Seasonal nearshore ocean acidification and deoxygenation in the Southern California Bight.

The California Current System experiences seasonal ocean acidification and hypoxia (OAH) owing to wind-driven upwelling, but little is known about the intensity, frequency, and depth distribution of OAH in the shallow nearshore environment. Here we present observations of OAH and dissolved inorganic carbon and nutrient parameters based on monthly transects from March 2017 to September 2018 extending from the surf zone to the ~ 40 m depth contour in La Jolla, California. Biologically concerning OAH conditions were observed at depths as shallow as 10 m and as close as 700 m to the shoreline. Below 20 m depth, 8% of observations were undersaturated with respect to aragonite, 28% of observations had a pHT less than 7.85, and 19% of observations were below the sublethal oxygen threshold of 157 µmol kg-1. These observations raise important questions about the impacts of OAH on coastal organisms and ecosystems and how future intensified upwelling may exacerbate these conditions.

The past, present, and future of coral reef growth in the Florida Keys.

Coral-reef degradation is driving global-scale reductions in reef-building capacity and the ecological, geological, and socioeconomic functions it supports. The persistence of those essential functions will depend on whether coral-reef management is able to rebalance the competing processes of reef accretion and erosion. Here, we reconstructed census-based carbonate budgets of 46 reefs throughout the Florida Keys from 1996 to 2019. We evaluated the environmental and ecological drivers of changing budget states and compared historical trends in reef-accretion potential to millennial-scale baselines of accretion from reef cores and future projections with coral restoration. We found that historically, most reefs had positive carbonate budgets, and many had reef-accretion potential comparable to the ~3 mm year-1 average accretion rate during the peak of regional reef building ~7000 years ago; however, declines in reef-building Acropora palmata and Orbicella spp. corals following a series of thermal stress events and coral disease outbreaks resulted in a shift from positive to negative budgets for most reefs in the region. By 2019, only ~15% of reefs had positive net carbonate production. Most of those reefs were in inshore, Lower Keys patch-reef habitats with low water clarity, supporting the hypothesis that environments with naturally low irradiance may provide a refugia from thermal stress. We caution that our estimated carbonate budgets are likely overly optimistic; comparison of reef-accretion potential to measured accretion from reef cores suggests that, by not accounting for the role of nonbiological physical and chemical erosion, census-based carbonate budgets may underestimate total erosion by ~1 mm year-1 (-1.15 kg CaCO3 m-2 year-1 ). Although the present state of Florida's reefs is dire, we demonstrate that the restoration of reef-building corals has the potential to help mitigate declines in reef accretion in some locations, which could allow some key ecosystem functions to be maintained until the threat of global climate change is addressed.

Scale dependence of coral reef oases and their environmental correlates.

Identifying relatively intact areas within ecosystems and determining the conditions favoring their existence is necessary for effective management in the context of widespread environmental degradation. In this study, we used 3766 surveys of randomly selected sites in the United States and U.S. Territories to identify the correlates of sites categorized as "oases" (defined as sites with relatively high total coral cover). We used occupancy models to evaluate the influence of 10 environmental predictors on the probability that an area (21.2-km2 cell) would harbor coral oases defined at four spatial extents: cross-basin, basin, region, and subregion. Across all four spatial extents, oases were more likely to occur in habitats with high light attenuation. The influence of the other environmental predictors on the probability of oasis occurrence were less consistent and varied with the scale of observation. Oases were most likely in areas of low human population density, but this effect was evident only at the cross-basin and subregional extents. At the regional and subregional extents oases were more likely where sea-surface temperature was more variable, whereas at the larger spatial extents the opposite was true. By identifying the correlates of oasis occurrence, the model can inform the prioritization of reef areas for management. Areas with biophysical conditions that confer corals with physiological resilience, as well as limited human impacts, likely support coral reef oases across spatial extents. Our approach is widely applicable to the development of conservation strategies to protect biodiversity and ecosystems in an era of magnified human disturbance.

Implications of salinity normalization of seawater total alkalinity in coral reef metabolism studies.

Salinity normalization of total alkalinity (TA) and dissolved inorganic carbon (DIC) data is commonly used to account for conservative mixing processes when inferring net metabolic modification of seawater by coral reefs. Salinity (S), TA, and DIC can be accurately and precisely measured, but salinity normalization of TA (nTA) and DIC (nDIC) can generate considerable and unrecognized uncertainties in coral reef metabolic rate estimates. While salinity normalization errors apply to nTA, nDIC, and other ions of interest in coral reefs, here, we focus on nTA due to its application as a proxy for net coral reef calcification and the importance for reefs to maintain calcium carbonate production under environmental change. We used global datasets of coral reef TA, S, and modeled groundwater discharge to assess the effect of different volumetric ratios of multiple freshwater TA inputs (i.e., groundwater, river, surface runoff, and precipitation) on nTA. Coral reef freshwater endmember TA ranged from -2 up to 3032 µmol/kg in hypothetical reef locations with freshwater inputs dominated by riverine, surface runoff, or precipitation mixing with groundwater. The upper bound of freshwater TA in these scenarios can result in an uncertainty in reef TA of up to 90 µmol/kg per unit S normalization if the freshwater endmember is erroneously assumed to have 0 µmol/kg alkalinity. The uncertainty associated with S normalization can, under some circumstances, even shift the interpretation of whether reefs are net calcifying to net dissolving, or vice versa. Moreover, the choice of reference salinity for normalization implicitly makes assumptions about whether biogeochemical processes occur before or after mixing between different water masses, which can add uncertainties of ±1.4% nTA per unit S normalization. Additional considerations in identifying potential freshwater sources of TA and their relative volumetric impact on seawater are required to reduce uncertainties associated with S normalization of coral reef carbonate chemistry data in some environments. However, at a minimum, researchers should minimize the range of salinities over which the normalization is applied, precisely measure salinity, and normalize TA values to a carefully selected reference salinity that takes local factors into account.

Coral calcification responses to the North Atlantic Oscillation and coral bleaching in Bermuda.

The North Atlantic Oscillation (NAO) has been hypothesized to drive interannual variability in Bermudan coral extension rates and reef-scale calcification through the provisioning of nutritional pulses associated with negative NAO winters. However, the direct influence of the NAO on Bermudan coral calcification rates remains to be determined and may vary between species and reef sites owing to implicit differences in coral life history strategies and environmental gradients across the Bermuda reef platform. In this study, we investigated the connection between negative NAO winters and Bermudan Diploria labyrinthiformis, Pseudodiploria strigosa, and Orbicella franksi coral calcification rates across rim reef, lagoon, and nearshore reef sites. Linear mixed effects modeling detected an inverse correlation between D. labyrinthiformis calcification rates and the winter NAO index, with higher rates associated with increasingly negative NAO winters. Conversely, there were no detectable correlations between P. strigosa or O. franksi calcification rates and the winter NAO index suggesting that coral calcification responses associated with negative NAO winters could be species-specific. The correlation between coral calcification rates and winter NAO index was significantly more negative at the outer rim of the reef (Hog Reef) compared to a nearshore reef site (Whalebone Bay), possibly indicating differential influence of the NAO as a function of the distance from the reef edge. Furthermore, a negative calcification anomaly was observed in 100% of D. labyrinthiformis cores in association with the 1988 coral bleaching event with a subsequent positive calcification anomaly in 1989 indicating a post-bleaching recovery in calcification rates. These results highlight the importance of assessing variable interannual coral calcification responses between species and across inshore-offshore gradients to interannual atmospheric modes such as the NAO, thermal stress events, and potential interactions between ocean warming and availability of coral nutrition to improve projections for future coral calcification rates under climate change.

Nearshore coral growth declining on the Mesoamerican Barrier Reef System.

Anthropogenic global change and local stressors are impacting coral growth and survival worldwide, altering the structure and function of coral reef ecosystems. Here, we show that skeletal extension rates of nearshore colonies of two abundant and widespread Caribbean corals (Siderastrea siderea, Pseudodiploria strigosa) declined across the Belize Mesoamerican Barrier Reef System (MBRS) over the past century, while offshore coral conspecifics exhibited relatively stable extension rates over the same temporal interval. This decline has caused nearshore coral extension rates to converge with those of their historically slower growing offshore coral counterparts. For both species, individual mass coral bleaching events were correlated with low rates of skeletal extension within specific reef environments, but no single bleaching event was correlated with low skeletal extension rates across all reef environments. We postulate that the decline in skeletal extension rates for nearshore corals is driven primarily by the combined effects of long-term ocean warming and increasing exposure to higher levels of land-based anthropogenic stressors, with acute thermally induced bleaching events playing a lesser role. If these declining trends in skeletal growth of nearshore S. siderea and P. strigosa continue into the future, the structure and function of these critical nearshore MBRS coral reef systems is likely to be severely impaired.

Environmental controls on modern scleractinian coral and reef-scale calcification.

Modern reef-building corals sustain a wide range of ecosystem services because of their ability to build calcium carbonate reef systems. The influence of environmental variables on coral calcification rates has been extensively studied, but our understanding of their relative importance is limited by the absence of in situ observations and the ability to decouple the interactions between different properties. We show that temperature is the primary driver of coral colony (Porites astreoides and Diploria labyrinthiformis) and reef-scale calcification rates over a 2-year monitoring period from the Bermuda coral reef. On the basis of multimodel climate simulations (Coupled Model Intercomparison Project Phase 5) and assuming sufficient coral nutrition, our results suggest that P. astreoides and D. labyrinthiformis coral calcification rates in Bermuda could increase throughout the 21st century as a result of gradual warming predicted under a minimum CO2 emissions pathway [representative concentration pathway (RCP) 2.6] with positive 21st-century calcification rates potentially maintained under a reduced CO2 emissions pathway (RCP 4.5). These results highlight the potential benefits of rapid reductions in global anthropogenic CO2 emissions for 21st-century Bermuda coral reefs and the ecosystem services they provide.

Heterotrophy mitigates the response of the temperate coral Oculina arbuscula to temperature stress.

Anthropogenic increases in atmospheric carbon dioxide concentration have caused global average sea surface temperature (SST) to increase by approximately 0.11°C per decade between 1971 and 2010 - a trend that is projected to continue through the 21st century. A multitude of research studies have demonstrated that increased SSTs compromise the coral holobiont (cnidarian host and its symbiotic algae) by reducing both host calcification and symbiont density, among other variables. However, we still do not fully understand the role of heterotrophy in the response of the coral holobiont to elevated temperature, particularly for temperate corals. Here, we conducted a pair of independent experiments to investigate the influence of heterotrophy on the response of the temperate scleractinian coral Oculina arbuscula to thermal stress. Colonies of O. arbuscula from Radio Island, North Carolina, were exposed to four feeding treatments (zero, low, moderate, and high concentrations of newly hatched Artemia sp. nauplii) across two independent temperature experiments (average annual SST (20°C) and average summer temperature (28°C) for the interval 2005-2012) to quantify the effects of heterotrophy on coral skeletal growth and symbiont density. Results suggest that heterotrophy mitigated both reduced skeletal growth and decreased symbiont density observed for unfed corals reared at 28°C. This study highlights the importance of heterotrophy in maintaining coral holobiont fitness under thermal stress and has important implications for the interpretation of coral response to climate change.

Temperature Regimes Impact Coral Assemblages along Environmental Gradients on Lagoonal Reefs in Belize.

Coral reefs are increasingly threatened by global and local anthropogenic stressors such as rising seawater temperature, nutrient enrichment, sedimentation, and overfishing. Although many studies have investigated the impacts of local and global stressors on coral reefs, we still do not fully understand how these stressors influence coral community structure, particularly across environmental gradients on a reef system. Here, we investigate coral community composition across three different temperature and productivity regimes along a nearshore-offshore gradient on lagoonal reefs of the Belize Mesoamerican Barrier Reef System (MBRS). A novel metric was developed using ultra-high-resolution satellite-derived estimates of sea surface temperatures (SST) to classify reefs as exposed to low (lowTP), moderate (modTP), or high (highTP) temperature parameters over 10 years (2003 to 2012). Coral species richness, abundance, diversity, density, and percent cover were lower at highTP sites relative to lowTP and modTP sites, but these coral community traits did not differ significantly between lowTP and modTP sites. Analysis of coral life history strategies revealed that highTP sites were dominated by hardy stress-tolerant and fast-growing weedy coral species, while lowTP and modTP sites consisted of competitive, generalist, weedy, and stress-tolerant coral species. Satellite-derived estimates of Chlorophyll-a (chl-a) were obtained for 13-years (2003-2015) as a proxy for primary production. Chl-a concentrations were highest at highTP sites, medial at modTP sites, and lowest at lowTP sites. Notably, thermal parameters correlated better with coral community traits between site types than productivity, suggesting that temperature (specifically number of days above the thermal bleaching threshold) played a greater role in defining coral community structure than productivity on the MBRS. Dominance of weedy and stress-tolerant genera at highTP sites suggests that corals utilizing these two life history strategies may be better suited to cope with warmer oceans and thus may warrant protective status under climate change.