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Response of benthic foraminifera to ocean acidification in their natural sediment environment: a long-term culturing experiment

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 6 months of incubation, foraminiferal assemblages were kept and treated in natural sediment 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 of incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Omega calc was much higher in the sediment than in the water column in nearly all pCO2 treatments and remained close to saturation. As a result, the life cycle (population density, growth and reproduction) of living assemblages varied markedly during the experimental period, but was largely unaffected by the pCO2 treatments applied. According to the size-frequency distribution, we conclude that foraminifera start reproduction at a diameter of 250 µm. Mortality of living Ammonia aomoriensis was unaffected, whereas size of large and dead tests decreased with elevated pCO2 from 285 µm (pCO2 from 430 to 1865 µatm) to 258 µm (pCO2 3247 µatm). The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of CaCO3 weight. Living individuals had a calcium carbonate production rate of 0.47 g/m**2/a, whereas dead empty tests accumulated a rate of 0.27 g /m**2/a. Although Omega calc was close to 1, approximately 30% of the empty tests of Ammonia aomoriensis showed dissolution features at high pCO2 of 3247 µatm during the last 2 months of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. Our results emphasize that the sensitivity to ocean acidification of the endobenthic foraminifera Ammonia aomoriensis in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

The influence of seawater pH on U/ Ca ratios in the scleractinian cold-water coral Lophelia pertusa

The increasing pCO2 in seawater is a serious threat for marine calcifiers and alters the biogeochemistry of the ocean. Therefore, the reconstruction of past-seawater properties and their impact on marine ecosystems is an important way to investigate the underlying mechanisms and to better constrain the effects of possible changes in the future ocean. Cold-water coral (CWC) ecosystems are biodiversity hotspots. Living close to aragonite undersaturation, these corals serve as living laboratories as well as archives to reconstruct the boundary conditions of their calcification under the carbonate system of the ocean. We investigated the reef-building CWC Lophelia pertusa as a recorder of intermediate ocean seawater pH. This species-specific field calibration is based on a unique sample set of live in situ collected L. pertusa and corresponding seawater samples. These data demonstrate that uranium speciation and skeletal incorporation for azooxanthellate scleractinian CWCs is pH dependent and can be reconstructed with an uncertainty of ±0.15. Our Lophelia U / Ca-pH calibration appears to be controlled by the high pH values and thus highlighting the need for future coral and seawater sampling to refine this relationship. However, this study recommends L. pertusa as a new archive for the reconstruction of intermediate water mass pH and hence may help to constrain tipping points for ecosystem dynamics and evolutionary characteristics in a changing ocean.

Seawater carbonate chemistry and distribution of coastal benthic foraminifera

Coastal ecosystems are subjected to both large natural variability and increasing anthropogenic impact on environmental parameters such as changes in salinity, temperature, and pH. This study documents the distribution of living benthic foraminifera under the influence of multiple environmental stressors in the Skagerrak-Baltic Sea region. Sediment core tops were studied at five sites along a transect from the Skagerrak to the Baltic Sea, with strong environmental gradients, especially in terms of salinity, pH, calcium carbonate saturation and dissolved oxygen concentration in the bottom water and pore water. We found that living foraminiferal densities and species richness were higher at the Skagerrak station, where the general living conditions were relatively beneficial for Foraminifera, with higher salinity and Ωcalc in the water column and higher pH and oxygen concentration in the bottom and pore water. The most common species reported at each station reflect the differences in the environmental conditions between the stations. The dominant species were Cassidulina laevigata and Hyalinea balthica in the Skagerrak, Stainforthia fusiformis, Nonionella aff. stella and Nonionoides turgida in the Kattegat and N. aff. stella and Nonionellina labradorica in the Öresund. The most adverse conditions, such as low salinity, low Ωcalc, low dissolved oxygen concentrations and low pH, were noted at the Baltic Sea stations, where the calcareous tests of the dominant living taxa Ammonia spp. and Elphidium spp. were partially to completely dissolved, probably due to a combination of different stressors affecting the required energy for biomineralization. Even though Foraminifera are able to live in extremely varying environmental conditions, the present results suggest that the benthic coastal ecosystems in the studied region, which are apparently affected by an increase in the range of environmental variability, will probably be even more influenced by a future increase in anthropogenic impacts, including coastal ocean acidification and deoxygenation.

Seawater carbonate chemistry and carbonate load of seagrass leaves

Seagrass meadows play a significant role in the formation of carbonate sediments, serving as a substrate for carbonate-producing epiphyte communities. The magnitude of the epiphyte load depends on plant structural and physiological parameters, related to the time available for epiphyte colonization. Yet, the carbonate accumulation is likely to also depend on the carbonate saturation state of seawater (Omega) that tends to decrease as latitude increases due to decreasing temperature and salinity. A decrease in carbonate accumulation with increasing latitude has already been demonstrated for other carbonate producing communities. The aim of this study was to assess whether there was any correlation between latitude and the epiphyte carbonate load and net carbonate production rate on seagrass leaves. Shoots from 8 different meadows of the Zostera genus distributed across a broad latitudinal range (27 °S to up to 64 °N) were sampled along with measurements of temperature and Omega. The Omega within meadows significantly decreased as latitude increased and temperature decreased. The mean carbonate content and load on seagrass leaves ranged from 17 % DW to 36 % DW and 0.4-2.3 mg CO3/cm**2, respectively, and the associated mean carbonate net production rate varied from 0.007 to 0.9 mg CO3/cm**2/d. Mean carbonate load and net production rates decreased from subtropical and tropical, warmer regions towards subpolar latitudes, consistent with the decrease in Omega. These results point to a latitudinal variation in the contribution of seagrass to the accumulation of carbonates in their sediments which affect important processes occurring in seagrass meadows, such as nutrient cycling, carbon sequestration and sediment accretion.

The benthic foraminiferal community in a naturally CO2-rich coastal habitat of the southwestern Baltic Sea

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 a consequence of higher alkalinity (AT) for most 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 characterised 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 (Omega calc) co-vary with the population densities of living Ammonia aomoriensis. Further, we observed that most of the tests of living calcifying foraminifera were intact. Only Ammonia aomorienis showed dissolution and recalcification structures on the tests, especially at undersaturated conditions. Therefore, the benthic community is subjected to 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 Omega calc levels.

Ecosystem functions of rare arable plants - field study: Araneae data

Partly taken from the materials and methods of https://doi.org/10.1016/j.baae.2022.12.003: To compare the activity densities of ground-dwelling predators between treatments with and without RAPs, spiders were sampled using pitfall traps, which were set up after each round of aphid counting (one per plot, twice per year; Brown & Matthews, 2016). The traps (with a volume of 400 ml and a width of 90 mm) were filled with a mixture of water and ethylene glycol (1:1; 120 ml) and dug at ground level into the middle of each plot. The traps were covered with a plastic roof and a metal grid (15 × 15 mm grid size) to avoid overflowing during rain and accidental rodent catches (Császár et al., 2018). The traps were activated for 7 days. Subsequently, all arthropods were transferred into 70% ethanol. Spiders were identified to species according to Nentwig et al. (2019). Spider hunting strategy (active hunter or web-builder) was used as the feeding trait according to Cardoso et al. (2011).

Ecosystem functions of rare arable plants - field study: Aphidoidea data

Taken from materials and methods of https://doi.org/10.1016/j.baae.2022.12.003: Aphids were counted on 50 randomly selected shoots in two crop rows (100 shoots in total) per plot and sampling round. To reduce edge effects, rows with less than 20 cm to the edge were excluded. Counting took place twice a year, that is, once during crop flowering (BBCH 61; beginning of aphid population growth) and once during crop milk ripening stage (BBCH 75).

Ecosystem functions of rare arable plants - field study: Soil data

Taken from the methods of https://doi.org/10.1016/j.agee.2020.107237: The effect of rare arable plants on soil nutrient concentration was measured by taking soil samples in the 1st and 2nd study year (March 2018 and August 2019). One soil sample per plot was taken to a 20 cm depth and analyzed by the AGROLAB Group (Landshut, Germany) for soil organic matter [%] and nitrogen concentration [%] (DIN EN 15936; 2012 and DIN EN 16168; 2012-11).

Ecosystem functions of rare arable plants - field study: Carabidae data

Partly taken from the materials and methods of https://doi.org/10.1016/j.baae.2022.12.003: To compare the activity densities of ground-dwelling predators between treatments with and without RAPs, carabids were sampled using pitfall traps, which were set up after each round of aphid counting (one per plot, twice per year; Brown & Matthews, 2016). The traps (with a volume of 400 ml and a width of 90 mm) were filled with a mixture of water and ethylene glycol (1:1; 120 ml) and dug at ground level into the middle of each plot. The traps were covered with a plastic roof and a metal grid (15 × 15 mm grid size) to avoid overflowing during rain and accidental rodent catches (Császár et al., 2018). The traps were activated for 7 days. Subsequently, all arthropods were transferred into 70% ethanol. Carabids were identified to species according to Hůrka (1996). Carabid feeding behavior was classified according to Homburg et al. (2014). To simplify the dataset, carabid feeding behavior was classified as predominantly granivorous (species mainly feed on seeds and fruits) or as carnivorous/omnivorous, because carnivorous and omnivorous species are potentially feeding on aphids and other non-plant material.

Ecosystem functions of rare arable plants - field study: Plants biomass data

Taken from the methods of https://doi.org/10.1016/j.agee.2020.107237: Vegetation surveys were performed once in July for both study years. Plant species were classified and each species' percent cover for both the arable plant community and the crop were visually estimated per plot. Species were divided into rare arable plants, other spontaneously occurring arable plants, the sown crop species, and volunteer crops that re-emerged after cultivation in previous years. To measure the productivity of our plots, both the crop biomass and arable plant biomass (rare arable plants + volunteer crops + spontaneous arable plants) were collected in July and August in both study years. For the arable plant biomass, three 0.5 m × 0.5 m sampling quadrats were randomly placed in the plot, harvested, and dried at 65 °C for 48 h. Crop biomass was measured after cutting, drying, and weighing three randomly selected crop rows per plot. To minimize edge effects, the outmost crop rows were excluded from the sampling. Arable plants and crop biomasses were projected as g m ⁻².

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