Abundance, Electron transport System (ETS) activity and respiration rates of pelagic decapods from the Benguela upwelling system during AFR258, D356 and MSM17/3

Decapods were sampled with a 1 m**2 MOCNESS (mainly upper 1000 m) in the northern Benguela Current during three cruises in December 2009, September/October 2010 and February 2011. Although pelagic decapods are abundant members of the micronekton community, information about their ecophysiology is very limited. Species-specific regional distribution limits were detected for various decapod species (e.g. Plesionika carinata, Sergestes arcticus, Pasiphaea semispinosa). Significant diel vertical migration patterns were determined for three caridean and three penaeiodean species. Biomass was variable and ranged from 23 to 2770 mg dry mass m**-2 with highest values for P. semispinosa. Fatty acid and stable isotope analyses revealed that the examined decapod species are omnivorous tocarnivorous except for the herbivorous to omnivorous species P. carinata. Calanid copepods such as Calanoides carinatus were identified as an important prey item especially for caridean species. Community consumption rates of pelagic decapods derived from respiration rates ranged from 7 mg C m**-2 d**-1 (231S) to 420 mg C m**-2 d**-1 (191S, 171S). A potential active respiratory carbon flux was calculated for migrating pelagic decapods with 4.4 mg C m**- d**-1 for the upper 200 m and with 2.6 mg C m**-2 d**-1 from the base of the euphotic zone to a depth of 600 m. Overall, pelagic decapods apparently play a more prominent role in the northern Benguela Current ecosystem than previously assumed and may exert a substantial predation impact on calanid copepods (up to 13% d**-1 of standing stock).

Carbonate chemistry, community metabolism, PAR, temperature and salinity of One Tree Island reef

There are few in situ studies showing how net community calcification (Gnet) of coral reefs is related to carbonate chemistry, and the studies to date have demonstrated different predicted rates of change. In this study, we measured net community production (Pnet), Gnet, and carbonate chemistry of a reef flat at One Tree Island, Great Barrier Reef. Diurnal pCO2 variability of 289-724 µatm was driven primarily by photosynthesis and respiration. The reef flat was found to be net autotrophic, with daily production of ? 35 mmol C/m**2/d and net calcification of ? 33 mmol C/m**2/d . Gnet was strongly related to Pnet, which drove a hysteresis pattern in the relationship between Gnet and aragonite saturation state (Omega ar). Although Pnet was the main driver of Gnet, Omega ar was still an important factor, where 95% of the variance in Gnet could be described by Pnet and Omega ar. Based on the observed in situ relationship, Gnet would be expected to reach zero when Omega ar is 2.5. It is unknown what proportion of a decline in Gnet would be through reduced calcification and what would occur through increased dissolution, but the results here support predictions that overall calcium carbonate production will decline in coral reefs as a result of ocean acidification.

Effects of elevated CO2 and temperature on an intertidal meiobenthic community

In the near future, the marine environment is likely to be subjected to simultaneous increases in temperature and decreased pH. The potential effects of these changes on intertidal, meiofaunal assemblages were investigated using a mesocosm experiment. Artificial Substrate Units containing meiofauna from the extreme low intertidal zone were exposed for 60 days to eight experimental treatments (four replicates for each treatment) comprising four pH levels: 8.0 (ambient control), 7.7 & 7.3 (predicted changes associated with ocean acidification), and 6.7 (CO2 point-source leakage from geological storage), crossed with two temperatures: 12 °C (ambient control) and 16 °C (predicted). Community structure, measured using major meiofauna taxa was significantly affected by pH and temperature. Copepods and copepodites showed the greatest decline in abundance in response to low pH and elevated temperature. Nematodes increased in abundance in response to low pH and temperature rise, possibly caused by decreased predation and competition for food owing to the declining macrofauna density. Nematode species composition changed significantly between the different treatments, and was affected by both seawater acidification and warming. Estimated nematode species diversity, species evenness, and the maturity index, were substantially lower at 16 °C, whereas trophic diversity was slightly higher at 16 °C except at pH 6.7. This study has demonstrated that the combination of elevated levels of CO2 and ocean warming may have substantial effects on structural and functional characteristics of meiofaunal and nematode communities, and that single stressor experiments are unlikely to encompass the complexity of abiotic and biotic interactions. At the same time, ecological interactions may lead to complex community responses to pH and temperature changes in the interstitial environment.

Spatial community shift from hard to soft corals in acidified water

Anthropogenic increases in the partial pressure of CO2 (pCO2) cause ocean acidification, declining calcium carbonate saturation states, reduced coral reef calcification and changes in the compositions of marine communities. Most projected community changes due to ocean acidification describe transitions from hard coral to non-calcifying macroalgal communities; other organisms have received less attention, despite the biotic diversity of coral reef communities. We show that the spatial distributions of both hard and soft coral communities in volcanically acidified, semi-enclosed waters off Iwotorishima Island, Japan, are related to pCO2 levels. Hard corals are restricted to non-acidified low- pCO2 (225 µatm) zones, dense populations of the soft coral Sarcophyton elegans dominate medium- pCO2 (831 µatm) zones, and both hard and soft corals are absent from the highest- pCO2 (1,465 µatm) zone. In CO2-enriched culture experiments, high- pCO2 conditions benefited Sarcophyton elegans by enhancing photosynthesis rates and did not affect light calcification, but dark decalcification (negative net calcification) increased with increasing pCO2. These results suggest that reef communities may shift from reef-building hard corals to non-reef-building soft corals under pCO2 levels (550-970 µatm) predicted by the end of this century, and that higher pCO2 levels would challenge the survival of some reef organisms.

Coral-algae metabolism and diurnal changes in the CO2-carbonate system of bulk sea water

Precise measurements were conducted in continuous flow seawater mesocosms located in full sunlight that compared metabolic response of coral, coral-macroalgae and macroalgae systems over a diurnal cycle. Irradiance controlled net photosynthesis (Pnet), which in turn drove net calcification (Gnet), and altered pH. Pnet exerted the dominant control on [CO3]2- and aragonite saturation state (Omega arag) over the diel cycle. Dark calcification rate decreased after sunset, reaching zero near midnight followed by an increasing rate that peaked at 03:00 h. Changes in Omega arag and pH lagged behind Gnet throughout the daily cycle by two or more hours. The flux rate Pnet was the primary driver of calcification. Daytime coral metabolism rapidly removes dissolved inorganic carbon (DIC) from the bulk seawater and photosynthesis provides the energy that drives Gnet while increasing the bulk water pH. These relationships result in a correlation between Gnet and Omega arag, with Omega arag as the dependent variable. High rates of H+ efflux continued for several hours following mid-day peak Gnet suggesting that corals have difficulty in shedding waste protons as described by the Proton Flux Hypothesis. DIC flux (uptake) followed Pnet and Gnet and dropped off rapidly following peak Pnet and peak Gnet indicating that corals can cope more effectively with the problem of limited DIC supply compared to the problem of eliminating H+. Over a 24 h period the plot of total alkalinity (AT) versus DIC as well as the plot of Gnet versus Omega arag revealed a circular hysteresis pattern over the diel cycle in the coral and coral-algae mesocosms, but not the macroalgae mesocosm. Presence of macroalgae did not change Gnet of the corals, but altered the relationship between Omega arag and Gnet. Predictive models of how future global changes will effect coral growth that are based on oceanic Omega arag must include the influence of future localized Pnet on Gnet and changes in rate of reef carbonate dissolution. The correlation between Omega arag and Gnet over the diel cycle is simply the response of the CO2-carbonate system to increased pH as photosynthesis shifts the equilibria and increases the [CO3]2- relative to the other DIC components of [HCO3]- and [CO2]. Therefore Omega arag closely tracked pH as an effect of changes in Pnet, which also drove changes in Gnet. Measurements of DIC flux and H+ flux are far more useful than concentrations in describing coral metabolism dynamics. Coral reefs are systems that exist in constant disequilibrium with the water column.

Seawater carbonate chemistry, community calcification and photosynthesis during experiments with coral reefs at Shiraho reef, Ishigaki Island, southwest Japan, 2009

Seven coral reef communities were defined on Shiraho fringing reef, Ishigaki Island, Japan. Net photosynthesis and calcification rates were measured by in situ incubations at 10 sites that included six of the defined communities, and which occupied most of the area on the reef flat and slope. Net photosynthesis on the reef flat was positive overall, but the reef flat acts as a source for atmospheric CO2, because the measured calcification/photosynthesis ratio of 2.5 is greater than the critical ratio of 1.67. Net photosynthesis on the reef slope was negative. Almost all excess organic production from the reef flat is expected to be effused to the outer reef and consumed by the communities there. Therefore, the total net organic production of the whole reef system is probably almost zero and the whole reef system also acts as a source for atmospheric CO2. Net calcification rates of the reef slope corals were much lower than those of the branching corals. The accumulation rate of the former was approximately 0.5 m kyr?1 and of the latter was ~0.7-5 m kyr?1. Consequently, reef slope corals could not grow fast enough to keep up with or catch up to rising sea levels during the Holocene. On the other hand, the branching corals grow fast enough to keep up with this rising sea level. Therefore, a transition between early Holocene and present-day reef communities is expected. Branching coral communities would have dominated while reef growth kept pace with sea level rise, and the reef was constructed with a branching coral framework. Then, the outside of this framework was covered and built up by reef slope corals and present-day reefs were constructed.

Seawater carbonate chemistry, nutrients, growth rate and phytoplnkton community response to increasing CO2 concentration during experiments, 2011

This paper reports for the first time upon the effects of increasing CO2 concentrations on a natural phytoplankton assemblage in a tropical estuary (the Godavari River Estuary in India). Two short-term (5-day) bottle experiments were conducted (with and without nutrient addition) during the pre-monsoon season when the partial pressure of CO2 in the surface water is quite low. The results reveal that the concentrations of total chlorophyll, the phytoplankton growth rate, the concentrations of particulate organic matter, the photosynthetic oxygen evolution rates, and the total bacterial count were higher under elevated CO2 treatments, as compared to ambient conditions (control). delta13C of particulate organic matter (POM) varied inversely with respect to CO2, indicating a clear signature of higher CO2 influx under the elevated CO2 levels. Whereas, delta13CPOM in the controls indicated the existence of an active bicarbonate transport system under limited CO2 supply. A considerable change in phytoplankton community structure was noticed, with marker pigment analysis by HPLC revealing that cyanobacteria were dominant over diatoms as CO2 concentrations increased. A mass balance calculation indicated that insufficient nutrients (N, P and Si) might have inhibited diatomgrowth compared to cyanobacteria, regardless of increased CO2 supply. The present study suggests that CO2 concentration and nutrient supply could have significant effects on phytoplankton physiology and community composition for natural phytoplankton communities in this region. However, this work was conducted during a non-discharge period (nutrient-limited conditions) and the responses of phytoplankton to increasing CO2 might not necessarily be the same during other seasons with high physicochemical variability. Further investigation is therefore needed.

Seawater carbonate chemistry in Hog reef 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.

Seawater carbonate chemistry and community calcification during Shiraha coral reef (Ishigaki Island, Japan) studies, 1995

Only about half of all the CO_2 that has been produced by the burning of fossil fuels now remains in the atmosphere. The CO_2 "missing" from the atmosphere is the subject of an important debate. It was thought that the great majority of the missing CO_2 has invaded the ocean, for this system naturally acts as a giant chemical regulator of the atmosphere. Although it is clear that ocean processes have a major role in the regulation of the carbon dioxide content of the atmosphere through air-sea exchange processes, recent studies of the oceanic carbon cycle and air-sea interaction indicate that oceanic carbon is in a quasi-steady state via the system of biological and physical processes in the ocean interior. It is difficult to determine whether the ocean has the capacity to take up the increasing air-born CO_2 released by human activities over the past five or six decades. To understand this enigma, we need a better understanding of the natural variability of the oceanic carbon cycle.

Seawater carbonate chemistry, stable carbon isotope fractionation and growth rate during experiments with marine phytoplankton community, 1999

Stable carbon isotope fractionation (%) of 7 marine phytoplankton species grown in different irradiance cycles was measured under nutrient-replete conditions at a high light intensity in batch cultures. Compared to experiments under continuous light, all species exhibited a significantly higher instantaneous growth rate (pi), defined as the rate of carbon fixation during the photo period, when cultivated at 12:12 h. 16:8 h, or 186 h light:dark (L/D) cycles. Isotopic fractionation by the diatoms Skeletonema costatum, Asterionella glacialis, Thalassiosira punctigera, and Coscinodiscus wailesii (Group I) was 4 to 6% lower in a 16:8 h L/D cycle than under continuous light, which we attribute to differences in pi. In contrast, E, in Phaeodactylum tn'cornutum, Thalassiosira weissflogii, and in the dinoflagellate Scrippsiella trochoidea (Group 11) was largely insensitive to day length-related differences in instantaneous growth rate. Since other studies have reported growth-rate dependent fractionation under N-limited conditions in P. tricornutum, pi-related effects on fractionation apparently depend on the factor controlling growth rate. We suggest that a general relationship between E, and pi/[C02,,,] may not exist. For 1 species of each group we tested the effect of variable CO2 concentration, [COz,,,], on isotopic fractionation. A decrease in [CO2,,,] from ca 26 to 3 pm01 kg-' caused a decrease in E, by less than 3%0 This indicates that variation in h in response to changes in day length has a similar or even greater effect on isotopic fractionation than [COz,,,] m some of the species tested. In both groups E, tended to be higher in smaller species at comparable growth rates. In 24 and 48 h time series the algal cells became progressively enriched in 13C during the day and the first hours of the dark period, followed by l3C depletion in the 2 h before beginning of the following Light period. The daily amplitude of the algal isotopic composition (613C), however, was <1.5%0, which demonstrates that diurnal variation in Fl3C is relatively small.

Seawater carbonate chemistry and phytoplankton and eubacterial community compositions in the northwest subarctic Pacific

On-deck CO2-Fe-manipulated incubation experiments were conducted using surface seawater collected from the Western Subarctic Gyre of the NW Pacific in the summer of 2008 to elucidate the impacts of ocean acidification and Fe enrichment on the abundance and community composition of phytoplankton and eubacteria in the study area. During the incubation, excluding the initial period, the mean partial pressures of CO2 in non-Fe-added bottles were 230, 419, 843, and 1124 µatm, whereas those in Fe-added treatments were 152, 394, 791, and 1008 µatm. Changes in the abundance and community composition of phytoplankton were estimated using HPLC pigment signatures with the program CHEMTAX and flow cytometry. A DGGE fingerprint technique targeting 16S rRNA gene fragments was also used to estimate changes in eubacterial phylotypes during incubation. The Fe addition induced diatom blooms, and subsequently stimulated the growth of heterotrophic bacteria such as Roseobacter, Phaeobacter, and Alteromonas in the post-bloom phase. In both the Fe-limited and Fe-replete treatments, concentrations of 19'-hexanoyloxyfucoxanthin, a haptophyte marker, and the cell abundance of coccolithophores decreased at higher CO2 levels (750 and 1000 ppm), whereas diatoms exhibited little response to the changes in CO2 availability. The abundances of Synechococcus and small eukaryotic phytoplankton (<10 µm) increased at the higher CO2 levels. DGGE band positions revealed that Methylobacterium of Alphaproteobacteria occurred solely at lower CO2 levels (180 and 380 ppm) during the post-bloom phase. These results suggest that increases in CO2 level could affect not only the community composition of phytoplankton but also that of eubacteria. As these microorganisms play critical roles in the biological carbon pump and microbial loop, our results indicate that the progression of ocean acidification can alter the biogeochemical processes in the study area.

Seawater carbonate chemistry, nitrogen concentration and macro community analyse, 2011

Rising anthropogenic CO2 emissions acidify the oceans, and cause changes to seawater carbon chemistry. Bacterial biofilm communities reflect environmental disturbances and may rapidly respond to ocean acidification. This study investigates community composition and activity responses to experimental ocean acidification in biofilms from the Australian Great Barrier Reef. Natural biofilms grown on glass slides were exposed for 11 d to four controlled pCO2 concentrations representing the following scenarios: A) pre-industrial (~300 ppm), B) present-day (~400 ppm), C) mid century (~560 ppm) and D) late century (~1140 ppm). Terminal restriction fragment length polymorphism and clone library analyses of 16S rRNA genes revealed CO2-correlated bacterial community shifts between treatments A, B and D. Observed bacterial community shifts were driven by decreases in the relative abundance of Alphaproteobacteria and increases of Flavobacteriales (Bacteroidetes) at increased CO2 concentrations, indicating pH sensitivity of specific bacterial groups. Elevated pCO2 (C + D) shifted biofilm algal communities and significantly increased C and N contents, yet O2 fluxes, measured using in light and dark incubations, remained unchanged. Our findings suggest that bacterial biofilm communities rapidly adapt and reorganize in response to high pCO2 to maintain activity such as oxygen production.

Effects of ocean acidification on population dynamics and community structure of crustose coralline algae

Calcification and growth of crustose coralline algae (CCA) are affected by elevated seawater pCO2 and associated changes in carbonate chemistry. However, the effects of ocean acidification (OA) on population and community-level responses of CCA have barely been investigated. We explored changes in community structure and population dynamics (size structure and reproduction) of CCA in response to OA. Recruited from an experimental flow-through system, CCA settled onto the walls of plastic aquaria and developed under exposure to one of three pCO2 treatments (control [present day, 389±6 ppm CO2], medium [753±11 ppm], and high [1267±19 ppm]). Elevated pCO2 reduced total CCA abundance and affected community structure, in particular the density of the dominant species Pneophyllum sp. and Porolithon onkodes. Meanwhile, the relative abundance of P. onkodes declined from 24% under control CO2 to 8.3% in high CO2 (65% change), while the relative abundance of Pneophyllum sp. remained constant. Population size structure of P. onkodes differed significantly across treatments, with fewer larger individuals under high CO2. In contrast, the population size structure and number of reproductive structures (conceptacles) per crust of Pneophyllum sp. was similar across treatments. The difference in the magnitude of the response of species abundance and population size structure between species may have the potential to induce species composition changes in the future. These results demonstrate that the impacts of OA on key coral reef builders go beyond declines in calcification and growth, and suggest important changes to aspects of population dynamics and community ecology.

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.

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.