Climate change will not only shift environmental means but will also increase the intensity of extreme events, exerting additional stress on ecosystems. While field observations on the ecological consequences of heat waves are emerging, experimental evidence is rare, and lacking at the community level. Using a novel "near-natural" outdoor mesocosms approach, this study tested whether marine summer heat waves have detrimental consequences for macrofauna of a temperate coastal community, and whether sequential heat waves provoke an increase or decrease of sensitivity to thermal stress. Three treatments were applied, defined and characterized through a statistical analysis of 15 years of temperature records from the experimental site: (1) no heat wave, (2) two heat waves in June and July followed by a summer heat wave in August and (3) the summer heat wave only. Overall, 50% of the species showed positive, negative or positive/negative responses in either abundance and/or biomass. We highlight four possible ways in which single species responded to either three subsequent heat waves or one summer heat wave: (1) absence of a response (tolerance, 50% of species), (2) negative accumulative effects by three subsequent heat waves (tellinid bivalve), (3) buffering by proceeding heat waves due to acclimation and/or shifts in phenology (spionid polychaete) and (4) an accumulative positive effect by subsequent heat waves (amphipod). The differential responses to single or sequential heat waves at the species level entailed shifts at the community level. Community-level differences between single and triple heat waves were more pronounced than those between regimes with vs. without heat waves. Detritivory was reduced by the single heat wave while suspension feeding was less common in the triple heat wave regime. Critical extreme events occur already today and will occur more frequently in a changing climate, thus, leading to detrimental impacts on coastal marine systems.

Marine heatwaves have been observed worldwide and are expected to increase in both frequency and intensity due to climate change. Such events may cause ecosystem reconfigurations arising from species range contraction or redistribution, with ecological, economic and social implications. Macrophytes such as the brown seaweed Fucus vesiculosus and the seagrass Zostera marina are foundation species in many coastal ecosystems of the temperate northern hemisphere. Hence, their response to extreme events can potentially determine the fate of associated ecosystems. Macrophyte functioning is intimately linked to the maintenance of photosynthesis, growth and reproduction, and resistance against pathogens, epibionts and grazers. We investigated morphological, physiological, pathological and chemical defence responses of western Baltic Sea F. vesiculosus and Z. marina populations to simulated near‐natural marine heatwaves. Along with (a) the control, which constituted no heatwave but natural stochastic temperature variability (0HW), two treatments were applied: (b) two late‐spring heatwaves (June, July) followed by a summer heatwave (August; 3HW) and (c) a summer heatwave only (1HW). The 3HW treatment was applied to test whether preconditioning events can modulate the potential sensitivity to the summer heatwave. Despite the variety of responses measured in both species, only Z. marina growth was impaired by the accumulative heat stress imposed by the 3HW treatment. Photosynthetic rate, however, remained high after the last heatwave indicating potential for recovery. Only epibacterial abundance was significantly affected in F. vesiculosus. Hence both macrophytes, and in particular F. vesiculosus, seem to be fairly tolerant to short‐term marine heatwaves at least at the intensities applied in this experiment (up to 5°C above mean temperature over a period of 9 days). This may partly be due to the fact that F. vesiculosus grows in a highly variable environment, and may have a high phenotypic plasticity.

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.This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made.Additional Supporting Information may be found in the online version of this article.

While it is well known that severe marine summer heatwaves can cause acute and dramatic die‐offs of seagrass meadows, the effect of trans‐seasonal warming and winter/spring heatwaves are yet poorly understood. This study simulated a 9‐months warming scenario on the common seagrass Zostera marina from winter into summer, using outdoor mesocosms, which provided near‐natural conditions. The relevance of the natural temperature pattern, as well as the 3.6°C warming, and their implications were further discussed in the context of a 22‐yr temperature time series of the study region. Survival of plants was high in winter independent of temperature. In spring, however, heat‐treated Z. marina flowered 1.5 months earlier and experienced high mortalities. Thereafter, plant survival, growth, and pigmentation were largely comparable between temperature regimes. Yet, a comparatively high mortality occurred in ambient plants, after an abnormally warm June. Final biomass was reduced by ~ 50% in heat‐treated plants. These results imply that warm winter‐to‐spring conditions can have severe effects on vital seagrass traits. Warming accelerates consumption of energy reserves triggering advanced flowering, similar to many terrestrial plants. Although, surviving heat‐treated plants were not able to re‐stock energy reserves throughout the high‐light summer as inferred from low plant biomass, these seemed rather resistant to summer heatwave events.