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2018
Takeshita, Y, Cyronak T, Martz TR, Kindeberg T, Andersson AJ.  2018.  Coral reef carbonate chemistry variability at different functional scales. Frontiers in Marine Science. 5   10.3389/fmars.2018.00175   AbstractWebsite

There is a growing recognition for the need to understand how seawater carbonate chemistry over coral reef environments will change in a high-CO2 world to better assess the impacts of ocean acidification on these valuable ecosystems. Coral reefs modify overlying water column chemistry through biogeochemical processes such as net community organic carbon production (NCR) and calcification (NCC). However, the relative importance and influence of these processes on seawater carbonate chemistry vary across multiple functional scales (defined here as space, time, and benthic community composition), and have not been fully constrained. Here, we use Bermuda as a case study to assess (1) spatiotemporal variability in physical and chemical parameters along a depth gradient at a rim reef location, (2) the spatial variability of total alkalinity (TA) and dissolved inorganic carbon (DIC) over distinct benthic habitats to infer NCC:NCP ratios [< several km(2); rim reef vs. seagrass and calcium carbonate (CaCO3) sediments] on diel timescales, and (3) compare how TA-DIC relationships and NCC:NCP vary as we expand functional scales from local habitats to the entire reef platform (10's of km(2)) on seasonal to interannual timescales. Our results demonstrate that TA-DIC relationships were strongly driven by local benthic metabolism and community composition over diel cycles. However, as the spatial scale expanded to the reef platform, the TA-DIC relationship reflected processes that were integrated over larger spatiotemporal scales, with effects of NCC becoming increasingly more important over NCR. This study demonstrates the importance of considering drivers across multiple functional scales to constrain carbonate chemistry variability over coral reefs.

Eyre, BD, Cyronak T, Drupp P, DeCarlo EH, Sachs JP, Andersson AJ.  2018.  Coral reefs will transition to net dissolving before end of century. Science. 359:908-911.   10.1126/science.aao1118   AbstractWebsite

Ocean acidification refers to the lowering of the ocean's pH due to the uptake of anthropogenic CO2 from the atmosphere. Coral reef calcification is expected to decrease as the oceans become more acidic. Dissolving calciumcarbonate (CaCO3) sands could greatly exacerbate reef loss associated with reduced calcification but is presently poorly constrained. Here we show that CaCO3 dissolution in reef sediments across five globally distributed sites is negatively correlated with the aragonite saturation state (War) of overlying seawater and that CaCO3 sediment dissolution is 10-fold more sensitive to ocean acidification than coral calcification. Consequently, reef sediments globally will transition from net precipitation to net dissolution when seawater War reaches 2.92 +/- 0.16 (expected circa 2050 CE). Notably, some reefs are already experiencing net sediment dissolution.

Cyronak, T, Andersson AJ, Langdon C, Albright R, Bates NR, Caldeira K, Carlton R, Corredor JE, Dunbar RB, Enochs I, Erez J, Eyre BD, Gattuso JP, Gledhill D, Kayanne H, Kline DI, Koweek DA, Lantz C, Lazar B, Manzello D, McMahon A, Melendez M, Page HN, Santos IR, Schulz KG, Shaw E, Silverman J, Suzuki A, Teneva L, Watanabe A, Yamamoto S.  2018.  Taking the metabolic pulse of the world's coral reefs. Plos One. 13   10.1371/journal.pone.0190872   AbstractWebsite

Worldwide, coral reef ecosystems are experiencing increasing pressure from a variety of anthropogenic perturbations including ocean warming and acidification, increased sedimentation, eutrophication, and overfishing, which could shift reefs to a condition of net calcium carbonate (CaCO3) dissolution and erosion. Herein, we determine the net calcification potential and the relative balance of net organic carbon metabolism (net community production; NCP) and net inorganic carbon metabolism (net community calcification; NCC) within 23 coral reef locations across the globe. In light of these results, we consider the suitability of using these two metrics developed from total alkalinity (TA) and dissolved inorganic carbon (DIC) measurements collected on different spatiotemporal scales to monitor coral reef biogeochemistry under anthropogenic change. All reefs in this study were net calcifying for the majority of observations as inferred from alkalinity depletion relative to offshore, although occasional observations of net dissolution occurred at most locations. However, reefs with lower net calcification potential (i.e., lower TA depletion) could shift towards net dissolution sooner than reefs with a higher potential. The percent influence of organic carbon fluxes on total changes in dissolved inorganic carbon (DIC) (i.e., NCP compared to the sum of NCP and NCC) ranged from 32% to 88% and reflected inherent biogeochemical differences between reefs. Reefs with the largest relative percentage of NCP experienced the largest variability in seawater pH for a given change in DIC, which is directly related to the reefs ability to elevate or suppress local pH relative to the open ocean. This work highlights the value of measuring coral reef carbonate chemistry when evaluating their susceptibility to ongoing global environmental change and offers a baseline from which to guide future conservation efforts aimed at preserving these valuable ecosystems.

2015
Yeakel, KL, Andersson AJ, Bates NR, Noyes TJ, Collins A, Garley R.  2015.  Shifts in coral reef biogeochemistry and resulting acidification linked to offshore productivity. Proceedings of the National Academy of Sciences of the United States of America. 112:14512-14517.   10.1073/pnas.1507021112   AbstractWebsite

Oceanic uptake of anthropogenic carbon dioxide (CO2) has acidified open-ocean surface waters by 0.1 pH units since preindustrial times. Despite unequivocal evidence of ocean acidification (OA) via open-ocean measurements for the past several decades, it has yet to be documented in near-shore and coral reef environments. A lack of long-term measurements from these environments restricts our understanding of the natural variability and controls of seawater CO2-carbonate chemistry and biogeochemistry, which is essential to make accurate predictions on the effects of future OA on coral reefs. Here, in a 5-y study of the Bermuda coral reef, we show evidence that variations in reef biogeochemical processes drive interannual changes in seawater pH and Omega(aragonite) that are partly controlled by offshore processes. Rapid acidification events driven by shifts toward increasing net calcification and net heterotrophy were observed during the summers of 2010 and 2011, with the frequency and extent of such events corresponding to increased offshore productivity. These events also coincided with a negative winter North Atlantic Oscillation (NAO) index, which historically has been associated with extensive offshore mixing and greater primary productivity at the Bermuda Atlantic Time-series Study (BATS) site. Our results reveal that coral reefs undergo natural interannual events of rapid acidification due to shifts in reef biogeochemical processes that may be linked to offshore productivity and ultimately controlled by larger-scale climatic and oceanographic processes.

2014
Andersson, AJ, Yeakel KL, Bates NR, de Putron SJ.  2014.  Partial offsets in ocean acidification from changing coral reef biogeochemistry. Nature Climate Change. 4:56-61.   10.1038/nclimate2050   AbstractWebsite

Concerns have been raised about how coral reefs will be affected by ocean acidification(1,2), but projections of future seawater CO2 chemistry have focused solely on changes in the pH and aragonite saturation state (Omega(a)) of open-ocean surface seawater conditions surrounding coral reefs(1-4) rather than the reef systems themselves. The seawater CO2 chemistry within heterogeneous reef systems can be significantly different from that of the open ocean depending on the residence time, community composition and the main biogeochemical processes occurring on the reef, that is, net ecosystem production (NEP = gross primary production autotrophic and heterotrophic respiration) and net ecosystem calcification (NEC = gross calcification gross CaCO3 dissolution), which combined act to modify seawater chemistry(5-7). On the basis of observations from the Bermuda coral reef, we show that a range of projected biogeochemical responses of coral reef communities to ocean acidification by the end of this century could partially offset changes in seawater pH and Omega(a) by an average of 12-24% and 15-31%, respectively.

2013
McLeod, E, Anthony KRN, Andersson A, Beeden R, Golbuu Y, Kleypas J, Kroeker K, Manzello D, Salm RV, Schuttenberg H, Smith JE.  2013.  Preparing to manage coral reefs for ocean acidification: lessons from coral bleaching. Frontiers in Ecology and the Environment. 11:20-27.   10.1890/110240   AbstractWebsite

Ocean acidification is a direct consequence of increasing atmospheric carbon dioxide concentrations and is expected to compromise the structure and function of coral reefs within this century. Research into the effects of ocean acidification on coral reefs has focused primarily on measuring and predicting changes in seawater carbon (C) chemistry and the biological and geochemical responses of reef organisms to such changes. To date, few ocean acidification studies have been designed to address conservation planning and management priorities. Here, we discuss how existing marine protected area design principles developed to address coral bleaching may be modified to address ocean acidification. We also identify five research priorities needed to incorporate ocean acidification into conservation planning and management: (1) establishing an ocean C chemistry baseline, (2) establishing ecological baselines, (3) determining species/habitat/community sensitivity to ocean acidification, (4) projecting changes in seawater carbonate chemistry, and (5) identifying potentially synergistic effects of multiple stressors.

Anthony, KRN, Diaz-Pulido G, Verlinden N, Tilbrook B, Andersson AJ.  2013.  Benthic buffers and boosters of ocean acidification on coral reefs. Biogeosciences. 10:4897-4909.   10.5194/bg-10-4897-2013   AbstractWebsite

Ocean acidification is a threat to marine ecosystems globally. In shallow-water systems, however, ocean acidification can be masked by benthic carbon fluxes, depending on community composition, seawater residence time, and the magnitude and balance of net community production (NCP) and calcification (NCC). Here, we examine how six benthic groups from a coral reef environment on Heron Reef (Great Barrier Reef, Australia) contribute to changes in the seawater aragonite saturation state (Omega(a)). Results of flume studies using intact reef habitats (1.2m by 0.4 m), showed a hierarchy of responses across groups, depending on CO2 level, time of day and water flow. At low CO2 (350-450 mu atm), macroalgae (Chnoospora implexa), turfs and sand elevated Omega(a) of the flume water by around 0.10 to 1.20 h(-1) - normalised to contributions from 1m(2) of benthos to a 1m deep water column. The rate of Omega(a) increase in these groups was doubled under acidification (560-700 mu atm) and high flow (35 compared to 8 cm s(-1)). In contrast, branching corals (Acropora aspera) increased Omega(a) by 0.25 h(-1) at ambient CO2 (350-450 mu atm) during the day, but reduced Omega(a) under acidification and high flow. Nighttime changes in Omega(a) by corals were highly negative (0.6-0.8 h(-1)) and exacerbated by acidification. Calcifying macroalgae (Halimeda spp.) raised Omega(a) by day (by around 0.13 h(-1)), but lowered Omega(a) by a similar or higher amount at night. Analyses of carbon flux contributions from benthic communities with four different compositions to the reef water carbon chemistry across Heron Reef flat and lagoon indicated that the net lowering of Omega(a) by coral-dominated areas can to some extent be countered by long water-residence times in neighbouring areas dominated by turfs, macroalgae and carbonate sand.

2012
Venti, A, Kadko D, Andersson AJ, Langdon C, Bates NR.  2012.  A multi-tracer model approach to estimate reef water residence times. Limnology and Oceanography-Methods. 10:1078-1095.   10.4319/lom.2012.10.1078   AbstractWebsite

We present a new method for obtaining the residence time of coral reef waters and demonstrate the successful application of this method by estimating rates of net ecosystem calcification (NEC) at four locations across the Bermuda platform and showing that the rates thus obtained are in reasonable agreement with independent estimates based on different methodologies. The contrast in Be-7 activity between reef and offshore waters can be related to the residence time of the waters over the reef through a time-dependent model that takes into account the rainwater flux of Be-7, the radioactive half-life of Be-7, and the rate of removal of Be-7 on particles estimated from Th-234. Sampling for Be-7 and Th-234 was conducted during the late fall and winter between 2008 and 2010. Model results yielded residence times ranging from 1.4 (+/- 0.7) days at the rim reef to 12 (+/- 4.0) days closer to shore. When combined with measurements of salinity-normalized total alkalinity anomalies, these residence times yielded platform-average NEC rates ranging from a maximum of 20.3 (+/- 7.0) mmolCaCO(3) m(-2) d(-1) in Nov 2008 to a minimum of 2.5 (+/- 0.8) mmolCaCO(3) m(-2) d(-1) in Feb 2009. The advantage of this new approach is that the rates of NEC obtained are temporally and spatially averaged. This novel approach for estimating NEC rates may be applicable to other coral reef ecosystems, providing an opportunity to assess how these rates may change in the context of ocean acidification.

Andersson, AJ, Mackenzie FT.  2012.  Revisiting four scientific debates in ocean acidification research. Biogeosciences. 9:893-905.   10.5194/bg-9-893-2012   AbstractWebsite

In recent years, ocean acidification has gained continuously increasing attention from scientists and a number of stakeholders and has raised serious concerns about its effects on marine organisms and ecosystems. With the increase in interest, funding resources, and the number of scientific investigations focusing on this environmental problem, increasing amounts of data and results have been produced, and a progressively growing and more rigorous understanding of this problem has begun to develop. Nevertheless, there are still a number of scientific debates, and in some cases misconceptions, that keep reoccurring at a number of forums in various contexts. In this article, we revisit four of these topics that we think require further thoughtful consideration including: (1) surface seawater CO2 chemistry in shallow water coastal areas, (2) experimental manipulation of marine systems using CO2 gas or by acid addition, (3) net versus gross calcification and dissolution, and (4) CaCO3 mineral dissolution and seawater buffering. As a summation of these topics, we emphasize that: (1) many coastal environments experience seawater pCO(2) that is significantly higher than expected from equilibrium with the atmosphere and is strongly linked to biological processes; (2) addition of acid, base or CO2 gas to seawater can all be useful techniques to manipulate seawater chemistry in ocean acidification experiments; (3) estimates of calcification or CaCO3 dissolution based on present techniques are measuring the net of gross calcification and dissolution; and (4) dissolution of metastable carbonate mineral phases will not produce sufficient alkalinity to buffer the pH and carbonate saturation state of shallow water environments on timescales of decades to hundreds of years to the extent that any potential negative effects on marine calcifiers will be avoided.

2011
Andersson, AJ, Mackenzie FT.  2011.  Technical comment on Kroeker et al. (2010) Meta-analysis reveals negative yet variable effects of ocean acidification on marine organisms. Ecology Letters, 13, 1419–1434. Ecology Letters. 14:E1-E2.: Blackwell Publishing Ltd   10.1111/j.1461-0248.2011.01646.x   AbstractWebsite

Meta-analysis of experimental results has been interpreted to imply that the calcification response of organisms depositing high Mg-calcite is more resilient to ocean acidification than organisms depositing aragonite/calcite. This conclusion might be biased by inadequate recognition and categorisation of high Mg-calcite according to mineral solubility.

2009
Andersson, AJ, Kuffner IB, Mackenzie FT, Jokiel PL, Rodgers KS, Tan A.  2009.  Net Loss of CaCO(3) from a subtropical calcifying community due to seawater acidification: mesocosm-scale experimental evidence. Biogeosciences. 6:1811-1823. AbstractWebsite

Acidification of seawater owing to oceanic uptake of atmospheric CO(2) originating from human activities such as burning of fossil fuels and land-use changes has raised serious concerns regarding its adverse effects on corals and calcifying communities. Here we demonstrate a net loss of calcium carbonate (CaCO(3)) material as a result of decreased calcification and increased carbonate dissolution from replicated subtropical coral reef communities (n=3) incubated in continuous-flow mesocosms subject to future seawater conditions. The calcifying community was dominated by the coral Montipora capitata. Daily average community calcification or Net Ecosystem Calcification (NEC=CaCO(3) production - dissolution) was positive at 3.3 mmol CaCO(3) m(-2) h(-1) under ambient seawater pCO(2) conditions as opposed to negative at -0.04 mmol CaCO(3) m(-2) h(-1) under seawater conditions of double the ambient pCO(2). These experimental results provide support for the conclusion that some net calcifying communities could become subject to net dissolution in response to anthropogenic ocean acidification within this century. Nevertheless, individual corals remained healthy, actively calcified (albeit slower than at present rates), and deposited significant amounts of CaCO(3) under the prevailing experimental seawater conditions of elevated pCO(2).

2008
Jokiel, PL, Rodgers KS, Kuffner IB, Andersson AJ, Cox EF, Mackenzie FT.  2008.  Ocean acidification and calcifying reef organisms: a mesocosm investigation. Coral Reefs. 27:473-483.   10.1007/s00338-008-0380-9   AbstractWebsite

A long-term (10 months) controlled experiment was conducted to test the impact of increased partial pressure of carbon dioxide (pCO(2)) on common calcifying coral reef organisms. The experiment was conducted in replicate continuous flow coral reef mesocosms flushed with unfiltered sea water from Kaneohe Bay, Oahu, Hawaii. Mesocosms were located in full sunlight and experienced diurnal and seasonal fluctuations in temperature and sea water chemistry characteristic of the adjacent reef flat. Treatment mesocosms were manipulated to simulate an increase in pCO(2) to levels expected in this century [midday pCO(2) levels exceeding control mesocosms by 365 +/- 130 mu atm (mean +/- sd)]. Acidification had a profound impact on the development and growth of crustose coralline algae (CCA) populations. During the experiment, CCA developed 25% cover in the control mesocosms and only 4% in the acidified mesocosms, representing an 86% relative reduction. Free-living associations of CCA known as rhodoliths living in the control mesocosms grew at a rate of 0.6 g buoyant weight year(-1) while those in the acidified experimental treatment decreased in weight at a rate of 0.9 g buoyant weight year(-1), representing a 250% difference. CCA play an important role in the growth and stabilization of carbonate reefs, so future changes of this magnitude could greatly impact coral reefs throughout the world. Coral calcification decreased between 15% and 20% under acidified conditions. Linear extension decreased by 14% under acidified conditions in one experiment. Larvae of the coral Pocillopora damicornis were able to recruit under the acidified conditions. In addition, there was no significant difference in production of gametes by the coral Montipora capitata after 6 months of exposure to the treatments.

Kuffner, IB, Andersson AJ, Jokiel PL, Rodgers KS, Mackenzie FT.  2008.  Decreased abundance of crustose coralline algae due to ocean acidification. Nature Geoscience. 1:114-117.   10.1038/ngeo100   AbstractWebsite

Owing to anthropogenic emissions, atmospheric concentrations of carbon dioxide could almost double between 2006 and 2100 according to business- as- usual carbon dioxide emission scenarios(1). Because the ocean absorbs carbon dioxide from the atmosphere(2-4), increasing atmospheric carbon dioxide concentrations will lead to increasing dissolved inorganic carbon and carbon dioxide in surface ocean waters, and hence acidification and lower carbonate saturation states(2,5). As a consequence, it has been suggested that marine calcifying organisms, for example corals, coralline algae, molluscs and foraminifera, will have difficulties producing their skeletons and shells at current rates(6,7), with potentially severe implications for marine ecosystems, including coral reefs(6,8 - 11). Here we report a seven-week experiment exploring the effects of ocean acidification on crustose coralline algae, a cosmopolitan group of calcifying algae that is ecologically important in most shallow-water habitats(12-14). Six outdoor mesocosms were continuously supplied with sea water from the adjacent reef and manipulated to simulate conditions of either ambient or elevated seawater carbon dioxide concentrations. The recruitment rate and growth of crustose coralline algae were severely inhibited in the elevated carbon dioxide mesocosms. Our findings suggest that ocean acidification due to human activities could cause significant change to benthic community structure in shallow-warm-water carbonate ecosystems.

2007
Andersson, AJ, Bates NR, Mackenzie FT.  2007.  Dissolution of carbonate sediments under rising pCO(2) and ocean acidification: Observations from Devil's Hole, Bermuda. Aquatic Geochemistry. 13:237-264.   10.1007/s10498-007-9018-8   AbstractWebsite

Rising atmospheric pCO(2) and ocean acidification originating from human activities could result in increased dissolution of metastable carbonate minerals in shallow-water marine sediments. In the present study, in situ dissolution of carbonate sedimentary particles in Devil's Hole, Bermuda, was observed during summer when thermally driven density stratification restricted mixing between the bottom water and the surface mixed layer and microbial decomposition of organic matter in the subthermocline layer produced pCO(2) levels similar to or higher than those levels anticipated by the end of the 21st century. Trends in both seawater chemistry and the composition of sediments in Devil's Hole indicate that Mg-calcite minerals are subject to selective dissolution under conditions of elevated pCO(2). The derived rates of dissolution based on observed changes in excess alkalinity and estimates of vertical eddy diffusion ranged from 0.2 mmol to 0.8 mmol CaCO3 m(-2) h(-1). On a yearly basis, this range corresponds to 175-701 g CaCO3 m(-2) year(-1); the latter rate is close to 50% of the estimate of the current average global coral reef calcification rate of about 1,500 g CaCO3 m(-2) year(-1). Considering a reduction in marine calcification of 40% by the year 2100, or 90% by 2300, as a result of surface ocean acidification, the combination of high rates of carbonate dissolution and reduced rates of calcification implies that coral reefs and other carbonate sediment environments within the 21st and following centuries could be subject to a net loss in carbonate material as a result of increasing pCO(2) arising from burning of fossil fuels.

2003
Andersson, AJ, Mackenzie FT, Ver LM.  2003.  Solution of shallow-water carbonates: An insignificant buffer against rising atmospheric CO2. Geology. 31:513-516.   10.1130/0091-7613(2003)031<0513:soscai>2.0.co;2   AbstractWebsite

Model predictions suggest that the saturation state of surface ocean waters with respect to carbonate minerals will decline during the twenty-first century owing to increased invasion of atmospheric CO2. As a result, calcareous organisms may have difficulty calcifying, leading to production of weaker skeletons and greater vulnerability to erosion. Alternatively, it has been suggested that there will be no significant impact on coral reef ecosystems because any changes in saturation state and pH will be restored by dissolution of metastable carbonate minerals. To resolve this controversy, we employ a physical-biogeochemical box model representative of the shallow-water ocean environment. Numerical simulations demonstrate that the carbonate saturation state of surface waters could significantly decrease and hamper the biogenic production of CaCO3 during the twenty-first century. Similarly, the average saturation state of marine pore waters could decline significantly, inducing dissolution of metastable carbonate phases within the pore-water-sediment system. Such dissolution could buffer the carbon chemistry of the pore waters, but overlying surface waters of reefs and other shallow-water carbonate environments will not accumulate sufficient alkalinity to buffer pH or carbonate saturation state changes owing to invasion of atmospheric CO2.