Description: Warming and acidification of the oceans as a consequence of increasing CO2-concentrations occur at large scales. Numerous studies have shown the impact of single stressors on individual species. However, studies on the combined effect of multiple stressors on a multi-species assemblage, which is ecologically much more realistic and relevant, are still scarce. Therefore, we orthogonally crossed the two factors warming and acidification in mesocosm experiments and studied their single and combined impact on the brown alga Fucus vesiculosus associated with its natural community (epiphytes and mesograzers) in the Baltic Sea in all seasons (from April 2013 to April 2014). We superimposed our treatment factors onto the natural fluctuations of all environmental variables present in the Benthocosms in so-called delta-treatments. Thereby we compared the physiological responses of F. vesiculosus (growth and metabolites) to the single and combined effects of natural Kiel Fjord temperatures and pCO2 conditions with a 5 °C temperature increase and/or pCO2 increase treatment (1100 ppm in the headspace above the mesocosms). Responses were also related to the factor photoperiod which changes over the course of the year. Our results demonstrate complex seasonal pattern. Elevated pCO2 positively affected growth of F. vesiculosus alone and/or interactively with warming. The response direction (additive, synergistic or antagonistic), however, depended on season and daylength. The effects were most obvious when plants were actively growing during spring and early summer. Our study revealed for the first time that it is crucial to always consider the impact of variable environmental conditions throughout all seasons. In summary, our study indicates that in future F. vesiculosus will be more affected by detrimental summer heat-waves than by ocean acidification although the latter consequently enhances growth throughout the year. The mainly negative influence of rising temperatures on the physiology of this keystone macroalga may alter and/or hamper its ecological functions in the shallow coastal ecosystem of the Baltic Sea.

Description: Fucus vesiculosus L. (Phaeophyceae) is the most abundant and hence ecologically most important primary producer, carbon sink and habitat provider in the western Baltic Sea. All F. vesiculosus L. specimens were collected on 23 April 2014 from a depth of 0.2-1 m in the non-tidal Kiel Fjord, western Baltic Sea (54°27'N; 10°12'E), where this species forms dense and almost monospecific stands on stones. After sampling the algal thalli were stored in a refrigerator box with water from the sampling site, transported to Bremerhaven and stored at 10 °C for one day in filtered seawater. Experiments were conducted with vegetative apical tips (6.7±0.5 cm length), the actively growing region of F. vesiculosus, which were randomly selected and cut from 144 different individuals prior to the experiments. These tips were acclimated to laboratory conditions for three days in filtered seawater at 10 °C before the start of the experiment. Furthermore, 30 additional vegetative apices were freeze-dried to document the initial biochemical status of F. vesiculosus in its native habitat. A temperature gradient was installed in a walk-in constant cooling chamber (15 °C) in nine water baths (5, 10, 15, 20, 24, 26, 27, 28 and 29 °C ± 0.1 °C) which were tempered by thermostats (5, 10 and 15 °C: Huber Variostat CC + Pilot ONE, Peter Huber Kältemaschinen GmbH, Offenburg, Germany; 20 and 28 °C: Haake DC3, Thermo Fisher Scientific Inc., Waltham, USA; 24, 26, 27 and 29 °C: Haake DC10). Every temperature treatment consisted of four 2 L glass beakers (n = 4). In each beaker four F. vesiculosus apices were grown in 2 µm-filtered North Sea water diluted with demineralized water in a ratio of 1:1 and enriched with nutrients after Provasoli (1968; 1/10 enrichment), leading to a salinity of about 15.6 which equaled habitat conditions. The algae were exposed to an irradiance of 130 µmol photons m-2 s-1 ±10 % (Powerstar HGI-TS 150 W, OSRAM GmbH, Bad Homburg, Germany) measured at the top of the beaker under a 16:8 h L:D cycle. The media in the beakers was changed every third or fourth day and aerated with artificial air containing 380 ppm CO2 (gas mixing device; HTK Hamburg GmbH, Hamburg, Germany). Before the experiment, the algae were acclimated to the final temperatures in steps of 5 °C for 2 days each, beginning at 10 °C. After 21 days exposure time, three out of four samples per replicate were freeze-dried for further biochemical analyses, and afterwards the thermostats were turned off to reduce the temperature to 16±0.4 °C for another 10 days permitting growth under post-culture conditions.

Description: Coastal marine ecosystems have been under high anthropogenic pressure and it can be assumed that prevalent local perturbation interacts with rising global stressors under proceeding climate change. Understanding their effective pathways and cumulative effects is of high relevance not only with regard to future risk assessment, but also for current ecosystem management. In benthic mesocosms, we factorially tested the effects of one global (combined elevated seawater temperature and CO2 concentration) and one local (nutrient enrichment) stressor on a common coastal Baltic seaweed system (Fucus vesiculosus). Both treatments in combination had additive negative impacts on the seaweed-epiphyte-mesograzer system by altering its regulatory mechanisms. That is, warming decreased the biomass of two mesograzer species (weakened top-down control), whereas moderate nutrient enrichment increased epiphyte biomass (intensified bottom-up control), which ultimately resulted in a significant biomass reduction of the foundation seaweed. Our results suggest that climate change impacts might be underestimated if local pressures are disregarded. Furthermore, they give implication for local ecological management as the mitigation of local perturbation may limit climate change impacts on marine ecosystems.

Description: 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.

Description: Ocean acidification and warming effects on the macroalgal species Fucus vesiculosus forma mytili were simulated in the tidal benthic mesocosm facility at the AWI Wadden Sea Station on the island of Sylt, Germany (55°01'19.2''N, 8°26'17.7''E). The SY1 experiment in spring 2014 (11 weeks from early April to late June) was based on a "Temp X pCO2" full-factorial setup (ambient or delta 5°C temperature X ambient or 1000 ppm pCO2) resulting in 4 treatment levels à 3 replicates. The seawater parameters (temperature, pH, salinity, total alkalinity, pCO2) and seawater inorganic nutrient concentrations (silicate, nitrite, phosphate, ammonium, total nitrogen oxide, nitrate) were measured on a weekly basis. The biomass growth rates (RGR) of Fucus vesiculosus forma mytili were calculated over time, and its physiological parameters (carbon, nitrogen, CN ratio, mannitol) were measured at the beginning and end of the experiment.

Description: The plea for using more “realistic,” community‐level, investigations to assess the ecological impacts of global change has recently intensified. Such experiments are typically more complex, longer, more expensive, and harder to interpret than simple organism‐level benchtop experiments. Are they worth the extra effort? Using outdoor mesocosms, we investigated the effects of ocean warming (OW) and acidification (OA), their combination (OAW), and their natural fluctuations on coastal communities of the western Baltic Sea during all four seasons. These communities are dominated by the perennial and canopy‐forming macrophyte Fucus vesiculosus—an important ecosystem engineer Baltic‐wide. We, additionally, assessed the direct response of organisms to temperature and pH in benchtop experiments, and examined how well organism‐level responses can predict community‐level responses to the dominant driver, OW. OW affected the mesocosm communities substantially stronger than acidification. OW provoked structural and functional shifts in the community that differed in strength and direction among seasons. The organism‐level response to OW matched well the community‐level response of a given species only under warm and cold thermal stress, that is, in summer and winter. In other seasons, shifts in biotic interactions masked the direct OW effects. The combination of direct OW effects and OW‐driven shifts of biotic interactions is likely to jeopardize the future of the habitat‐forming macroalga F. vesiculosus in the Baltic Sea. Furthermore, we conclude that seasonal mesocosm experiments are essential for our understanding of global change impact because they take into account the important fluctuations of abiotic and biotic pressures.