An enigmatic decoupling between heat stress and coral bleaching on the Great Barrier Reef

Ocean warming threatens the functioning of coral reef ecosystems by inducing mass coral bleaching and mortality events. The link between temperature and coral bleaching is now well-established based on observations that mass bleaching events usually occur when seawater temperatures are anomalously high. However, times of high heat stress but without coral bleaching are equally important because they can inform an understanding of factors that regulate temperature-induced bleaching. Here, we investigate the absence of mass coral bleaching on the Great Barrier Reef (GBR) during austral summer 2004. Using four gridded sea surface temperature data products, validated with in situ temperature loggers, we demonstrate that the summer of 2004 was among the warmest summers of the satellite era (1982–2017) on the GBR. At least half of the GBR experienced temperatures that were high enough to initiate bleaching in other years, yet mass bleaching was not reported during 2004. The absence of bleaching is not fully explained by wind speed or cloud cover. Rather, 2004 is clearly differentiated from bleaching years by the slow speed of the East Australian Current (EAC) offshore of the GBR. An anomalously slow EAC during summer 2004 may have dampened the upwelling of nutrient-rich waters onto the GBR shelf, potentially mitigating bleaching due to the lower susceptibility of corals to heat stress in low-nutrient conditions. Although other factors such as irradiance or acclimatization may have played a role in the absence of mass bleaching, 2004 remains a key case study for demonstrating the dynamic nature of coral responses to marine heatwaves

Coral biomineralization, climate proxies and the sensitivity of coral reefs to CO2-driven climate change

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution February 2017Scleractinian corals extract calcium (Ca2+) and carbonate (CO2−3) ions from seawater to construct their calcium carbonate (CaCO3) skeletons. Key to the coral biomineralization process is the active elevation of the CO2−3 concentration of the calcifying fluid to achieve rapid nucleation and growth of CaCO3 crystals. Coral skeletons contain valuable records of past climate variability and contribute to the formation of coral reefs. However, limitations in our understanding of coral biomineralization hinder the accuracy of (1) coral-based reconstructions of past climate, and (2) predictions of coral reef futures as anthropogenic CO2 emissions drive declines in seawater CO2−3 concentration. In this thesis, I investigate the mechanism of coral biomineralization and evaluate the sensitivity of coral reef CaCO3 production to seawater carbonate chemistry. First, I conducted abiogenic CaCO3 precipitation experiments that identified the U/Ca ratio as a proxy for fluid CO2−3 concentration. Based on these experimental results, I developed a quantitative coral biomineralization model that predicts temperature can be reconstructed from coral skeletons by combining Sr/Ca - which is sensitive to both temperature and CO2−3 - with U/Ca into a new proxy called “Sr-U”. I tested this prediction with 14 corals from the Pacific Ocean and the Red Sea spanning mean annual temperatures of 25.7-30.1°C and found that Sr-U has uncertainty of only 0.5°C, twice as accurate as conventional coral-based thermometers. Second, I investigated the processes that differentiate reef-water and open-ocean carbonate chemistry, and the sensitivity of ecosystem-scale calcification to these changes. On Dongsha Atoll in the northern South China Sea, metabolic activity of resident organisms elevates reef-water CO2−3 twice as high as the surrounding open ocean, driving rates of ecosystem calcification higher than any other coral reef studied to date. When high temperatures stressed the resident coral community, metabolic activity slowed, with dramatic effects on reef-water chemistry and ecosystem calcification. Overall, my thesis highlights how the modulation of CO2−3, by benthic communities on the reef and individual coral polyps in the colony, controls the sensitivity of coral reefs to future ocean acidification and influences the climate records contained in the skeleton.This research was funded by a National Science Foundation (NSF) Graduate Research Fellowship, NSF grants OCE 1041106, OCE 1338320, and OCE 1220529, by a thematic project at Academia Sinica, Taiwan, the WHOI Ocean Ventures Fund, and by the WHOI Coastal Ocean Institute

Factors affecting B/Ca ratios in synthetic aragonite

Author Posting. © The Author(s), 2015. This is the author's version of the work. It is posted here for personal use, not for redistribution. The definitive version was published in Chemical Geology 437 (2016): 67-76, doi:10.1016/j.chemgeo.2016.05.007.Measurements of B/Ca ratios in marine carbonates have been suggested to record seawater carbonate chemistry, however experimental calibration of such proxies based on inorganic partitioning remains limited. Here we conducted a series of synthetic aragonite precipitation experiments to evaluate the factors influencing the partitioning of B/Ca between aragonite and seawater. Our results indicate that the B/Ca ratio of synthetic aragonites depends primarily on the relative concentrations of borate and carbonate ions in the solution from which the aragonite precipitates; not on bicarbonate concentration as has been previously suggested. The influence of temperature was not significant over the range investigated (20 – 40°C), however, partitioning may be influenced by saturation state (and/or growth rate). Based on our experimental results, we suggest that aragonite B/Ca ratios can be utilized as a proxy of [CO32-]. Boron isotopic composition (δ11B) is an established pH proxy, thus B/Ca and δ11B together allow the full carbonate chemistry of the solution from which the aragonite precipitated to be calculated. To the extent that aragonite precipitation by marine organisms is affected by seawater chemistry, B/Ca may also prove useful in reconstructing seawater chemistry. A simplified boron purification protocol based on amberlite resin and the organic buffer TRIS is also described.This work was supported by the Australian Research Council (ARC) Centre of Excellence for Coral Reef Studies. Research conducted at WHOI was supported by NSF grant OCE-1338320. M.H. was supported by an ARC Super Science Fellowship and an NSF International Postdoctoral Fellowship. T.D. was supported by a NSF Graduate Research Fellowship. M.M. was supported by a Western Australian Premiers Fellowship and an ARC Laureate Fellowship

Coral reef drag coefficients – water depth dependence

Author Posting. © American Meteorological Society, 2017. This article is posted here by permission of American Meteorological Society for personal use, not for redistribution. The definitive version was published in Journal of Physical Oceanography 47 (2017): 1061-1075, doi:10.1175/JPO-D-16-0248.1.A major challenge in modeling the circulation over coral reefs is uncertainty in the drag coefficient because existing estimates span two orders of magnitude. Current and pressure measurements from five coral reefs are used to estimate drag coefficients based on depth-average flow, assuming a balance between the cross-reef pressure gradient and the bottom stress. At two sites wind stress is a significant term in the cross-reef momentum balance and is included in estimating the drag coefficient. For the five coral reef sites and a previous laboratory study, estimated drag coefficients increase as the water depth decreases consistent with open channel flow theory. For example, for a typical coral reef hydrodynamic roughness of 5 cm, observational estimates, and the theory indicate that the drag coefficient decreases from 0.4 in 20 cm of water to 0.005 in 10 m of water. Synthesis of results from the new field observations with estimates from previous field and laboratory studies indicate that coral reef drag coefficients range from 0.2 to 0.005 and hydrodynamic roughnesses generally range from 2 to 8 cm. While coral reef drag coefficients depend on factors such as physical roughness and surface waves, a substantial fraction of the scatter in estimates of coral reef drag coefficients is due to variations in water depth.The Red Sea field program was supported by Awards USA 00002 and KSA 00011 made by KAUST to S. Lentz and J. Churchill. The Palau field program was funded by NSF Award OCE-1220529

Relationship between water and aragonite barium concentrations in aquaria reared juvenile corals

This paper is not subject to U.S. copyright. The definitive version was published in Geochimica et Cosmochimica Acta 209 (2017): 123-134, doi:10.1016/j.gca.2017.04.006.Coral barium to calcium (Ba/Ca) ratios have been used to reconstruct records of upwelling, river and groundwater discharge, and sediment and dust input to the coastal ocean. However, this proxy has not yet been explicitly tested to determine if Ba inclusion in the coral skeleton is directly proportional to seawater Ba concentration and to further determine how additional factors such as temperature and calcification rate control coral Ba/Ca ratios. We measured the inclusion of Ba within aquaria reared juvenile corals (Favia fragum) at three temperatures (∼27.7, 24.6 and 22.5 °C) and three seawater Ba concentrations (73, 230 and 450 nmol kg−1). Coral polyps were settled on tiles conditioned with encrusting coralline algae, which complicated chemical analysis of the coral skeletal material grown during the aquaria experiments. We utilized Sr/Ca ratios of encrusting coralline algae (as low as 3.4 mmol mol−1) to correct coral Ba/Ca for this contamination, which was determined to be 26 ± 11% using a two end member mixing model. Notably, there was a large range in Ba/Ca across all treatments, however, we found that Ba inclusion was linear across the full concentration range. The temperature sensitivity of the distribution coefficient is within the range of previously reported values. Finally, calcification rate, which displayed large variability, was not correlated to the distribution coefficient. The observed temperature dependence predicts a change in coral Ba/Ca ratios of 1.1 μmol mol−1 from 20 to 28 °C for typical coastal ocean Ba concentrations of 50 nmol kg−1. Given the linear uptake of Ba by corals observed in this study, coral proxy records that demonstrate peaks of 10–25 μmol mol−1 would require coastal seawater Ba of between 60 and 145 nmol kg−1. Further validation of the coral Ba/Ca proxy requires evaluation of changes in seawater chemistry associated with the environmental perturbation recorded by the coral as well as verification of these results for Porites species, which are widely used in paleo reconstructions.M.E.G. was supported by a NDSEG graduate fellowship. Funding for this research came from the NSF Chemical Oceanography program (OCE-0751525) and the Coastal Ocean Institute, the Ocean and Climate Change Institute and the Ocean Ventures Fund at Woods Hole Oceanographic Institution

Climate modulates internal wave activity in the Northern South China Sea

Author Posting. © American Geophysical Union, 2015. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Geophysical Research Letters 42 (2015): 831–838, doi:10.1002/2014GL062522.Internal waves (IWs) generated in the Luzon Strait propagate into the Northern South China Sea (NSCS), enhancing biological productivity and affecting coral reefs by modulating nutrient concentrations and temperature. Here we use a state-of-the-art ocean data assimilation system to reconstruct water column stratification in the Luzon Strait as a proxy for IW activity in the NSCS and diagnose mechanisms for its variability. Interannual variability of stratification is driven by intrusions of the Kuroshio Current into the Luzon Strait and freshwater fluxes associated with the El Niño–Southern Oscillation. Warming in the upper 100 m of the ocean caused a trend of increasing IW activity since 1900, consistent with global climate model experiments that show stratification in the Luzon Strait increases in response to radiative forcing. IW activity is expected to increase in the NSCS through the 21st century, with implications for mitigating climate change impacts on coastal ecosystems.This work was supported by NSF award 1220529 to Anne Cohen, by the Academia Sinica (Taiwan) through a thematic project grant to G.T.F.W. and Anne Cohen, by the Alfred P. Sloan Foundation and the WHOI Oceans and Climate Change Institute/Moltz Fellowship through awards to K.B.K., and by an NSF Graduate Research Fellowship to T.M.D.2015-08-1

Mineralogy of Deep-Sea Coral Aragonites as a Function of Aragonite Saturation State

In an ocean with rapidly changing chemistry, studies have assessed coral skeletal health under projected ocean acidification (OA) scenarios by characterizing morphological distortions in skeletal architecture and measuring bulk properties, such as net calcification and dissolution. Few studies offer more detailed information on skeletal mineralogy. Since aragonite crystallography will at least partially govern the material properties of coral skeletons, such as solubility and strength, it is important to understand how it is influenced by environmental stressors. Here, we take a mineralogical approach using micro X-ray diffraction (XRD) and whole pattern Rietveld refinement analysis to track crystallographic shifts in deep-sea coral Lophelia pertusa samples collected along a natural seawater aragonite saturation state gradient (Ωsw = 1.15–1.44) in the Gulf of Mexico. Our results reveal statistically significant linear relationships between rising Ωsw and increasing unit cell volume driven by an anisotropic lengthening along the b-axis. These structural changes are similarly observed in synthetic aragonites precipitated under various saturation states, indicating that these changes are inherent to the crystallography of aragonite. Increased crystallographic disorder via widening of the full width at half maximum of the main (111) XRD peaks trend with increased Ba substitutions for Ca, however, trace substitutions by Ba, Sr, and Mg do not trend with crystal lattice parameters in our samples. Instead, we observe a significant trend of increasing calcite content as a function of both decreasing unit cell parameters as well as decreasing Ωsw. This may make calcite incorporation an important factor to consider in coral crystallography, especially under varying aragonite saturation states (ΩAr). Finally, by defining crystallography-based linear relationships between ΩAr of synthetic aragonite analogs and lattice parameters, we predict internal calcifying fluid saturation state (Ωcf = 11.1–17.3 calculated from b-axis lengths; 15.2–25.2 calculated from unit cell volumes) for L. pertusa, which may allow this species to calcify despite the local seawater conditions. This study will ideally pave the way for future studies to utilize quantitative XRD in exploring the impact of physical and chemical stressors on biominerals

Comparison of equatorial Pacific sea surface temperature variability and trends with Sr/Ca records from multiple corals

Author Posting. © American Geophysical Union, 2016. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Paleoceanography 31 (2016): 252–265, doi:10.1002/2015PA002897.Coral Sr/Ca is widely used to reconstruct past ocean temperatures. However, some studies report different Sr/Ca-temperature relationships for conspecifics on the same reef, with profound implications for interpretation of reconstructed temperatures. We assess whether these differences are attributable to small-scale oceanographic variability or “vital effects” associated with coral calcification and quantify the effect of intercolony differences on temperature estimates and uncertainties. Sr/Ca records from four massive Porites colonies growing on the east and west sides of Jarvis Island, central equatorial Pacific, were compared with in situ logger temperatures spanning 2002–2012. In general, Sr/Ca captured the occurrence of interannual sea surface temperature events but their amplitude was not consistently recorded by any of the corals. No long-term trend was identified in the instrumental data, yet Sr/Ca of one coral implied a statistically significant cooling trend while that of its neighbor implied a warming trend. Slopes of Sr/Ca-temperature regressions from the four different colonies were within error, but offsets in mean Sr/Ca rendered the regressions statistically distinct. Assuming that these relationships represent the full range of Sr/Ca-temperature calibrations in Jarvis Porites, we assessed how well Sr/Ca of a nonliving coral with an unknown Sr/Ca-temperature relationship can constrain past temperatures. Our results indicate that standard error of prediction methods underestimate the actual error as we could not reliably reconstruct the amplitude or frequency of El Niño–Southern Oscillation events as large as ± 2°C. Our results underscore the importance of characterizing the full range of temperature-Sr/Ca relationships at each study site to estimate true error.This study was supported by an NSF Graduate Research Fellowship to A.A. and by NSF-OCE-0926986 and NSF-OCE-1031971.2016-08-0

Validation of the remotely sensed nighttime sea surface temperature in the shallow waters at the Dongsha Atoll

© The Author(s), 2017. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Terrestrial, Atmospheric and Oceanic Sciences 28 (2017): 517-524, doi:10.3319/TAO.2017.03.30.01.Fine scale temperature structures, which are commonly found in the top few meters of shallow water columns, may result in deviations of the remotely sensed night-time sea surface temperatures (SST) by the MODIS-Aqua sensor (SSTsat) from the bulk sea surface temperatures (SSTbulk) that they purport to represent. The discrepancies between SSTsat and SSTbulk recorded by temperature loggers at eight stations with bottom depths of 2 - 20 m around the Dongsha Atoll (DSA) between June 2013 and May 2015 were examined. The SSTsat had an average cool bias error of -0.43 ± 0.59°C. The bias error was larger in the warmer (> 26°C) waters which were presumably more strongly stratified. The root mean square error (RMSE) between SSTsat and SSTbulk, ±0.73°C, was 25% larger than that reported in the open northern South China Sea. An operational calibration algorithm was developed to increase the accuracy in the estimation of SSTbulk from SSTsat. In addition to removing the cool bias error, this algorithm also reduced the RMSE to virtually the same level as that found in the open northern South China Sea. With the application of the algorithm, in June 2015, the average SST in the lagoon of the DSA was raised by about 0.5°C to 31.1 ± 0.4°C, and the area of lagoon with SSTbulk above 31°C, the median value of the physiological temperature threshold of reef organisms, was increased by 69% to about three quarters of the lagoon.This work was supported in part by the Key Research and Development Program of Shandong Province (grant no. 2015GSF117017) and Ocean University of China (grant no. 201513037 and 201512011) to Pan, and the Academia Sinica through grant titled “Ocean Acidification: Comparative biogeochemistry in shallow-water tropical coral reef ecosystems in a naturally acidic marine environment” to Wong

Physical processes determine spatial structure in water temperature and residence time on a wide reef flat

Author Posting. © American Geophysical Union, 2020. This article is posted here by permission of American Geophysical Union for personal use, not for redistribution. The definitive version was published in Journal of Geophysical Research: Oceans 125(12), (2020): e2020JC016543, https://doi.org/10.1029/2020JC016543.On coral reefs, flow determines residence time of water influencing physical and chemical environments and creating observable microclimates within the reef structure. Understanding the physical mechanisms driving environmental variability on shallow reefs, which distinguishes them from the open ocean, is important for understanding what contributes to thermal resilience of coral communities and predicting their response to future anomalies. In June 2014, a field experiment conducted at Dongsha Atoll in the northern South China Sea investigated the physical forces that drive flow over a broad shallow reef flat. Instrumentation included current and pressure sensors and a distributed temperature sensing system, which resolved spatially and temporally continuous temperature measurements over a 3‐km cross‐reef section from the lagoon to reef crest. Spectral analysis shows that while diurnal variability was significant across the reef flat—a result expected from daily solar heating—temperature also varied at higher frequencies near the reef crest. These spatially variable temperature regimes, or thermal microclimates, are influenced by circulation on the wide reef flat, with spatially and temporally variable contributions from tides, wind, and waves. Through particle tracking simulations, we find the residence time of water is shorter near the reef crest (3.6 h) than near the lagoon (8.6 h). Tidal variability in flow direction on the reef flat leads to patterns in residence time that are different than what would be predicted from unidirectional flow. Circulation on the reef also determines the source (originating from offshore vs. the lagoon) of the water present on the reef flat.We thank S. Tyler, and J. Selker from the Center for Transformative Environmental Monitoring Programs (CTEMPs), funded by the National Science Foundation (EAR awards 1440596 and 1440506), for timely and effective provision of experimental design support, logistical support and equipment for the project. Support for S. Lentz is from NSF Grant No. OCE‐1558343. Support for A. Cohen from NSF Grant No. 1220529, by the Academia Sinica (Taiwan) through a thematic project grant to G. Wong and A. Cohen. Support for E. Reid and K. Davis is from National Science Foundation (NSF) Grant No. OCE‐1753317, and support to E. Reid from the Environmental Engineering Henry Samueli Endowed Fellowship and the UCI Oceans Graduate Fellowship.2021-05-2