Impacts of ocean acidification and warming on Arctic calcifying key species : benthic coralline red alga (Lithothamnion glaciale) and pelagic thecosome pteropods (Limacina helicina & L. retroversa) = Auswirkung von Ozeanversauerung und Ozeanerwärmung auf die Biologie arktischer kalzifizierender Schlüsselarten: Benthische koralline Rotalge (Lithothamnion glaciale) und pelagische thekosmome Flügelschnecken (Limacina helicina & L. retroversa)

Over the last two centuries human activities significantly changed the Earth system. Atmospheric CO2 concentrations were increased by ~25% in only 50 years (1965 to 2015) through combustion of fossil fuels and changes in land use. Consequences of these activities are a warming climate and the ‘acidification’ of the world oceans. A doubling of surface ocean carbon dioxide- (CO2) and proton- (H+) and a 40% decrease of the carbonate ion (CO32-) concentrations until the end of this century will probably affect many marine organisms. Due to low temperatures, the Arctic Surface Ocean will be one of the first to experience mean annual corrosive conditions for calcified organisms. This makes the Arctic Ocean a bellwether for global impacts of ocean acidification (OA) on marine life. Biogenic calcium carbonate often establishes ecological functions like being the fundament for species rich habitats in the case of hermatypic corals and crustose coralline algae (CCA) or impacting global carbon cycling embodied by coccolithophores, foraminifera and thecosome pteropods. The marine carbon cycle influences the destination of human CO2 emissions and the development of the global climate. Therefore, the ecological importance of CCA and pteropods, their involvement in the marine carbon cycle and their anticipated vulnerability to OA call for research on their fate in the future ocean. This doctoral thesis investigates the effects of warming and ocean acidification on growth, calcification, dissolution, corrosion and survival of Arctic calcifying keystone species Lithothamnion glaciale (CCA) and Limacina helicina and retroversa (thecosome pteropods)

Impact of ocean acidification and elevated temperatures on early juveniles of the polar shelled pteropod Limacina helicina: mortality, shell degradation, and shell growth

Due to their aragonitic shell thecosome pteropods may be particularly vulnerable to ocean acidification driven by anthropogenic CO2 emissions. This applies specifically to species inhabiting Arctic surface waters that are projected to become locally undersaturated with respect to aragonite as early as 2016. This study investigated the effects of rising pCO2 partial pressures and elevated temperature on pre-winter juveniles of the polar pteropod Limacina helicina. After a 29 days experiment in September/October 2009 at three different temperatures and under pCO2 scenarios projected for this century, mortality, shell degradation, shell diameter and shell increment were investigated. Temperature and pCO2 had a significant effect on mortality, but temperature was the overriding factor. Shell diameter, shell increment and shell degradation were significantly impacted by pCO2 but not by temperature. Mortality was 46% higher at 8 °C compared to 3 °C (in situ), and 14% higher at 1100 μatm CO2 as compared to 230 μatm CO2. Shell diameter and increment were reduced by 10% and 12% at 1100 μatm CO2 as compared to 230 μatm CO2, respectively, and shell degradation was 41% higher at elevated compared to ambient pCO2 partial pressures. We conclude that pre-winter juveniles will be negatively affected by both rising temperature and pCO2 which may result in a possible abundance decline of the overwintering population, the basis for next year's reproduction

Technical Note: A mobile sea-going mesocosm system - new opportunities for ocean change research

One of the great challenges in ocean change research is to understand and forecast the effects of environmental changes on pelagic communities and the associated impacts on biogeochemical cycling. Mesocosms, experimental enclosures designed to approximate natural conditions, and in which environmental factors can be manipulated and closely monitored, provide a powerful tool to close the gap between single species laboratory experiments and observational and correlative approaches applied in field surveys. Existing pelagic mesocosm systems are stationary and/or restricted to well-protected waters. To allow mesocosm experimentation in a range of hydrographic conditions and in areas considered most sensitive to ocean change, we developed a mobile, sea-going mesocosm facility, the Kiel Off-Shore Mesocosms for Future Ocean Simulations (KOSMOS). The KOSMOS platform, which can be transported and deployed by mid-sized research vessels, is designed for operation in moored and free-floating mode under low to moderate wave conditions (up to 2.5 m wave heights). It encloses a water column 2 m in diameter and 15 to 25 m deep (~50–75 m3 in volume) without disrupting the vertical structure or disturbing the enclosed plankton community. Several new developments in mesocosm design and operation were implemented to (i) minimize differences in starting conditions between mesocosms, (ii) allow for extended experimental duration, (iii) precisely determine the mesocosm volume, (iv) determine air–sea gas exchange, and (v) perform mass balance calculations. After multiple test runs in the Baltic Sea, which resulted in continuous improvement of the design and handling, the KOSMOS platform successfully completed its first full-scale experiment in the high Arctic off Svalbard (78° 56.2′ N, 11° 53.6′ E) in June/July 2010. The study, which was conducted in the framework of the European Project on Ocean Acidification (EPOCA), focused on the effects of ocean acidification on a natural plankton community and its impacts on biogeochemical cycling and air/sea exchange of climate relevant gases. This manuscript describes the mesocosm hardware, its deployment and handling, CO2 manipulation, sampling and cleaning, including some further modifications conducted based on the experiences gained during this study

Phytoplankton Blooms at Increasing Levels of Atmospheric Carbon Dioxide: Experimental Evidence for Negative Effects on Prymnesiophytes and Positive on Small Picoeukaryotes

Anthropogenic emissions of carbon dioxide (CO2) and the ongoing accumulation in the surface ocean together with concomitantly decreasing pH and calcium carbonate saturation states have the potential to impact phytoplankton community composition and therefore biogeochemical element cycling on a global scale. Here we report on a recent mesocosm CO2 perturbation study (Raunefjorden, Norway), with a focus on organic matter and phytoplankton dynamics. Cell numbers of three phytoplankton groups were particularly affected by increasing levels of seawater CO2 throughout the entire experiment, with the cyanobacterium Synechococcus and picoeukaryotes (prasinophytes) profiting, and the coccolithophore Emiliania huxleyi (prymnesiophyte) being negatively impacted. Combining these results with other phytoplankton community CO2 experiments into a data-set of global coverage suggests that, whenever CO2 effects are found, prymnesiophyte (especially coccolithophore) abundances are negatively affected, while the opposite holds true for small picoeukaryotes belonging to the class of prasinophytes, or the division of chlorophytes in general. Future reductions in calcium carbonate-producing coccolithophores, providing ballast which accelerates the sinking of particulate organic matter, together with increases in picoeukaryotes, an important component of the microbial loop in the euphotic zone, have the potential to impact marine export production, with feedbacks to Earth’s climate system.publishedVersio

Temporal biomass dynamics of an Arctic plankton bloom in response to increasing levels of atmospheric carbon dioxide

Ocean acidification and carbonation, driven by anthropogenic emissions of carbon dioxide (CO2), have been shown to affect a variety of marine organisms and are likely to change ecosystem functioning. High latitudes, especially the Arctic, will be the first to encounter profound changes in carbonate chemistry speciation at a large scale, namely the under-saturation of surface waters with respect to aragonite, a calcium carbonate polymorph produced by several organisms in this region. During a CO2 perturbation study in Kongsfjorden on the west coast of Spitsbergen (Norway), in the framework of the EU-funded project EPOCA, the temporal dynamics of a plankton bloom was followed in nine mesocosms, manipulated for CO2 levels ranging initially from about 185 to 1420 μatm. Dissolved inorganic nutrients were added halfway through the experiment. Autotrophic biomass, as identified by chlorophyll a standing stocks (Chl a), peaked three times in all mesocosms. However, while absolute Chl a concentrations were similar in all mesocosms during the first phase of the experiment, higher autotrophic biomass was measured as high in comparison to low CO2 during the second phase, right after dissolved inorganic nutrient addition. This trend then reversed in the third phase. There were several statistically significant CO2 effects on a variety of parameters measured in certain phases, such as nutrient utilization, standing stocks of particulate organic matter, and phytoplankton species composition. Interestingly, CO2 effects developed slowly but steadily, becoming more and more statistically significant with time. The observed CO2-related shifts in nutrient flow into different phytoplankton groups (mainly dinoflagellates, prasinophytes and haptophytes) could have consequences for future organic matter flow to higher trophic levels and export production, with consequences for ecosystem productivity and atmospheric CO2.publishedVersio

Influence of Ocean Acidification on a Natural Winter-to-Summer Plankton Succession : First Insights from a Long-Term Mesocosm Study Draw Attention to Periods of Low Nutrient Concentrations

Every year, the oceans absorb about 30% of anthropogenic carbon dioxide (CO2) leading to a re-equilibration of the marine carbonate system and decreasing seawater pH. Today, there is increasing awareness that these changes-summarized by the term ocean acidification (OA)-could differentially affect the competitive ability of marine organisms, thereby provoking a restructuring of marine ecosystems and biogeochemical element cycles. In winter 2013, we deployed ten pelagic mesocosms in the Gullmar Fjord at the Swedish west coast in order to study the effect of OA on plankton ecology and biogeochemistry under close to natural conditions. Five of the ten mesocosms were left unperturbed and served as controls (similar to 380 mu atm pCO(2)), whereas the others were enriched with CO2-saturated water to simulate realistic end-of-the-century carbonate chemistry conditions (mu 760 mu atm pCO(2)). We ran the experiment for 113 days which allowed us to study the influence of high CO2 on an entire winter-to-summer plankton succession and to investigate the potential of some plankton organisms for evolutionary adaptation to OA in their natural environment. This paper is the first in a PLOS collection and provides a detailed overview on the experimental design, important events, and the key complexities of such a "long-term mesocosm" approach. Furthermore, we analyzed whether simulated end-of-the-century carbonate chemistry conditions could lead to a significant restructuring of the plankton community in the course of the succession. At the level of detail analyzed in this overview paper we found that CO2-induced differences in plankton community composition were non-detectable during most of the succession except for a period where a phytoplankton bloom was fueled by remineralized nutrients. These results indicate: (1) Long-term studies with pelagic ecosystems are necessary to uncover OA-sensitive stages of succession. (2) Plankton communities fueled by regenerated nutrients may be more responsive to changing carbonate chemistry than those having access to high inorganic nutrient concentrations and may deserve particular attention in future studies.Peer reviewe