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Bates, NR, Amat A, Andersson AJ.  2010.  Feedbacks and responses of coral calcification on the Bermuda reef system to seasonal changes in biological processes and ocean acidification. Biogeosciences. 7:2509-2530.   10.5194/bg-7-2509-2010   AbstractWebsite

Despite the potential impact of ocean acidification on ecosystems such as coral reefs, surprisingly, there is very limited field data on the relationships between calcification and seawater carbonate chemistry. In this study, contemporaneous in situ datasets of seawater carbonate chemistry and calcification rates from the high-latitude coral reef of Bermuda over annual timescales provide a framework for investigating the present and future potential impact of rising carbon dioxide (CO(2)) levels and ocean acidification on coral reef ecosystems in their natural environment. A strong correlation was found between the in situ rates of calcification for the major framework building coral species Diploria labyrinthiformis and the seasonal variability of [CO(3)(2-)] and aragonite saturation state Omega(aragonite), rather than other environmental factors such as light and temperature. These field observations provide sufficient data to hypothesize that there is a seasonal 'Carbonate Chemistry Coral Reef Ecosystem Feedback' (CREF hypothesis) between the primary components of the reef ecosystem (i.e., scleractinian hard corals and macroalgae) and seawater carbonate chemistry. In early summer, strong net autotrophy from benthic components of the reef system enhance [CO(3)(2-)] and Omega(aragonite) conditions, and rates of coral calcification due to the photosynthetic uptake of CO(2). In late summer, rates of coral calcification are suppressed by release of CO(2) from reef metabolism during a period of strong net heterotrophy. It is likely that this seasonal CREF mechanism is present in other tropical reefs although attenuated compared to high-latitude reefs such as Bermuda. Due to lower annual mean surface seawater [CO(3)(2-)] and Omega(aragonite) in Bermuda compared to tropical regions, we anticipate that Bermuda corals will experience seasonal periods of zero net calcification within the next decade at [CO(3)(2-)] and Omega(aragonite) thresholds of similar to 184 mu moles kg(-1) and 2.65. However, net autotrophy of the reef during winter and spring (as part of the CREF hypothesis) may delay the onset of zero NEC or decalcification going forward by enhancing [CO(3)(2-)] and Omega(aragonite). The Bermuda coral reef is one of the first responders to the negative impacts of ocean acidification, and we estimate that calcification rates for D. labyrinthiformis have declined by > 50% compared to pre-industrial times.

Guest, JR, Edmunds PJ, Gates RD, Kuffner IB, Andersson AJ, Barnes BB, Chollett I, Courtney TA, Elahi R, Gross K, Lenz EA, Mitarai S, Mumby PJ, Nelson HR, Parker BA, Putnam HM, Rogers CS, Toth LT.  2018.  A framework for identifying and characterising coral reef "oases" against a backdrop of degradation. Journal of Applied Ecology. 55:2865-2875.   10.1111/1365-2664.13179   AbstractWebsite

1. Human activities have led to widespread ecological decline; however, the severity of degradation is spatially heterogeneous due to some locations resisting, escaping, or rebounding from disturbances. 2. We developed a framework for identifying oases within coral reef regions using long-term monitoring data. We calculated standardised estimates of coral cover (z-scores) to distinguish sites that deviated positively from regional means. We also used the coefficient of variation (CV) of coral cover to quantify how oases varied temporally, and to distinguish among types of oases. We estimated "coral calcification capacity" (CCC), a measure of the coral community's ability to produce calcium carbonate structures and tested for an association between this metric and z-scores of coral cover. 3. We illustrated our z-score approach within a modelling framework by extracting z-scores and CVs from simulated data based on four generalized trajectories of coral cover. We then applied the approach to time-series data from long-term reef monitoring programmes in four focal regions in the Pacific (the main Hawaiian Islands and Mo'orea, French Polynesia) and western Atlantic (the Florida Keys and St. John, US Virgin Islands). Among the 123 sites analysed, 38 had positive z-scores for median coral cover and were categorised as oases. 4. Synthesis and applications. Our framework provides ecosystem managers with a valuable tool for conservation by identifying "oases" within degraded areas. By evaluating trajectories of change in state (e.g., coral cover) among oases, our approach may help in identifying the mechanisms responsible for spatial variability in ecosystem condition. Increased mechanistic understanding can guide whether management of a particular location should emphasise protection, mitigation or restoration. Analysis of the empirical data suggest that the majority of our coral reef oases originated by either escaping or resisting disturbances, although some sites showed a high capacity for recovery, while others were candidates for restoration. Finally, our measure of reef condition (i.e., median z-scores of coral cover) correlated positively with coral calcification capacity suggesting that our approach identified oases that are also exceptional for one critical component of ecological function.

Andersson, A.  2015.  A fundamental paradigm for coral reef carbonate sediment dissolution. Frontiers in Marine Science. 2   10.3389/fmars.2015.00052   Abstract

The long-term success of coral reefs depends on a positive balance of calcium carbonate production exceeding dissolution, erosion, and material export. As a result of ocean acidification, coral reefs could transition from net accretion to net erosion owing to decreasing rates of calcification and increasing rates of chemical dissolution and bioerosion. Here, I present a fundamental paradigm that aims to explain the main driver of carbonate sediment dissolution on coral reefs based on theory and a new empirical dataset of pore water carbonate chemistry from the Bermuda coral reef platform. The paradigm shows that carbonate sediment dissolution is most strongly controlled by the extent of organic matter decomposition in the sediments, but that the magnitude of dissolution is influenced by how much decomposition is required to reach pore water undersaturation with respect to the most soluble bulk carbonate mineral phase present in the sediments, a condition defined as the Carbonate Critical Threshold (CCT). Decomposition of organic matter beyond the CCT under aerobic conditions results in stoichiometric proportional dissolution of carbonate sediments. As ocean acidification proceeds over the next several decades, the extent of organic matter decomposition required to reach the CCT will decrease, carbonate dissolution will increase, and subsequently the accumulation of carbonate sediments will decrease. Since drastic reductions in anthropogenic CO2 emission are unlikely in the foreseeable future, the paradigm shows that active controls and reduction of organic matter input to coral reefs at the local scale might be an effective mitigation strategy to prevent or delay coral reefs transitioning to a state of net dissolution.

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Beman, JM, Chow CE, King AL, Feng YY, Fuhrman JA, Andersson A, Bates NR, Popp BN, Hutchins DA.  2011.  Global declines in oceanic nitrification rates as a consequence of ocean acidification. Proceedings of the National Academy of Sciences of the United States of America. 108:208-213.   10.1073/pnas.1011053108   AbstractWebsite

Ocean acidification produced by dissolution of anthropogenic carbon dioxide (CO2) emissions in seawater has profound consequences for marine ecology and biogeochemistry. The oceans have absorbed one-third of CO2 emissions over the past two centuries, altering ocean chemistry, reducing seawater pH, and affecting marine animals and phytoplankton in multiple ways. Microbially mediated ocean biogeochemical processes will be pivotal in determining how the earth system responds to global environmental change; however, how they may be altered by ocean acidification is largely unknown. We show here that microbial nitrification rates decreased in every instance when pH was experimentally reduced (by 0.05-0.14) at multiple locations in the Atlantic and Pacific Oceans. Nitrification is a central process in the nitrogen cycle that produces both the greenhouse gas nitrous oxide and oxidized forms of nitrogen used by phytoplankton and other microorganisms in the sea; at the Bermuda Atlantic Time Series and Hawaii Ocean Time-series sites, experimental acidification decreased ammonia oxidation rates by 38% and 36%. Ammonia oxidation rates were also strongly and inversely correlated with pH along a gradient produced in the oligotrophic Sargasso Sea (r(2) = 0.87, P < 0.05). Across all experiments, rates declined by 8-38% in low pH treatments, and the greatest absolute decrease occurred where rates were highest off the California coast. Collectively our results suggest that ocean acidification could reduce nitrification rates by 3-44% within the next few decades, affecting oceanic nitrous oxide production, reducing supplies of oxidized nitrogen in the upper layers of the ocean, and fundamentally altering nitrogen cycling in the sea.

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Reid, PC, Fischer AC, Lewis-Brown E, Meredith MP, Sparrow M, Andersson AJ, Antia A, Bates NR, Bathmann U, Beaugrand G, Brix H, Dye S, Edwards M, Furevik T, Gangsto R, Hatun H, Hopcroft RR, Kendall M, Kasten S, Keeling R, Le Quere C, Mackenzie FT, Malin G, Mauritzen C, Olafsson J, Paull C, Rignot E, Shimada K, Vogt M, Wallace C, Wang ZM, Washington R.  2009.  Impacts of the Oceans on Climate Change. Advances in Marine Biology, Vol 56. 56( Sims DW, Ed.).:1-150., San Diego: Elsevier Academic Press Inc   10.1016/s0065-2881(09)56001-4   Abstract

The oceans play a key role in climate regulation especially in part buffering (neutralising) the effects of increasing levels of greenhouse gases in the atmosphere and rising global temperatures. This chapter examines how the regulatory processes performed by the oceans alter as a response to climate change and assesses the extent to which positive feedbacks from the ocean may exacerbate climate change. There is clear evidence for rapid change in the oceans. As the main heat store for the world there has been an accelerating change in sea temperatures over the last few decades, which has contributed to rising sea-level. The oceans are also the main store of carbon dioxide (CO(2)), and are estimated to have taken up similar to 40% of anthropogenic-sourced CO(2) from the atmosphere since the beginning of the industrial revolution. A proportion of the carbon uptake is exported via the four ocean 'carbon pumps' (Solubility, Biological, Continental Shelf and Carbonate Counter) to the deep ocean reservoir. Increases in sea temperature and changing planktonic systems and ocean currents may lead to a reduction in the uptake of CO(2) by the ocean; some evidence suggests a suppression of parts of the marine carbon sink is already underway. While the oceans have buffered climate change through the uptake of CO(2) produced by fossil fuel burning this has already had an impact on ocean chemistry through ocean acidification and will continue to do so. Feedbacks to climate change from acidification may result from expected impacts on marine organisms (especially corals and calcareous plankton), ecosystems and biogeochemical cycles. The polar regions of the world are showing the most rapid responses to climate change. As a result of a strong ice-ocean influence, small changes in temperature, salinity and ice cover may trigger large and sudden changes in regional climate with potential downstream feedbacks to the climate of the rest of the world. A warming Arctic Ocean may lead to further releases of the potent greenhouse gas methane from hydrates and permafrost. The Southern Ocean plays a critical role in driving, modifying and regulating global climate change via the carbon cycle and through its impact on adjacent Antarctica. The Antarctic Peninsula has shown some of the most rapid rises in atmospheric and oceanic temperature in the world, with an associated retreat of the majority of glaciers. Parts of the West Antarctic ice sheet are deflating rapidly, very likely due to a change in the flux of oceanic heat to the undersides of the floating ice shelves. The final section on modelling feedbacks from the ocean to climate change identifies limitations and priorities for model development and associated observations. Considering the importance of the oceans to climate change and our limited understanding of climate-related ocean processes, our ability to measure the changes that are taking place are conspicuously inadequate. The chapter highlights the need for a comprehensive, adequately funded and globally extensive ocean observing system to be implemented and sustained as a high priority. Unless feedbacks from the oceans to climate change are adequately included in climate change models, it is possible that the mitigation actions needed to stabilise CO(2) and limit temperature rise over the next century will be underestimated.

Morse, JW, Andersson AJ, Mackenzie FT.  2006.  Initial responses of carbonate-rich shelf sediments to rising atmospheric pCO(2) and "ocean acidification": Role of high Mg-calcites. Geochimica Et Cosmochimica Acta. 70:5814-5830.   10.1016/j.gca.2006.08.017   AbstractWebsite

Carbonate-rich sediments at shoal to shelf depths (< 200 m) represent a major CaCO(3) reservoir that can rapidly react to the decreasing saturation state of seawater with respect to carbonate minerals, produced by the increasing partial pressure of atmospheric carbon dioxide (pCO(2)) and "acidification" of ocean waters. Aragonite is usually the most abundant carbonate mineral in these sediments. However, the second most abundant (typically similar to 24 wt%) carbonate mineral is high Mg-calcite (Mg-calcite) whose solubility can exceed that of aragonite making it the "first responder" to the decreasing saturation state of seawater. For the naturally occurring biogenic Mg-calcites, dissolution experiments have been used to predict their "stoichiometric solubilities" as a function of mol% MgCO(3). The only valid relationship that one can provisionally use for the metastable stabilities for Mg-calcite based on composition is that for the synthetically produced phases where metastable equilibrium has been achieved from both under- and over-saturation. Biogenic Mg-calcites exhibit a large offset in solubility from that of abiotic Mg-calcite and can also exhibit a wide range of solubilities for biogenic Mg-calcites of similar Mg content. This indicates that factors other than the Mg content can influence the solubility of these mineral phases. Thus, it is necessary to turn to observations of natural sediments where changes in the saturation state of surrounding waters occur in order to determine their likely responses to the changing saturation state in upper oceanic waters brought on by increasing pCO(2). In the present study, we investigate the responses of Mg-calcites to rising pCO(2) and "ocean acidification" by means of a simple numerical model based on the experimental range of biogenic Mg-calcite solubilities as a function of Mg content in order to bracket the behavior of the most abundant Mg-calcite phases in the natural environment. In addition, observational data from Bermuda and the Great Bahama Bank are also presented in order to project future responses of these minerals. The numerical simulations suggest that Mg-calcite minerals will respond to rising pCO(2) by sequential dissolution according to mineral stability, progressively leading to removal of the more soluble phases until the least soluble phases remain. These results are confirmed by laboratory experiments and observations from Bermuda. As a consequence of continuous increases in atmospheric CO, from burning of fossil fuels, the average composition of contemporary carbonate sediments could change, i.e., the average Mg content in the sediments may slowly decrease. Furthermore, evidence from the Great Bahama Bank indicates that the amount of abiotic carbonate production is likely to decline as pCO(2) continues to rise. (c) 2006 Elsevier Inc. All rights reserved.

Edmunds, PJ, Comeau S, Lantz C, Andersson A, Briggs C, Cohen A, Gattuso JP, Grady JM, Gross K, Johnson M, Muller EB, Ries JB, Tambutte S, Tambutte E, Venn A, Carpenter RC.  2016.  Integrating the effects of ocean acidification across functional scales on tropical coral reefs. Bioscience. 66:350-362.   10.1093/biosci/biw023   AbstractWebsite

There are concerns about the future of coral reefs in the face of ocean acidification and warming, and although studies of these phenomena have advanced quickly, efforts have focused on pieces of the puzzle rather than integrating them to evaluate ecosystem-level effects. The field is now poised to begin this task, but there are information gaps that first must be overcome before progress can be made. Many of these gaps focus on calcification at the levels of cells, organisms, populations, communities, and ecosystem, and their closure will be made difficult by the complexity of the interdependent processes by which coral reefs respond to ocean acidification, with effects scaling from cells to ecosystems and from microns to kilometers. Existing ecological theories provide an important and largely untapped resource for overcoming these difficulties, and they offer great potential for integrating the effects of ocean acidification across scales on coral reefs.

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Mackenzie, FT, Andersson AJ, Arvidson RS, Guidry MW, Lerman A.  2011.  Land-sea carbon and nutrient fluxes and coastal ocean CO(2) exchange and acidification: Past, present, and future. Applied Geochemistry. 26:S298-S302.   10.1016/j.apgeochem.2011.03.087   AbstractWebsite

Epochs of changing atmospheric CO(2) and seawater CO(2)-carbonic acid system chemistry and acidification have occurred during the Phanerozoic at various time scales. On the longer geologic time scale, as sea level rose and fell and continental free board decreased and increased, respectively, the riverine fluxes of Ca, Mg, DIC, and total alkalinity to the coastal ocean varied and helped regulate the C chemistry of seawater, but nevertheless there were major epochs of ocean acidification (OA). On the shorter glacial-interglacial time scale from the Last Glacial Maximum (LGM) to late preindustrial time, riverine fluxes of DIC, total alkalinity, and N and P nutrients increased and along with rising sea level, atmospheric PCO(2) and temperature led, among other changes, to a slightly deceasing pH of coastal and open ocean waters, and to increasing net ecosystem calcification and decreasing net heterotrophy in coastal ocean waters. From late preindustrial time to the present and projected into the 21st century, human activities, such as fossil fuel and land-use emissions of CO(2) to the atmosphere, increasing application of N and P nutrient subsidies and combustion N to the landscape, and sewage discharges of C, N, P have led, and will continue to lead, to significant modifications of coastal ocean waters. The changes include a rapid decline in pH and carbonate saturation state (modern problem of ocean acidification), a shift toward dissolution of carbonate substrates exceeding production, potentially leading to the "demise" of the coral reefs, reversal of the direction of the sea-to-air flux of CO(2) and enhanced biological production and burial of organic C, a small sink of anthropogenic CO(2), accompanied by a continuous trend toward increasing autotrophy in coastal waters. (C) 2011 Elsevier Ltd. All rights reserved.

Andersson, AJ, Mackenzie FT, Bates NR.  2008.  Life on the margin: implications of ocean acidification on Mg-calcite, high latitude and cold-water marine calcifiers. Marine Ecology-Progress Series. 373:265-273.   10.3354/meps07639   AbstractWebsite

Future anthropogenic emissions of CO(2) and the resulting ocean acidification may have severe consequences for marine calcifying organisms and ecosystems. Marine calcifiers depositing calcitic hard parts that contain significant concentrations of magnesium, i.e. Mg-calcite, and calcifying organisms living in high latitude and/or cold-water environments are at immediate risk to ocean acidification and decreasing seawater carbonate saturation because they are currently immersed in seawater that is just slightly supersaturated with respect to the carbonate phases they secrete. Under the present rate Of CO(2) emissions, model calculations show that high latitude ocean waters could reach undersaturation with respect to aragonite in just a few decades. Thus, before this happens these waters will be undersaturated with respect to Mg-calcite minerals of higher solubility than that of aragonite. Similarly, tropical surface seawater could become undersaturated with respect to Mg-calcite minerals containing >= 12 mole percent (mol%) MgCO(3) during this century. As a result of these changes in surface seawater chemistry and further penetration of anthropogenic CO(2) into the ocean interior, we suggest that (1) the magnesium content of calcitic hard parts will decrease in many ocean environments, (2) the relative proportion of calcifiers depositing stable carbonate minerals, such as calcite and low Mg-calcite, will increase and (3) the average magnesium content of carbonate sediments will decrease. Furthermore, the highest latitude and deepest depth at which cold-water corals and other calcifiers currently exist will move towards lower latitudes and shallower depth, respectively. These changes suggest that anthropogenic emissions of CO(2) may be currently pushing the oceans towards an episode characteristic of a 'calcite sea.'

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Parson, RJ, Nelson CA, Carlson CA, Denman CC, Andersson AJ, Kledzik AL, Vergin KL, McNally SP, Treusch AH, Giovannoni SJ.  2014.  Marine bacterioplankton community turnover within seasonally hypoxic waters of a subtropical sound: Devil’s Hole, Bermuda. Environmental Microbiology.   10.1111/1462-2920.12445   Abstract

Understanding bacterioplankton community dynamics in coastal hypoxic environments is relevant to global biogeochemistry because coastal hypoxia is increasing worldwide. The temporal dynamics of bacterioplankton communities were analysed throughout the illuminated water column of Devil's Hole, Bermuda during the 6-week annual transition from a strongly stratified water column with suboxic and high-pCO2 bottom waters to a fully mixed and ventilated state during 2008. A suite of culture-independent methods provided a quantitative spatiotemporal characterization of bacterioplankton community changes, including both direct counts and rRNA gene sequencing. During stratification, the surface waters were dominated by the SAR11 clade of Alphaproteobacteria and the cyanobacterium Synechococcus. In the suboxic bottom waters, cells from the order Chlorobiales prevailed, with gene sequences indicating members of the genera Chlorobium and Prosthecochloris – anoxygenic photoautotrophs that utilize sulfide as a source of electrons for photosynthesis. Transitional zones of hypoxia also exhibited elevated levels of methane- and sulfur-oxidizing bacteria relative to the overlying waters. The abundance of both Thaumarcheota and Euryarcheota were elevated in the suboxic bottom waters (> 109 cells l−1). Following convective mixing, the entire water column returned to a community typical of oxygenated waters, with Euryarcheota only averaging 5% of cells, and Chlorobiales and Thaumarcheota absent.

Mackenzie, FT, Andersson AJ.  2013.  The marine carbon system and ocean acidification during Phanerozoic time. Geochemical Perspectives. 2:1-227.   10.7185/geochempersp.2.1   AbstractWebsite

The global CO2-carbonic acid-carbonate system of seawater, although certainly a well-researched topic of interest in the past, has risen to the fore in recent years because of the environmental issue of ocean acidification (often simply termed OA). Despite much previous research, there remain pressing questions about how this most important chemical system of seawater operated at the various time scales of the deep time of the Phanerozoic Eon (the past 545 Ma of Earth's history), interglacial-glacial time, and the Anthropocene (the time of strong human influence on the behaviour of the system) into the future of the planet. One difficulty in any analysis is that the behaviour of the marine carbon system is not only controlled by internal processes in the ocean, but it is intimately linked to the domains of the atmosphere, continental landscape, and marine carbonate sediments.

Langdon, CR, Gatusso JP, Andersson AJ.  2010.  Measurements of calcification abd dissolution of benthic organisms and communities. Guidebest practices in ocean acidification Reserach and data reporting . Abstract
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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.

Venti, A, Andersson A, Langdon C.  2014.  Multiple driving factors explain spatial and temporal variability in coral calcification rates on the Bermuda platform. Coral Reefs. 33:979-997.   10.1007/s00338-014-1191-9   AbstractWebsite

Experimental studies have shown that coral calcification rates are dependent on light, nutrients, food availability, temperature, and seawater aragonite saturation (Omega (arag)), but the relative importance of each parameter in natural settings remains uncertain. In this study, we applied Calcein fluorescent dyes as time indicators within the skeleton of coral colonies (n = 3) of Porites astreoides and Diploria strigosa at three study sites distributed across the northern Bermuda coral reef platform. We evaluated the correlation between seasonal average growth rates based on coral density and extension rates with average temperature, light, and seawater Omega (arag) in an effort to decipher the relative importance of each parameter. The results show significant seasonal differences among coral calcification rates ranging from summer maximums of 243 +/- A 58 and 274 +/- A 57 mmol CaCO3 m(-2) d(-1) to winter minimums of 135 +/- A 39 and 101 +/- A 34 mmol CaCO3 m(-2) d(-1) for P. astreoides and D. strigosa, respectively. We also placed small coral colonies (n = 10) in transparent chambers and measured the instantaneous rate of calcification under light and dark treatments at the same study sites. The results showed that the skeletal growth of D. strigosa and P. astreoides, whether hourly or seasonal, was highly sensitive to Omega (arag). We believe this high sensitivity, however, is misleading, due to covariance between light and Omega (arag), with the former being the strongest driver of calcification variability. For the seasonal data, we assessed the impact that the observed seasonal differences in temperature (4.0 A degrees C), light (5.1 mol photons m(-2) d(-1)), and Omega (arag) (0.16 units) would have on coral growth rates based on established relationships derived from laboratory studies and found that they could account for approximately 44, 52, and 5 %, respectively, of the observed seasonal change of 81 +/- A 14 mmol CaCO3 m(-2) d(-1). Using short-term light and dark incubations, we show how the covariance of light and Omega (arag) can lead to the false conclusion that calcification is more sensitive to Omega (arag) than it really is.

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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).

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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.

Andersson, AJ, Gledhill D.  2013.  Ocean Acidification and Coral Reefs: Effects on Breakdown, Dissolution, and Net Ecosystem Calcification. Annual Review of Marine Science. 5:321-348.   doi:10.1146/annurev-marine-121211-172241   AbstractWebsite

The persistence of carbonate structures on coral reefs is essential in providing habitats for a large number of species and maintaining the extraordinary biodiversity associated with these ecosystems. As a consequence of ocean acidification (OA), the ability of marine calcifiers to produce calcium carbonate (CaCO3) and their rate of CaCO3 production could decrease while rates of bioerosion and CaCO3 dissolution could increase, resulting in a transition from a condition of net accretion to one of net erosion. This would have negative consequences for the role and function of coral reefs and the eco-services they provide to dependent human communities. In this article, we review estimates of bioerosion, CaCO3 dissolution, and net ecosystem calcification (NEC) and how these processes will change in response to OA. Furthermore, we critically evaluate the observed relationships between NEC and seawater aragonite saturation state (Ωa). Finally, we propose that standardized NEC rates combined with observed changes in the ratios of dissolved inorganic carbon to total alkalinity owing to net reef metabolism may provide a biogeochemical tool to monitor the effects of OA in coral reef environments.

Andersson, AJ.  2014.  The oceanic CaCO3 cycle. Treatise on Geochemistry. Vol. 8( Holland HD, Turekian KK, Eds.)., Oxford: Elsevier   10.1016/B978-0-08-095975-7.00619-7  
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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.

Mackenzie, FT, Lerman A, Andersson AJ.  2004.  Past and present of sediment and carbon biogeochemical cycling models. Biogeosciences. 1:11-32. AbstractWebsite

The global carbon cycle is part of the much more extensive sedimentary cycle that involves large masses of carbon in the Earth's inner and outer spheres. Studies of the carbon cycle generally followed a progression in knowledge of the natural biological, then chemical, and finally geological processes involved, culminating in a more or less integrated picture of the biogeochemical carbon cycle by the 1920s. However, knowledge of the ocean's carbon cycle behavior has only within the last few decades progressed to a stage where meaningful discussion of carbon processes on an annual to millennial time scale can take place. In geologically older and pre-industrial time, the ocean was generally a net source of CO2 emissions to the atmosphere owing to the mineralization of land-derived organic matter in addition to that produced in situ and to the process of CaCO3 precipitation. Due to rising atmospheric CO2 concentrations because of fossil fuel combustion and land use changes, the direction of the air-sea CO2 flux has reversed, leading to the ocean as a whole being a net sink of anthropogenic CO2. The present thickness of the surface ocean layer, where part of the anthropogenic CO2 emissions are stored, is estimated as of the order of a few hundred meters. The oceanic coastal zone net air-sea CO2 exchange flux has also probably changed during industrial time. Model projections indicate that in preindustrial times, the coastal zone may have been net heterotrophic, releasing CO2 to the atmosphere from the imbalance between gross photosynthesis and total respiration. This, coupled with extensive CaCO3 precipitation in coastal zone environments, led to a net flux of CO2 out of the system. During industrial time the coastal zone ocean has tended to reverse its trophic status toward a non-steady state situation of net autotrophy, resulting in net uptake of anthropogenic CO2 and storage of carbon in the coastal ocean, despite the significant calcification that still occurs in this region. Furthermore, evidence from the inorganic carbon cycle indicates that deposition and net storage of CaCO3 in sediments exceed inflow of inorganic carbon from land and produce CO2 emissions to the atmosphere. In the shallow-water coastal zone, increase in atmospheric CO2 during the last 300 years of industrial time may have reduced the rate of calcification, and continuation of this trend is an issue of serious environmental concern in the global carbon balance.

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.

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Sato, KN, Andersson AJ, Day JMD, Taylor JRA, Frank MB, Jung JY, McKittrick J, Levin LA.  2018.  Response of sea urchin fitness traits to environmental gradients across the Southern California oxygen minimum zone. Frontiers in Marine Science. 5   10.3389/fmars.2018.00258   AbstractWebsite

Marine calcifiers are considered to be among the most vulnerable taxa to climate-forced environmental changes occurring on continental margins with effects hypothesized to occur on microstructural, biomechanical, and geochemical properties of carbonate structures. Natural gradients in temperature, salinity, oxygen, and pH on an upwelling margin combined with the broad depth distribution (100-1,100 m) of the pink fragile sea urchin, Strongylocentrotus (formerly Allocentrotus) fragilis, along the southern California shelf and slope provide an ideal system to evaluate potential effects of multiple climate variables on carbonate structures in situ. We measured, for the first time, trait variability across four distinct depth zones using natural gradients as analogues for species-specific implications of oxygen minimum zone (OMZ) expansion, deoxygenation and ocean acidification. Although S. fragilis may likely be tolerant of future oxygen and pH decreases predicted during the twenty-first century, we determine from adults collected across multiple depth zones that urchin size and potential reproductive fitness (gonad index) are drastically reduced in the OMZ core (450-900 m) compared to adjacent zones. Increases in porosity and mean pore size coupled with decreases in mechanical nanohardness and stiffness of the calcitic endoskeleton in individuals collected from lower pH(Total) (7.57-7.59) and lower dissolved oxygen (13-42 mu mol kg(-1)) environments suggest that S. fragilis may be potentially vulnerable to crushing predators if these conditions become more widespread in the future. In addition, elemental composition indicates that S. fragilis has a skeleton composed of the low Mg-calcite mineral phase of calcium carbonate (mean Mg/Ca = 0.02 mol mol(-1)), with Mg/Ca values measured in the lower end of values reported for sea urchins known to date. Together these findings suggest that ongoing declines in oxygen and pH will likely affect the ecology and fitness of a dominant echinoid on the California margin.

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.

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Andersson, AJ, Krug LA, Bates NR, Doney SC.  2013.  Sea-air CO2 flux in the North Atlantic subtropical gyre: Role and influence of Sub-Tropical Mode Water formation. Deep-Sea Research Part Ii-Topical Studies in Oceanography. 91:57-70.   10.1016/j.dsr2.2013.02.022   AbstractWebsite

The uptake of atmospheric carbon dioxide (CO2) into the mid-latitudes of the North Atlantic Ocean through the production of wintertime Sub-Tropical Mode Water (STMW) also known as Eighteen Degree Water (EDW) is poorly quantified and constrained. Nonetheless, it has been proposed that the EDW could serve as an important short-term sink of anthropogenic CO2. The objective of the present investigation was to determine sea-air CO2 gas exchange rates and seawater CO2 dynamics during wintertime formation of EDW in the North Atlantic Ocean. During 2006 and 2007, several research cruises were undertaken as part of the CLIMODE project across the northwest Atlantic Ocean with the intent to study the pre-conditioning, formation, and the evolution of EDW. Sea-air CO2 exchange rates were calculated based on measurements of atmospheric pCO(2), surface seawater pCO(2) and wind speed with positive values denoting a net flux from the surface ocean to the atmosphere. Average sea-air CO2 flux calculated along cruise tracks in the formation region equaled -18 +/- 6 mmol CO2 m(-2) d(-1) and -14 +/- 9 mmol CO2 m(-2) d(-1) in January of 2006 and March of 2007, respectively. Average sea-air CO2 flux in newly formed outcropping EDW in February and March of 2007 equaled -28 +/- 10 mmol CO2 m(-2) d(-1). These estimates exceeded previous flux estimates in this region by 40-185%. The magnitude of CO2 flux was mainly controlled by the observed variability in wind speed and Delta pCO(2) with smaller changes owing to variability in sea surface temperature. Small but statistically significant difference (4.1 +/- 2.6 mu mol kg(-1)) in dissolved inorganic carbon (DIC) was observed in two occurrences of newly formed EDW in February and March of 2007. This difference was explained either by differences in the relative contribution from different water masses involved in the initial formation process of EDW or temporal changes owing to sea-air CO2 exchange (similar to 25%) and vertical and/or lateral mixing (similar to 75%) with water masses high in DIC from the cold side of the Gulf Stream and/or from below the permanent thermocline. Based on the present estimate of sea-air CO2 flux in newly formed EDW and a formation rate of 9.3 Sv y (Sverdrup year = 10(6) m(3) s(-1) flow sustained for 1 year), CO2 uptake by newly formed EDW may constitute 3-6% of the total North Atlantic CO2 sink. However, advection of surface waters that carry an elevated burden of anthropogenic CO2 that are transported to the formation region and transformed to mode water may contribute additional CO2 to the total net uptake and sequestration of anthropogenic CO2 to the ocean interior. (c) 2013 Elsevier Ltd. All rights reserved.

Bresnahan, PJ, Wirth T, Martz TR, Andersson AJ, Cyronak T, D’Angelo S, Pennise J, Melville KW, Lenain L, Statom N.  2016.  A sensor package for mapping pH and oxygen from mobile platforms. Methods in Oceanography. 17:1-13.   10.1016/j.mio.2016.04.004   Abstract

A novel chemical sensor package named “WavepHOx” was developed in order to facilitate measurement of surface ocean pH, dissolved oxygen, and temperature from mobile platforms. The system comprises a Honeywell Durafet pH sensor, Aanderaa optode oxygen sensor, and chloride ion selective electrode, packaged into a hydrodynamic, lightweight housing. The WavepHOx has been deployed on a stand-up paddleboard and a Liquid Robotics Wave Glider in multiple near-shore settings in the Southern California Bight. Integration of the WavepHOx into these mobile platforms has enabled high spatiotemporal resolution pH and dissolved oxygen data collection. It is a particularly valuable tool for mapping shallow, fragile, or densely vegetated ecosystems which cannot be easily accessed by other platforms. Results from three surveys in San Diego, California, are reported. We show pH and dissolved oxygen variability >0.3 and >50% saturation, respectively, over tens to hundreds of meters to highlight the degree of natural spatial variability in these vegetated ecosystems. When deployed during an extensive discrete sampling program, the WavepHOx pH had a root mean squared error of 0.028 relative to pH calculated from fifty six measurements of total alkalinity and dissolved inorganic carbon, confirming its capacity for accurate, high spatiotemporal resolution data collection.