Water chemistry and fluffy layer geochemistry and mineral content at four sites in the western Baltic Sea

The fluffy layer was sampled repeatedly during nine expeditions between October 1996 and December 1998 at four stations situated along a S-N-transect from the Oder Estuary to the Arkona Basin. Geochemical and mineralogical analyses of the fluff show regional differences (trends) in composition, attributed to provenance and to hydrographical conditions along their transport pathways. Temporal variability is very high at the shallow water station of the estuary, and decreases towards the deeper stations in the north. In the shallow water area, intensive resuspension of the fluff due to wind-driven waves and currents leads to an average residence time of only one to two days. Near-bottom lateral transport of the fluff is the main process that transfers the fine grained material, containing both nutrients and contaminants, from the coastal zone into the deeper basins of the Baltic Sea. Seasonal effects (e.g. biogenic production in relation to trace metal variation) are observed at the Tromper Wiek station, where the residence time of the fluffy material is in the scale of seasons. Thus, the fluffy layer offers suitable material for environmental monitoring programs.

Oxygen isotope ratios on calcite from the Piànico lake record

Stable oxygen isotope analyses at annual, 2-, 5-, 10- and 20-varve sample resolutions were carried out on two selected varve intervals from the interglacial sediment record of the Piànico palaeolake. These sediments are particularly suitable for ultra-high-resolution isotope analyses on lacustrine endogenic calcite because of the exceptionally well-preserved varve structure. A bias through detrital contamination can be excluded because microscopically controlled sampling enabled selecting detritus-free samples. The studied sediment intervals comprise 352 and 88 continuous varve series formed during periods of rapid climate change at the onset and end of a marked millennial-scale cool interval during the Piànico Interglacial. The most intriguing result is a pronounced short-term oscillation in the bi-annually resolved isotope record superimposed on the general decreasing and increasing d18O trends at the climatic transitions that is recorded at lower sample resolution. Spectral analyses of the bi-annual time series reveal periodicities indicating solar and NAO controls on the d18O record. Multiple d18O measurements from endogenic calcite of individual varves showed variations of up to 0.6 per mil, thus larger than the observed inter-annual variability and most likely explained by seasonal effects.

Warming and acidification experiment on early life-stage F. vesiculosus under natural fluctuation and seasonal variations (spring 2013 - 2014)

Genetic diversity of baltic F. vesiculosus is low compared to other populations which might jeopardize their potential for adaptation to climate change. Especially the early life-stage F. vesiculosus may be threaten by ocean warming and acidification. To test this, we exposed F. vesiculosus germlings to warming and acidification in the near-natural scenario in the "Kiel Outdoor Benthocosms" maintaining the natural variation of the Kiel Fjord, Germany (54°27 'N, 10°11 'W) in all seasons (spring 2013 - 2014). Warming was simulated by using a delta treatment adding 5 °C and by increasing pCO2 at 1000 µatm. Warming positively affected germlings' growth in spring and in summer but decreased non-photochemical quenching in spring and survival in summer. Acidified conditions showed much weaker effects than warming. The high genotypic variation in stress sensitivity as well as the enhanced survival at high diversity levels indicate higher potential for adaptation for genetically diverse populations. We conclude that the combination of stressors and season determines the sensitivity to environmental stress and that genetic variation is crucial for the adaptation to climate change stress.

Temperature and photoperiod effects on sex determination in Leuresthes tenuis (fish)

In some gonochoristic species, sex is influenced not only by genotype at conception but also by the environment that offspring experience during early ontogeny (termed environmental sex determination or ESD). ESD is thought to be adaptive when seasonal variations in environmental conditions provide a sex-specific fitness advantage. In vertebrates, temperature is the most common determinant of sex, and seasonal variation in temperature serves as a temporal cue of environmental quality such as length of the growing season. Some environments, however, lack strong seasonal temperature fluctuations and other cues, particularly photoperiod, may provide a more reliable indicator of the environment offspring enter. We tested this hypothesis by rearing the offspring of the California grunion (Leuresthes tenuis, Ayres), which experiences low seasonal temperature variation in nature, under common garden conditions at three temperature and two photoperiod treatments. Our experiments revealed that both temperature and photoperiod significantly affected sex ratios in L. tenuis. More females were produced at cooler temperatures and longer day lengths, which is consistent with female biased sex ratios early in the breeding season, and likely adaptive through increased female size and fecundity. To our knowledge, this is the first documented case of photoperiod-dependent sex determination in a gonochoristic vertebrate.

Experiment: Food availability outweighs ocean acidification effects in juvenile Mytilus edulis

Ocean acidification is expected to decrease calcification rates of bivalves. Nevertheless in many coastal areas high pCO2 variability is encountered already today. Kiel Fjord (Western Baltic Sea) is a brackish (12-20 g kg-1) and CO2 enriched habitat, but the blue mussel Mytilus edulis dominates the benthic community. In a coupled field and laboratory study we examined the annual pCO2 variability in this habitat and the combined effects of elevated pCO2 and food availability on juvenile M. edulis growth and calcification. In the laboratory experiment, mussel growth and calcification were found to chiefly depend on food supply, with only minor impacts of pCO2 up to 3350 µatm. Kiel Fjord was characterized by strong seasonal pCO2 variability. During summer, maximal pCO2 values of 2500 µatm were observed at the surface and >3000 µatm at the bottom. However, the field growth experiment revealed seven times higher growth and calcification rates of M. edulis at a high pCO2 inner fjord field station (mean pCO2 ca. 1000 µatm) in comparison to a low pCO2 outer fjord station (ca. 600 µatm). In addition, mussels were able to outcompete the barnacle Amphibalanus improvisus at the high pCO2 site. High mussel productivity at the inner fjord site was enabled by higher particulate organic carbon concentrations. Kiel Fjord is highly impacted by eutrophication, which causes bottom water hypoxia and consequently high seawater pCO2. At the same time, elevated nutrient concentrations increase the energy availability for filter feeding organisms such as mussels. Thus M. edulis can dominate over a seemingly more acidification resistant species such as A. improvisus. We conclude that benthic stages of M. edulis tolerate high ambient pCO2 when food supply is abundant and that important habitat characteristics such as species interactions and energy availability need to be considered to predict species vulnerability to ocean acidification.

Ocean acidification outweighs nutrient effects in structuring seagrass epiphyte communities

1. Developing a framework for assessing interactions between multiple anthropogenic stressors remains an important goal in environmental research. In coastal ecosystems, the relative effects of aspects of global climate change (e.g. CO2 concentrations) and localized stressors (e.g. eutrophication), in combination, have received limited attention. 2. Using a long-term (11 month) field experiment, we examine how epiphyte assemblages in a tropical seagrass meadow respond to factorial manipulations of dissolved carbon dioxide (CO2(aq)) and nutrient enrichment. In situ CO2(aq) manipulations were conducted using clear, open-top chambers, which replicated carbonate parameter forecasts for the year 2100. Nutrient enrichment consisted of monthly additions of slow-release fertilizer, nitrogen (N) and phosphorus (P), to the sediments at rates equivalent to theoretical maximum rates of anthropogenic loading within the region (1.54 g N/m**2/d and 0.24 g P m**2/d). 3. Epiphyte community structure was assessed on a seasonal basis and revealed declines in the abundance of coralline algae, along with increases in filamentous algae under elevated CO2(aq). Surprisingly, nutrient enrichment had no effect on epiphyte community structure or overall epiphyte loading. Interactions between CO2(aq) and nutrient enrichment were not detected. Furthermore, CO2(aq)-mediated responses in the epiphyte community displayed strong seasonality, suggesting that climate change studies in variable environments should be conducted over extended time-scales. 4. Synthesis. The observed responses indicate that for certain locations, global stressors such as ocean acidification may take precedence over local eutrophication in altering the community structure of seagrass epiphyte assemblages. Given that nutrient-driven algal overgrowth is commonly cited as a widespread cause of seagrass decline, our findings highlight that alternate climate change forces may exert proximate control over epiphyte community structure.

Small Changes in pH Have Direct Effects on Marine Bacterial Community Composition: A Microcosm Approach

As the atmospheric CO2 concentration rises, more CO2 will dissolve in the oceans, leading to a reduction in pH. Effects of ocean acidification on bacterial communities have mainly been studied in biologically complex systems, in which indirect effects, mediated through food web interactions, come into play. These approaches come close to nature but suffer from low replication and neglect seasonality. To comprehensively investigate direct pH effects, we conducted highly-replicated laboratory acidification experiments with the natural bacterial community from Helgoland Roads (North Sea). Seasonal variability was accounted for by repeating the experiment four times (spring, summer, autumn, winter). Three dilution approaches were used to select for different ecological strategies, i.e. fast-growing or low-nutrient adapted bacteria. The pH levels investigated were in situ seawater pH (8.15-8.22), pH 7.82 and pH 7.67, representing the present-day situation and two acidification scenarios projected for the North Sea for the year 2100. In all seasons, both automated ribosomal intergenic spacer analysis and 16S ribosomal amplicon pyrosequencing revealed pH-dependent community shifts for two of the dilution approaches. Bacteria susceptible to changes in pH were different members of Gammaproteobacteria, Flavobacteriaceae, Rhodobacteraceae, Campylobacteraceae and further less abundant groups. Their specific response to reduced pH was often context-dependent. Bacterial abundance was not influenced by pH. Our findings suggest that already moderate changes in pH have the potential to cause compositional shifts, depending on the community assembly and environmental factors. By identifying pH-susceptible groups, this study provides insights for more directed, in-depth community analyses in large-scale and long-term experiments.

Effects of in situ CO2 enrichment on structural characteristics, photosynthesis, and growth of the Mediterranean seagrass Posidonia oceanica

Seagrass is expected to benefit from increased carbon availability under future ocean acidification. This hypothesis has been little tested by in situ manipulation. To test for ocean acidification effects on seagrass meadows under controlled CO2/pH conditions, we used a Free Ocean Carbon Dioxide Enrichment (FOCE) system which allows for the manipulation of pH as continuous offset from ambient. It was deployed in a Posidonia oceanica meadow at 11 m depth in the Northwestern Mediterranean Sea. It consisted of two benthic enclosures, an experimental and a control unit both 1.7 m**3, and an additional reference plot in the ambient environment (2 m**2) to account for structural artifacts. The meadow was monitored from April to November 2014. The pH of the experimental enclosure was lowered by 0.26 pH units for the second half of the 8-month study. The greatest magnitude of change in P. oceanica leaf biometrics, photosynthesis, and leaf growth accompanied seasonal changes recorded in the environment and values were similar between the two enclosures. Leaf thickness may change in response to lower pH but this requires further testing. Results are congruent with other short-term and natural studies that have investigated the response of P. oceanica over a wide range of pH. They suggest any benefit from ocean acidification, over the next century (at a pH of 7.7 on the total scale), on Posidonia physiology and growth may be minimal and difficult to detect without increased replication or longer experimental duration. The limited stimulation, which did not surpass any enclosure or seasonal effect, casts doubts on speculations that elevated CO2 would confer resistance to thermal stress and increase the buffering capacity of meadows.

Synergistic effects of ocean acidification and warming on overwintering pteropods in the Arctic

Ocean acidification and warming will be most pronounced in the Arctic Ocean. Aragonite shell-bearing pteropods in the Arctic are expected to be among the first species to suffer from ocean acidification. Carbonate undersaturation in the Arctic will first occur in winter and because this period is also characterized by low food availability, the overwintering stages of polar pteropods may develop into a bottleneck in their life cycle. The impacts of ocean acidification and warming on growth, shell degradation (dissolution), and mortality of two thecosome pteropods, the polar Limacina helicina and the boreal L. retroversa, were studied for the first time during the Arctic winter in the Kongsfjord (Svalbard). The abundance of L. helicina and L. retroversa varied from 23.5 to 120 ind /m2 and 12 to 38 ind /m2, and the mean shell size ranged from 920 to 981 µm and 810 to 823 µm, respectively. Seawater was aragonite-undersaturated at the overwintering depths of pteropods on two out of ten days of our observations. A 7-day experiment [temperature levels: 2 and 7 °C, pCO2 levels: 350, 650 (only for L. helicina) and 880 ?atm] revealed a significant pCO2 effect on shell degradation in both species, and synergistic effects between temperature and pCO2 for L. helicina. A comparison of live and dead specimens kept under the same experimental conditions indicated that both species were capable of actively reducing the impacts of acidification on shell dissolution. A higher vulnerability to increasing pCO2 and temperature during the winter season is indicated compared with a similar study from fall 2009. Considering the species winter phenology and the seasonal changes in carbonate chemistry in Arctic waters, negative climate change effects on Arctic thecosomes are likely to show up first during winter, possibly well before ocean acidification effects become detectable during the summer season.

The synergistic effects of ocean acidification and organic metabolism on calcium carbonate (CaCO3) dissolution in coral reef sediments

Ocean acidification (OA) is expected to reduce the net ecosystem calcification (NEC) rates and overall accretion of coral reef ecosystems. However, despite the fact that sediments are the most abundant form of calcium carbonate (CaCO3) in coral reef ecosystems and their dissolution may be more sensitive to OA than biogenic calcification, the impacts of OA induced sediment dissolution on coral reef NEC rates and CaCO3 accretion are poorly constrained. Carbon dioxide addition and light attenuation experiments were performed at Heron Island, Australia in an attempt to tease apart the influence of OA and organic metabolism (e.g. respiratory CO2 production) on CaCO3 dissolution. Overall, CaCO3 dissolution rates were an order of magnitude more sensitive to elevated CO2 and decreasing seawater aragonite saturation state (Omega Ar; 300-420% increase in dissolution per unit decrease in Omega Ar) than published reductions in biologically mediated calcification due to OA. Light attenuation experiments led to a 70% reduction in net primary production (NPP), which subsequently induced an increase in daytime (115%) and net diel (375%) CaCO3 dissolution rates. High CO2 and low light acted in synergy to drive a 575% increase in net diel dissolution rates. Importantly, disruptions to the balance of photosynthesis and respiration (P/R) had a significant effect on daytime CaCO3 dissolution, while average water column ?Ar was the main driver of nighttime dissolution rates. A simple model of platform-integrated dissolution rates was developed demonstrating that seasonal changes in photosynthetically active radiation (PAR) can have an important effect on platform integrated CaCO3 sediment dissolution rates. The considerable response of CaCO3 sediment dissolution to elevated CO2 means that much of the response of coral reef communities and ecosystems to OA could be due to increases in CaCO3 sediment and framework dissolution, and not decreases in biogenic calcification.