ALBERT

Description: 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.

Description: The dramatic decline of biodiversity worldwide has raised a general concern on the impacts this process could have for the well-being of humanity. Human societies strongly depend on the benefits provided by natural ecosystems, which are the result of biogeochemical processes governed by species activities and their interaction with abiotic compartments. After decades of experimental research on the biodiversity-functioning relationship, a relative agreement has been reached on the mechanisms underlying the impacts that biodiversity loss can have on ecosystem processes. However, a general consensus is still missing. We suggest that the reason preventing an integration of existing knowledge is the scale discrepancy between observations on global change impacts and biodiversity-functioning experiments. The present chapter provides an overview of global change impacts on biodiversity across various ecological scales and its consequences for ecosystem functioning, highlighting what is known and where knowledge gaps still persist. Furthermore, the reader will be introduced to a set of tools that allow a multi-scale analysis of how global change drivers impact ecosystem functioning.

Description: Warming is one of the most dramatic aspects of climate change and threatens future ecosystem functioning. It may alter primary productivity and thus jeopardize carbon sequestration, a crucial ecosystem service provided by coastal environments. Fucus vesiculosus is an important canopy-forming macroalga in the Baltic Sea, and its main consumer is Idotea balthica. The objective of this study is to understand how temperature impacts a simplified food web composed of macroalgae and herbivores to quantify the effect on organic carbon storage. The organisms were exposed to a temperature gradient from 5 to 25 °C. We measured and modeled primary production, respiration, growth and epiphytic load on the surface of Fucus and respiration, growth and egestion of Idotea. The results show that temperature affects physiological responses of Fucus and Idotea separately. However, Idotea proved more sensitive to increasing temperatures than the primary producers. The lag between the collapse of the grazer and the decline of Fucus and epiphytes above 20 °C allows an increase of carbon storage of the primary productivity at higher temperatures. Therefore, along the temperature gradient, the simplified food web stores carbon in a non-monotonic way (reaching minimum at 20 °C). Our work stresses the need of considering the combined metabolic performance of all organisms for sound predictions on carbon circulation in food webs.

Description: Fluctuations in abundance of dominant species can cause competitive release of resources with consequences on community structure and functioning. In the present study, changes in the intertidal macroinfauna community of an exposed sandy beach were evaluated during two contrasting periods characterized by low and high densities of the yellow clam Amarilladesma mactroides. The increase in clam abundance and biomass was associated with a significant decrease in abundance of the rest of the community. In particular, a decline was observed for the pea crab Austinixa patagoniensis, a commensal species that lives in the burrows of the shrimp Sergio mirim. Our study demonstrates that fluctuations in clam abundance lead to long-term changes in community structure, suggesting the presence of competitive interactions. The environmental stability over the two periods strengthens the hypothesis that the competition between species is crucial for shaping the ecological community. Stable isotope analysis allows discarding trophic competition as mechanism of exclusion. Image maps reveal complementary distribution of species, showing the relevance of the spatial competition, which is mediated by changes in abundance of a third species. Indeed, high densities of A. mactroides reduce the available area for the establishment of the S. mirim burrows, limiting the foraging behavior of its commensal, the pea crab. Such an interaction drives density-dependent exclusion of the pea crab from the intertidal zone following the establishment of the yellow clam population. This study illustrates that spatial competition triggered by the increase of a bed-forming species can have community-wide consequences in exposed sandy beaches

Description: Environmental changes have been rapidly increasing in the last decades, causing unprecedented shifts in biodiversity. The impacts of biodiversity changes on ecosystem processes depend on the traits of affected species and their functional redundancy at the community level. The generated data on biodiversity-functioning in marine environments are still fragmentary and predictions on how species, communities and ecosystems will respond to the ongoing global changes are still uncertain. This selection of manuscripts presents the efforts of researchers around the world towards a better understanding on the mechanisms driving biodiversity and functioning patterns in marine ecosystems. The issue is composed of studies about first records of diversity and single species patterns in overlooked marine communities, effects of pollution in shaping species composition, foundation species and the impact of their loss on local communities, and the relevance of ecological interactions and species’ traits in structuring marine food webs. We conclude that more field and experimental studies combined to modelling are needed for understanding mechanisms that currently determine the structure and functioning of ecosystems and for improving predictions under global change scenarios.

Description: 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.

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.

Description: The world is going through severe climatic changes and this transition period is characterized by peculiar phenomena. Despite the mean global temperature increase, episodes of higher frequency and longer heatwaves will occur more often. Heatwaves have already disrupted resilience of macrophyte species, resulting in massive mortality of seagrasses and macroalgae. Macrophytes are responsible for considerable amounts of carbon storage and sequestration in marine coastal ecosystems. Many macrophytes are also considered foundation species since they harbor high species diversity, which results in large capacity to structure a community. Thus, heatwaves are able to cause shifts in biodiversity and ecosystem functioning. The goal of this thesis was to improve the understanding of the mechanistic effect of climate change on temperate coastal marine ecosystems based on macrophytes meadows. The first chapter showed that grazing intensity (that also depends on temperature) contributed to control the amount of carbon stored on the macroalgae and modified the trend to a non-linear pattern. The lowest carbon storage capacity was found between 20 and 22 °C, following the consumption intensity of the grazers. The second chapter, through the analysis of attributes and the application of ecological network analysis, I demonstrated that three sequential heatwaves jeopardized the capacity of the ecosystem to store carbon, since photosynthesis declined and the size of the ecosystem decreased. Moreover, three heatwaves caused a simplification on the pathways of carbon circulation, which makes the ecosystem more vulnerable to further disturbances. In the third chapter, I analyzed with a qualitative network model (loop analysis) the impact of sequential heatwaves on the capacity of the ecosystem to deliver services. The provision of water purification and climate regulation services was impaired, while the capacity of habitat provision did not change after the exposure to heatwaves. Therefore, the conclusions of this thesis are that: (1) both changes in temperature regimes (i.e. average constant and heatwaves) have profound effects on single species physiological performance and modify trophic interactions, thus altering energy circulation in food webs; (2) three consecutive heatwaves during spring/summer represented a threat for the health of the benthic ecosystem studied due to the reduction of productivity and the lowered diversity of energy flows, which increased vulnerability of the system; (3) besides having affected the functioning of the ecosystem, the heatwaves also harmed the capacity of the ecosystem to provide water purification and climate regulation services. This thesis showed that more knowledge on ecosystem functioning and services can be generated by combining experimental and modeling approaches. Furthermore, focusing on whole ecosystems rather than on isolated responses of single organisms expands our comprehension on the effects of changes in temperature regimes, which may assist further climate change mitigation measures.

Description: 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.

Description: Highlights: • Networks indicators reveal structure of the food webs depicted by ecosystem models. • Fishing mortality affects the structure and functioning of the food webs. • Increasing fishing mortality of all fish groups triggers strong indicator response. • Overfishing endangers ecosystem resilience. Marine ecosystems are exposed to multiple stressors, mainly fisheries that, whenever mismanaged, may cause irreversible damages to whole food webs. Ecosystem models have been applied to forecast fisheries impact on fish stocks and marine food webs. These impacts have been studied through the use of multiple indicators that help to understand ecosystem responses to stressors. This study focused on a category of ecological indicators derived from the network theory to quantify energy flows inside the food web. These indicators were computed using two ecosystem models applied to the Eastern English Channel (i.e. Atlantis and OSMOSE). This work aimed at investigating how several ecological network indicators respond to different levels of fishing pressure and evaluating their robustness to model structure and fishing strategies. We applied a gradient of fishing mortality using two ecosystem models and carried out ecological network analysis to obtain network-derived indicators. The results revealed that the indicators response is highly driven by the food web structure, although the model assumptions buffered some results. The indicators computed from OSMOSE outputs were more sensitive to changes in fishing pressure than those from Atlantis. However, once the food web from Atlantis was simplified to mimic the structure of OSMOSE model, the indicators of the modified Atlantis became more sensitive to the intensity of fishing pressure. The indicators related to amount of energy flow and to the organization of the flows in the food web were sensitive to the increase of fishing mortality for all fishing strategies. These indicators suggested that increasing fishing mortality jeopardizes the amount of energy mobilized by the food webs and simplifies the ecological interactions, which has implications for the resilience of marine ecosystems. The study shed light on the trophic networks structure and functioning of the ecosystems whenever exposed to distur-bances. Furthermore, these indicators might be adequate for whole ecosystem assessments of health and contribute to ecosystem management.