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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.
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.
Anthropogenic CO2 emissions are rapidly changing seawater temperature, pH and carbonate chemistry. This study compares the embryonic development under high pCO2conditions across the south-north distribution range of the marine clam Limecola balthicain NW Europe. The combined effects of elevated temperature and reduced pH on hatching success and size varied strongly between the three studied populations, with the Gulf of Finland population appearing most endangered under the conditions predicted to occur by 2100. These results demonstrate that the assessment of marine faunal population persistence to future climatic conditions needs to consider the interactive effects of co-occurring physico-chemical alterations in seawater within the local context that determines population fitness, adaptation potential and the system resilience to environmental change.
Culturing experiments were performed with the benthic foraminifer Ammonia aomoriensis from Flensburg Fjord, western Baltic Sea. The experiments simulated a projected rise in atmospheric CO2 concentrations. We exposed specimens to 5 seawater pCO2 levels ranging from 618 µatm (pH 7.9) to 3130 µatm (pH 7.2) for 6 wk. Growth rates and mortality differed significantly among pCO2 treatments. The highest increase of mean test diameter (19%) was observed at 618 µatm. At partial pressures >1829 µatm, the mean test diameter was observed to decrease, by up to 22% at 3130 µatm. At pCO2 levels of 618 and 751 µatm, A. aomoriensis tests were found intact after the experiment. The outer chambers of specimens incubated at 929 and 1829 µatm were severely damaged by corrosion. Visual inspection of specimens incubated at 3130 µatm revealed wall dissolution of all outer chambers, only their inner organic lining stayed intact. Our results demonstrate that pCO2 values of >=929 µatm in Baltic Sea waters cause reduced growth of A. aomoriensis and lead to shell dissolution. The bottom waters in Flensburg Fjord and adjacent areas regularly experience pCO2 levels in this range during summer and fall. Increasing atmospheric CO2 concentrations are likely to extend and intensify these periods of undersaturation. This may eventually slow down calcification in A. aomoriensis to the extent that net carbonate precipitation terminates. The possible disappearance of this species from the Baltic Sea and other areas prone to seasonal undersaturation would likely cause significant shifts in shallow-water benthic ecosystems in the near future.
Increased maintenance costs at cellular, and consequently organism level, are thought to be involved in shaping the sensitivity of marine calcifiers to ocean acidification (OA). Yet, knowledge of the capacity of marine calcifiers to undergo metabolic adaptation is sparse. In Kiel Fjord, blue mussels thrive despite periodically high seawater PCO2, making this population interesting for studying metabolic adaptation under OA. Consequently, we conducted a multi-generation experiment and compared physiological responses of F1 mussels from 'tolerant' and 'sensitive' families exposed to OA for 1 year. Family classifications were based on larval survival; tolerant families settled at all PCO2 levels (700, 1120, 2400 µatm) while sensitive families did not settle at the highest PCO2 (>=99.8% mortality). We found similar filtration rates between family types at the control and intermediate PCO2 level. However, at 2400 µatm, filtration and metabolic scope of gill tissue decreased in tolerant families, indicating functional limitations at the tissue level. Routine metabolic rates (RMR) and summed tissue respiration (gill and outer mantle tissue) of tolerant families were increased at intermediate PCO2, indicating elevated cellular homeostatic costs in various tissues. By contrast, OA did not affect tissue and routine metabolism of sensitive families. However, tolerant mussels were characterised by lower RMR at control PCO2 than sensitive families, which had variable RMR. This might provide the energetic scope to cover increased energetic demands under OA, highlighting the importance of analysing intra-population variability. The mechanisms shaping such difference in RMR and scope, and thus species' adaptation potential, remain to be identified.
Phytoplankton populations can display high levels of genetic diversity that, when reflected by phenotypic variability, may stabilize a species response to environmental changes. We studied the effects of increased temperature and CO2 availability as predicted consequences of global change, on 16 genetically different isolates of the diatom Skeletonema marinoi from the Adriatic Sea and the Skagerrak (North Sea), and on eight strains of the PST (paralytic shellfish toxin)-producing dinoflagellate Alexandrium ostenfeldii from the Baltic Sea. Maximum growth rates were estimated in batch cultures of acclimated isolates grown for five to 10 generations in a factorial design at 20 and 24 °C, and present day and next century applied atmospheric pCO2, respectively. In both species, individual strains were affected in different ways by increased temperature and pCO2. The strongest response variability, buffering overall effects, was detected among Adriatic S. marinoi strains. Skagerrak strains showed a more uniform response, particularly to increased temperature, with an overall positive effect on growth. Increased temperature also caused a general growth stimulation in A. ostenfeldii, despite notable variability in strain-specific response patterns. Our data revealed a significant relationship between strain-specific growth rates and the impact of pCO2 on growth-slow growing cultures were generally positively affected, while fast growing cultures showed no or negative responses to increased pCO2. Toxin composition of A. ostenfeldii was consistently altered by elevated temperature and increased CO2 supply in the tested strains, resulting in overall promotion of saxitoxin production by both treatments. Our findings suggest that phenotypic variability within populations plays an important role in the adaptation of phytoplankton to changing environments, potentially attenuating short-term effects and forming the basis for selection. In particular, A. ostenfeldii blooms may expand and increase in toxicity under increased water temperature and atmospheric pCO2 conditions, with potentially severe consequences for the coastal ecosystem.
Seaweeds are key species of the Baltic Sea benthic ecosystems. They are the substratum of numerous fouling epibionts like bryozoans and tubeworms. Several of these epibionts bear calcified structures and could be impacted by the high pCO2 events of the late summer upwellings in the Baltic nearshores. Those events are expected to increase in strength and duration with global change and ocean acidification. If calcifying epibionts are impacted by transient acidification as driven by upwelling events, their increasing prevalence could cause a shift of the fouling communities toward fleshy species. The aim of the present study was to test the sensitivity of selected seaweed macrofoulers to transient elevation of pCO2 in their natural microenvironment, i.e. the boundary layer covering the thallus surface of brown seaweeds. Fragments of the macroalga Fucus serratus bearing an epibiotic community composed of the calcifiers Spirorbis spirorbis (Annelida) and Electra pilosa (Bryozoa) and the non-calcifier Alcyonidium hirsutum (Bryozoa) were maintained for 30 days under three pCO2 conditions: natural 460±59 µatm, present-day upwelling1193±166 µatm and future upwelling 3150±446 µatm. Only the highest pCO2 caused a significant reduction of growth rates and settlement of S. spirorbis individuals. Additionally, S. spirorbis settled juveniles exhibited enhanced calcification of 40% during daylight hours compared to dark hours, possibly reflecting a day-night alternation of an acidification-modulating effect by algal photosynthesis as opposed to an acidification-enhancing effect of algal respiration. E. pilosa colonies showed significantly increased growth rates at intermediate pCO2 (1193 µatm) but no response to higher pCO2. No effect of acidification on A. hirsutum colonies growth rates was observed. The results suggest a remarkable resistance of the algal macro-epibionts to levels of acidification occurring at present day upwellings in the Baltic. Only extreme future upwelling conditions impacted the tubeworm S. spirorbis, but not the bryozoans.
About 30% of the anthropogenically released CO2 is taken up by the oceans; such uptake causes surface ocean pH to decrease and is commonly referred to as ocean acidification (OA). Foraminifera are one of the most abundant groups of marine calcifiers, estimated to precipitate ca. 50 % of biogenic calcium carbonate in the open oceans. We have compiled the state of the art literature on OA effects on foraminifera, because the majority of OA research on this group was published within the last three years. Disparate responses of this important group of marine calcifiers to OA were reported, highlighting the importance of a process-based understanding of OA effects on foraminifera. We cultured the benthic foraminifer Ammonia sp. under a range of carbonate chemistry manipulation treatments to identify the parameter of the carbonate system causing the observed effects. This parameter identification is the first step towards a process-based understanding. We argue that CO3 is the parameter affecting foraminiferal size-normalized weights (SNWs) and growth rates. Based on the presented data, we can confirm the strong potential of Ammonia sp. foraminiferal SNW as a CO3 proxy.
Ocean warming and acidification may substantially affect the reproduction of keystone species such as Fucus vesiculosus (Phaeophyceae). In four consecutive benthic mesocosm experiments, we compared the reproductive biology and quantified the temporal development of Baltic Sea Fucus fertility under the single and combined impact of elevated seawater temperature and pCO2 (1100 ppm). In an additional experiment, we investigated the impact of temperature (0–25°C) on the maturation of North Sea F. vesiculosus receptacles. A marked seasonal reproductive cycle of F. vesiculosus became apparent in the course of 1 year. The first appearance of receptacles on vegetative apices and the further development of immature receptacles of F. vesiculosus in autumn were unaffected by warming or elevated pCO2. During winter, elevated pCO2 in both ambient and warmed temperatures increased the proportion of mature receptacles significantly. In spring, warming and, to a lesser extent, elevated pCO2 accelerated the maturation of receptacles and advanced the release of gametes by up to 2 weeks. Likewise, in the laboratory, maturation and gamete release were accelerated at 15–25°C relative to colder temperatures. In summary, elevated pCO2 and/or warming do not influence receptacle appearance in autumn, but do accelerate the maturation process during spring, resulting in earlier gamete release. Temperature and, to a much lesser extent, pCO2 affect the temporal development of Fucus fertility. Thus, rising temperatures will mainly shift or disturb the phenology of F. vesiculosus in spring and summer, which may alter and/or hamper its ecological functions in shallow coastal ecosystems of the Baltic Sea.
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