Seawater carbonate chemistry and benthic marine community during experiments, 2011

Ocean acidification is predicted to impact all areas of the oceans and affect a diversity of marine organisms. However, the diversity of responses among species prevents clear predictions about the impact of acidification at the ecosystem level. Here, we used shallow water CO2 vents in the Mediterranean Sea as a model system to examine emergent ecosystem responses to ocean acidification in rocky reef communities. We assessed in situ benthic invertebrate communities in three distinct pH zones (ambient, low, and extreme low), which differed in both the mean and variability of seawater pH along a continuous gradient. We found fewer taxa, reduced taxonomic evenness, and lower biomass in the extreme low pH zones. However, the number of individuals did not differ among pH zones, suggesting that there is density compensation through population blooms of small acidification-tolerant taxa. Furthermore, the trophic structure of the invertebrate community shifted to fewer trophic groups and dominance by generalists in extreme low pH, suggesting that there may be a simplification of food webs with ocean acidification. Despite high variation in individual species' responses, our findings indicate that ocean acidification decreases the diversity, biomass, and trophic complexity of benthic marine communities. These results suggest that a loss of biodiversity and ecosystem function is expected under extreme acidification scenarios.

Groundfish survey data from the German Bight from 1902-08, 1919-23, and 1930-1932 and ICES International Bottom Trawl Survey (IBTS) quarter 3 data from 1991 to 2009

Groundfish survey data from the German Bight from 1902-08, 1919-23, and 1930-1932 and ICES International Bottom Trawl Survey (IBTS) quarter 3 data from 1991 to 2009 were analysed with respect to species frequencies, maximum length, trends in catch-per-unit-effort, species richness parameters (SNR) and presence of large fish (Phi40), the latter defined as average presence of species per haul with specimens larger than 40 cm given. Four different periods are distinguished: (a) before 1914 with medium commercial CPUE and low landings, Phi40 approx. 2, high abundance in elasmobranchs and SNR conditions indicating highly diverse assemblages, (b) conditions immediately after 1918 with higher commercial CPUE, recovering landings, Phi40 at > 4 in 1919, and SNR conditions indicating highly diverse assemblages, (c) conditions from 1920 to the early 1930's with decreasing commercial CPUE, increased landings, decreasing Phi40, SNR conditions similar to later years indicating less diverse assemblages, and a decrease in elasmobranchs. In the IBTS series (d), Phi40 remains low indicating an increased rarity of large specimens, and SNR characteristics are similar to the third period. Dab, whiting and grey gurnard have increased considerably in the IBTS series as compared to the historic data. Phi40 is suggested an alternative indicator reflecting community functional diversity when weight based indicators cannot be applied.

Seasonality affects macroalgal community response to increases in pCO2

Ocean acidification is expected to alter marine systems, but there is uncertainty about its effects due to the logistical difficulties of testing its large-scale and long-term effects. Responses of biological communities to increases in carbon dioxide can be assessed at CO2 seeps that cause chronic exposure to lower seawater pH over localised areas of seabed. Shifts in macroalgal communities have been described at temperate and tropical pCO2 seeps, but temporal and spatial replication of these observations is needed to strengthen confidence our predictions, especially because very few studies have been replicated between seasons. Here we describe the seawater chemistry and seasonal variability of macroalgal communities at CO2 seeps off Methana (Aegean Sea). Monitoring from 2011 to 2013 showed that seawater pH decreased to levels predicted for the end of this century at the seep site with no confounding gradients in Total Alkalinity, salinity, temperature or wave exposure. Most nutrient levels were similar along the pH gradient; silicate increased significantly with decreasing pH, but it was not limiting for algal growth at all sites. Metal concentrations in seaweed tissues varied between sites but did not consistently increase with pCO2. Our data on the flora are consistent with results from laboratory experiments and observations at Mediterranean CO2 seep sites in that benthic communities decreased in calcifying algal cover and increased in brown algal cover with increasing pCO2. This differs from the typical macroalgal community response to stress, which is a decrease in perennial brown algae and proliferation of opportunistic green algae. Cystoseira corniculata was more abundant in autumn and Sargassum vulgare in spring, whereas the articulated coralline alga Jania rubens was more abundant at reference sites in autumn. Diversity decreased with increasing CO2 regardless of season. Our results show that benthic community responses to ocean acidification are strongly affected by season.

Seawater carbonate chemistry and community calcification during a Molokai reef (Hawaii) study 2006

The severity of the impact of elevated atmospheric pCO2 to coral reef ecosystems depends, in part, on how seawater pCO2 affects the balance between calcification and dissolution of carbonate sediments. Presently, there are insufficient published data that relate concentrations of pCO2 and CO3 to in situ rates of reef calcification in natural settings to accurately predict the impact of elevated atmospheric pCO2 on calcification and dissolution processes. Rates of net calcification and dissolution, CO3 concentrations, and pCO2 were measured, in situ, on patch reefs, bare sand, and coral rubble on the Molokai reef flat in Hawaii. Rates of calcification ranged from 0.03 to 2.30 mmol CaCO3 m**-2 h**-1 and dissolution ranged from -0.05 to -3.3 mmol CaCO3 m**-2 h**-1. Calcification and dissolution varied diurnally with net calcification primarily occurring during the day and net dissolution occurring at night. These data were used to calculate threshold values for pCO2 and CO3 at which rates of calcification and dissolution are equivalent. Results indicate that calcification and dissolution are linearly correlated with both CO3 and pCO2. Threshold pCO2 and CO3 values for individual substrate types showed considerable variation. The average pCO2 threshold value for all substrate types was 654±195 µatm and ranged from 467 to 1003 µatm. The average CO3 threshold value was 152±24 µmol/kg, ranging from 113 to 184 µmol/kg. Ambient seawater measurements of pCO2 and CO3 indicate that CO3 and pCO2 threshold values for all substrate types were both exceeded, simultaneously, 13% of the time at present day atmospheric pCO2 concentrations. It is predicted that atmospheric pCO2 will exceed the average pCO2 threshold value for calcification and dissolution on the Molokai reef flat by the year 2100.

Seawater carbonate chemistry and particulate organic particles during an incubation experiments with natural phytoplankton community, 2002

Incubation experiments with natural phytoplankton revealed a relationship between CO2 concentration and the production of transparent exopolymer particles (TEP), with TEP production being linearly related to theoretical CO2 uptake rates. The effect of different CO2 concentrations on TEP production was examined during incubation experiments with natural phytoplankton sampled at two different locations in the central Baltic Sea in summer 1999.

Seawater carbonate chemistry and community calcification during Miyako Island (Japan) coral reef studies, 1994

Coral reefs are characterized by enormous carbonate production of the organisms. It is known that rapid calcification is linked to photosynthesis under control of the carbonate equilibrium in seawater. We have established a model simulating the coexisting states of photosynthesis and calcification in order to examine the effects of photosynthesis and calcification on the carbonate system in seawater. Supposing that the rates of photosynthesis and calcification are proportional to concentrations of their inorganic carbon source, the model calculations indicate that three kinds of unique interactions of the organic and inorganic carbon productions are expected. These are photosynthetic enhancement of calcification, calcification which benefits photosynthesis and carbonate dissolution induced by respiration. The first effect appears when the photosynthetic rate is more than approximately 1.2 larger than that of calcification. This effect is caused by the increase of CO3 content and carbonate saturation degree in seawater. If photosynthesis use molecular carbon dioxide, the second effect occurs when the calcification rate is more than approximately 1.6 times larger than that of photosynthesis. Time series model experiments indicate that photosynthesis and calcification potentially enhance each other and that organic and inorganic carbon is produced more efficiently in the coexisting system than in the isolated reactions. These coexisting effects on production enhancement of photosynthesis and calcification are expected to appear not only in the internal pool of organisms but also in a reef environment which is isolated from the outer ocean during low tide. According to the measurements on the fringing type Shiraho Reef in the Ryukyu Islands, the diurnal change of water properties (pH, total alkalinity, total carbon dioxide and carbonate saturation degree) were conspicuous. This environment offers an appropriate condition for the appearance of these coexisting effects. The photosynthetic enhancement of calcification and the respiratory inducement of decalcification were observed during day-time and night-time slack-water periods, respectively. These coexisting effects, especially the photosynthetic enhancement of calcification, appear to play important roles for fluorishing coral reef communities.

Seawater carbonate chemistry and calcification rate in the Bermuda reef community, 2010

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 (CO2) 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 [CO32-] 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 [CO32-] and omega aragonite conditions, and rates of coral calcification due to the photosynthetic uptake of CO2. In late summer, rates of coral calcification are suppressed by release of CO2 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 [CO32-] 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 [CO32-] and omega aragonite thresholds of ~184 micro 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 [CO32-] 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.

Response of a natural phytoplankton community from the Qingdao coast (Yellow Sea, China) to variable CO2 levels over a short-term incubation experiment

Since marine phytoplankton play a vital role in stabilizing earth's climate by removing significant amount of atmospheric CO2, their responses to increasing CO2 levels are indeed vital to address. The responses of a natural phytoplankton community from the Qingdao coast (NW Yellow Sea, China) was studied under different CO2 levels in microcosms. HPLC pigment analysis revealed the presence of diatoms as a dominant microalgal group; however, members of chlorophytes, prasinophytes, cryptophytes and cyanophytes were also present. delta 13CPOM values indicated that the phytoplankton community probably utilized bicarbonate ions as dissolved inorganic carbon source through a carbon concentration mechanism (CCM) under low CO2 levels, and diffusive CO2 uptake increased upon the increase of external CO2 levels. Although, considerable increase in phytoplankton biomass was noticed in all CO2 treatments, CO2-induced effects were absent. Higher net nitrogen uptake under low CO2 levels could be related to the synthesis of CCM components. Flow cytometry analysis showed slight reduction in the abundance of Synechococcus and pico-eukaryotes under the high CO2 treatments. Diatoms did not show any negative impact in response to increasing CO2 levels; however, chlorophytes revealed a reverse tend. Heterotrophic bacterial count enhanced with increasing CO2 levels and indicated higher abundance of labile organic carbon. Thus, the present study indicates that any change in dissolved CO2 concentrations in this area may affect phytoplankton physiology and community structure and needs further long-term study.

Effects of ocean acidification on Posidonia oceanica epiphytic community and shoot productivity

1. Biological interactions can alter predictions that are based on single-species physiological response. It is known that leaf segments of the seagrass Posidonia oceanica will increase photosynthesis with lowered pH, but it is not clear whether the outcome will be altered when the whole plant and its epiphyte community, with different respiratory and photosynthetic demands, are included. In addition, the effects on the Posidonia epiphyte community have rarely been tested under controlled conditions, at near-future pH levels. 2. In order to better evaluate the effects of pH levels as projected for the upcoming decades on seagrass meadows, shoots of P. oceanica with their associated epiphytes were exposed in the laboratory to three pH levels (ambient: 8.1, 7.7 and 7.3, on the total scale) for 4 weeks. Net productivity, respiration, net calcification and leaf fluorescence were measured on several occasions. At the end of the study, epiphyte community abundance and composition, calcareous mass and crustose coralline algae growth were determined. Finally, photosynthesis vs. irradiance curves (PE) was produced from segments of secondary leaves cleaned of epiphytes and pigments extracted. 3. Posidonia leaf fluorescence and chlorophyll concentrations did not differ between pH treatments. Net productivity of entire shoots and epiphyte-free secondary leaves increased significantly at the lowest pH level yet limited or no stimulation in productivity was observed at the intermediate pH treatment. Under both pH treatments, significant decreases in epiphytic cover were observed, mostly due to the reduction of crustose coralline algae. The loss of the dominant epiphyte producer yet similar photosynthetic response for epiphyte-free secondary leaves and shoots suggests a minimal contribution of epiphytes to shoot productivity under experimental conditions. 4. Synthesis. Observed responses indicate that under future ocean acidification conditions foreseen in the next century an increase in Posidonia productivity is not likely despite the partial loss of epiphytic coralline algae which are competitors for light. A decline in epiphytic cover could, however, reduce the feeding capacity of the meadow for invertebrates. In situ long-term experiments that consider both acidification and warming scenarios are needed to improve ecosystem-level predictions.

Shallow water marine sediment bacterial community shifts along a natural CO2 gradient in the Mediterranean Sea Off vulcano, Italy

The effects of increasing atmospheric CO(2) on ocean ecosystems are a major environmental concern, as rapid shoaling of the carbonate saturation horizon is exposing vast areas of marine sediments to corrosive waters worldwide. Natural CO(2) gradients off Vulcano, Italy, have revealed profound ecosystem changes along rocky shore habitats as carbonate saturation levels decrease, but no investigations have yet been made of the sedimentary habitat. Here, we sampled the upper 2 cm of volcanic sand in three zones, ambient (median pCO(2) 419 µatm, minimum Omega (arag) 3.77), moderately CO(2)-enriched (median pCO(2) 592 µatm, minimum Omega (arag) 2.96), and highly CO(2)-enriched (median pCO(2) 1611 µatm, minimum Omega (arag) 0.35). We tested the hypothesis that increasing levels of seawater pCO(2) would cause significant shifts in sediment bacterial community composition, as shown recently in epilithic biofilms at the study site. In this study, 454 pyrosequencing of the V1 to V3 region of the 16S rRNA gene revealed a shift in community composition with increasing pCO(2). The relative abundances of most of the dominant genera were unaffected by the pCO(2) gradient, although there were significant differences for some 5 % of the genera present (viz. Georgenia, Lutibacter, Photobacterium, Acinetobacter, and Paenibacillus), and Shannon Diversity was greatest in sediments subject to long-term acidification (>100 years). Overall, this supports the view that globally increased ocean pCO(2) will be associated with changes in sediment bacterial community composition but that most of these organisms are resilient. However, further work is required to assess whether these results apply to other types of coastal sediments and whether the changes in relative abundance of bacterial taxa that we observed can significantly alter the biogeochemical functions of marine sediments.

Short- and long-term conditioning of a temperate marine diatom community to acidification and warming

Ocean acidification and greenhouse warming will interactively influence competitive success of key phytoplankton groups such as diatoms, but how long-term responses to global change will affect community structure is unknown. We incubated a mixed natural diatom community from coastal New Zealand waters in a short-term (two-week) incubation experiment using a factorial matrix of warming and/or elevated pCO2 and measured effects on community structure. We then isolated the dominant diatoms in clonal cultures and conditioned them for 1 year under the same temperature and pCO2 conditions from which they were isolated, in order to allow for extended selection or acclimation by these abiotic environmental change factors in the absence of interspecific interactions. These conditioned isolates were then recombined into 'artificial' communities modelled after the original natural assemblage and allowed to compete under conditions identical to those in the short-term natural community experiment. In general, the resulting structure of both the unconditioned natural community and conditioned 'artificial' community experiments was similar, despite differences such as the loss of two species in the latter. pCO2 and temperature had both individual and interactive effects on community structure, but temperature was more influential, as warming significantly reduced species richness. In this case, our short-term manipulative experiment with a mixed natural assemblage spanning weeks served as a reasonable proxy to predict the effects of global change forcing on diatom community structure after the component species were conditioned in isolation over an extended timescale. Future studies will be required to assess whether or not this is also the case for other types of algal communities from other marine regimes.

Effect of acidification on an Arctic phytoplankton community from Disko Bay, West Greenland

Long-term measurements (i.e. months) of in situ pH have not previously been reported from the Arctic; this study shows fluctuations between pH 7.5 and 8.3 during the spring bloom 2012 in a coastal area of Disko Bay, West Greenland. The effect of acidification on phytoplankton from this area was studied at both the community and species level in experimental pH treatments within (pH 8.0, 7.7 and 7.4) and outside (pH 7.1) in situ pH. The growth rate of the phytoplankton community decreased during the experimental acidification from 0.50 ± 0.01/day (SD) at pH 8.0 to 0.22 ± 0.01/day at pH 7.1. Nevertheless, the response to acidification was species-specific and divided into 4 categories: I, least affected; II, affected only at pH 7.1; III, gradually affected and IV, highly affected. In addition, the colony size and chain length of selected species were affected by the acidification. Our findings show that coastal phytoplankton from Disko Bay is naturally exposed to pH fluctuations exceeding the experimental pH range used in most ocean acidification studies. We emphasize that studies on ocean acidification should include in situ pH before assumptions on the effect of acidification on marine organisms can be made.