentation at Pelekane Bay, Hawai'i.

Coral reefs are being critically impacted by anthropogenic processes throughout the world. Long term monitoring is essential to the understanding of coral reef response to human impacts and the effectiveness of corrective management efforts. Here we reevaluated a valuable coral reef baseline established in Pelekane Bay, Hawai'i during 1976 and subsequently resurveyed in 1996. During this time interval substantial impacts occurred followed by extensive corrective measures. Coral and fish communities showed dramatic declines from 1977 to 1996 due to massive harbor construction and suboptimal land management practices on the watershed. More recently, corrective measures in the form of watershed stabilization and fishing regulations have been implemented. Consequently our 2012 survey reveals that coral cover since 1996 has increased slightly accompanied by a significant increase in fish abundance, diversity, and evenness. This improvement can be attributed to lower fishing pressure since 1996 due to reduced shoreline access, tighter fishing regulations and increased monitoring of legal and illegal fishing activities. Stabilization of the coral community can be attributed partially to reduced sedimentation resulting from watershed restoration that included installation of sediment check dams, control of feral ungulates, controlled grazing and replanting of native vegetation. Insights into the mechanism that removes sediment from reefs was provided by a major storm event and a tsunami that remobilized and flushed out sediment deposits. The increase in herbivorous fishes probably played a role in reducing algal competition in favor of corals. The data suggest that the precipitous reef decline in this area has been arrested and offers support for the corrective actions previously undertaken.

Response of reef corals on a fringing reef flat to elevated suspended-sediment concentrations: Moloka'i, Hawai'i.

A long-term (10 month exposure) experiment on effects of suspended sediment on the mortality, growth, and recruitment of the reef corals Montipora capitata and Porites compressa was conducted on the shallow reef flat off south Moloka'i, Hawai'i. Corals were grown on wire platforms with attached coral recruitment tiles along a suspended solid concentration (SSC) gradient that ranged from 37 mg l(-1) (inshore) to 3 mg l(-1) (offshore). Natural coral reef development on the reef flat is limited to areas with SSCs less than 10 mg l(-1) as previously suggested in the scientific literature. However, the experimental corals held at much higher levels of turbidity showed surprisingly good survivorship and growth. High SSCs encountered on the reef flat reduced coral recruitment by one to three orders of magnitude compared to other sites throughout Hawai'i. There was a significant correlation between the biomass of macroalgae attached to the wire growth platforms at the end of the experiment and percentage of the corals showing mortality. We conclude that lack of suitable hard substrate, macroalgal competition, and blockage of recruitment on available substratum are major factors accounting for the low natural coral coverage in areas of high turbidity. The direct impact of high turbidity on growth and mortality is of lesser importance.

Use of integrated landscape indicators to evaluate the health of linked watersheds and coral reef environments in the Hawaiian islands.

A linkage between the condition of watersheds and adjacent nearshore coral reef communities is an assumed paradigm in the concept of integrated coastal management. However, quantitative evidence for this "catchment to sea" or "ridge to reef" relationship on oceanic islands is lacking and would benefit from the use of appropriate marine and terrestrial landscape indicators to quantify and evaluate ecological status on a large spatial scale. To address this need, our study compared the Hawai'i Watershed Health Index (HI-WHI) and Reef Health Index (HI-RHI) derived independently of each other over the past decade. Comparisons were made across 170 coral reef stations at 52 reef sites adjacent to 42 watersheds throughout the main Hawaiian Islands. A significant positive relationship was shown between the health of watersheds and that of adjacent reef environments when all sites and depths were considered. This relationship was strongest for sites facing in a southerly direction, but diminished for north facing coasts exposed to persistent high surf. High surf conditions along the north shore increase local wave driven currents and flush watershed-derived materials away from nearshore waters. Consequently, reefs in these locales are less vulnerable to the deposition of land derived sediments, nutrients and pollutants transported from watersheds to ocean. Use of integrated landscape health indices can be applied to improve regional-scale conservation and resource management.

Projected changes to growth and mortality of Hawaiian corals over the next 100 years.

BACKGROUND: Recent reviews suggest that the warming and acidification of ocean surface waters predicated by most accepted climate projections will lead to mass mortality and declining calcification rates of reef-building corals. This study investigates the use of modeling techniques to quantitatively examine rates of coral cover change due to these effects. METHODOLOGY/PRINCIPAL FINDINGS: Broad-scale probabilities of change in shallow-water scleractinian coral cover in the Hawaiian Archipelago for years 2000-2099 A.D. were calculated assuming a single middle-of-the-road greenhouse gas emissions scenario. These projections were based on ensemble calculations of a growth and mortality model that used sea surface temperature (SST), atmospheric carbon dioxide (CO(2)), observed coral growth (calcification) rates, and observed mortality linked to mass coral bleaching episodes as inputs. SST and CO(2) predictions were derived from the World Climate Research Programme (WCRP) multi-model dataset, statistically downscaled with historical data. CONCLUSIONS/SIGNIFICANCE: The model calculations illustrate a practical approach to systematic evaluation of climate change effects on corals, and also show the effect of uncertainties in current climate predictions and in coral adaptation capabilities on estimated changes in coral cover. Despite these large uncertainties, this analysis quantitatively illustrates that a large decline in coral cover is highly likely in the 21(st) Century, but that there are significant spatial and temporal variances in outcomes, even under a single climate change scenario.