Description: It is expected that the calcification of foraminifera will be negatively affected by the ongoing acidification of the oceans. Compared to the open oceans, these organisms are subjected to much more adverse carbonate system conditions in coastal and estuarine environments such as the southwestern Baltic Sea, where benthic foraminifera are abundant. This study documents the seasonal changes of carbonate chemistry and the ensuing response of the foraminiferal community with bi-monthly resolution in Flensburg Fjord. In comparison to the surface pCO2, which is close to equilibrium with the atmosphere, we observed large seasonal fluctuations of pCO2 in the bottom and sediment pore waters. The sediment pore water pCO2 was constantly high during the entire year ranging from 1244 to 3324 μatm. Nevertheless, in contrast to the bottom water, sediment pore water was slightly supersaturated with respect to calcite as consequence of higher alkalinity (AT) for the most time of the year. Foraminiferal assemblages were dominated by two calcareous species, Ammonia aomoriensis and Elphidium incertum, and the agglutinated Ammotium cassis. The one year-cycle was characterized by seasonal community shifts. Our results revealed that there is no dynamic response of foraminiferal population density and diversity to elevated sediment pore water pCO2. Surprisingly, the fluctuations of sediment pore water undersaturation (Ωcalc) co-vary with the population densities of living Ammonia aomoriensis. Further, we observed that most of the tests of living calcifying specimens were intact. Only Ammonia aomorienis showed dissolution and recalcification structures on the tests, especially at undersaturated conditions. Therefore, the benthic community is subjected to constantly high pCO2 and tolerates elevated levels as long as sediment pore water remains supersaturated. Model calculations inferred that increasing atmospheric CO2 concentrations will finally lead to a perennial undersaturation in sediment pore waters. Whereas benthic foraminifera indeed may cope with a high sediment pore water pCO2, the steady undersaturation of sediment pore waters would likely cause a significant higher mortality of the dominating Ammonia aomoriensis. This shift may eventually lead to changes in the benthic foraminiferal communities in Flensburg Fjord, as well as in other regions experiencing naturally undersaturated Ωcalc levels.

Description: Calcifying foraminifera are expected to be endangered by ocean acidification, However, the response of a complete community kept in natural sediment and over multiple generations under controlled laboratory conditions has not been constrained to date. During 5 six month incubation, foraminiferal assemblages were treated with pCO2 enriched seawater of 430, 907, 1865 and 3247 μatm pCO2. The fauna was dominated by Ammonia aomoriensis and Elphidium species, whereas agglutinated species were rare. After 6 months incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Òcalc was much higher in the sedi10 ment than in the water column in all pCO2 treatments and remained close to saturation. As a result, the life cycle of living assemblages was largely unaffected by the tested pCO2 treatments. Growth rates, reproduction and mortality, and therefore population densities and size-frequency distribution of Ammonia aomoriensis varied markedly during the experimental period. Growth rates varied between 25 and 50 μm per month, 15 which corresponds to an addition of 1 or 2 new chambers per month. According to the size-frequency distribution, foraminifera start reproduction at a diameter of 250 μm. Mortality of large foraminifera was recognized, commencing at a test size of 285 μm at a pCO2 ranging from 430 to 1865 μatm, and of 258 μm at 3247 μatm. The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of dry 20 weight. Living individuals had a calcium carbonate production rate of 0.47 gm−2 yr−1, whereas dead empty tests accumulated at a rate of 0.27 gm−2a−1. Although Òcalc was close to 1, some empty tests of Ammonia aomoriensis showed dissolution features at the end of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. This species specific response could be explained by differences in 25 the elemental test composition, in particular the higher Mg-concentrations in Ammonia aomoriensis tests. Our results emphasize that the sensitivity to ocean acidification of endobenthic foraminifera in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

Description: Bivalve calcification, particularly of the early larval stages, is highly sensitive to the change in ocean carbonate chemistry resulting from atmospheric CO2 uptake. Earlier studies suggested that declining seawater [CO32−] and thereby lowered carbonate saturation affect shell production. However, disturbances of physiological processes such as acid-base regulation by adverse seawater pCO2 and pH can affect calcification in a secondary fashion. In order to determine the exact carbonate system component by which growth and calcification are affected it is necessary to utilize more complex carbonate chemistry manipulations. As single factors, pCO2 had no effects and [HCO3-] and pH had only limited effects on shell growth, while lowered [CO32−] strongly impacted calcification. Dissolved inorganic carbon (CT) limiting conditions led to strong reductions in calcification, despite high [CO32−], indicating that [HCO3-] rather than [CO32−] is the inorganic carbon source utilized for calcification by mytilid mussels. However, as the ratio [HCO3-] / [H+] is linearly correlated with [CO32−] it is not possible to differentiate between these under natural seawater conditions. An equivalent of about 80 μmol kg−1 [CO32−] is required to saturate inorganic carbon supply for calcification in bivalves. Below this threshold biomineralization rates rapidly decline. A comparison of literature data available for larvae and juvenile mussels and oysters originating from habitats differing substantially with respect to prevailing carbonate chemistry conditions revealed similar response curves. This suggests that the mechanisms which determine sensitivity of calcification in this group are highly conserved. The higher sensitivity of larval calcification seems to primarily result from the much higher relative calcification rates in early life stages. In order to reveal and understand the mechanisms that limit or facilitate adaptation to future ocean acidification, it is necessary to better understand the physiological processes and their underlying genetics that govern inorganic carbon assimilation for calcification.