Aquatic habitats are often characterized by both high diversity and the threat of multiple anthropogenic stressors. Our research deals with temporal and spatial aspects of two of the main threats for biodiversity, namely eutrophication and fragmentation. It is known that pulsed nutrient addition creates temporal differences in environmental conditions, promoting higher diversity by preventing the best competitor from dominating. Furthermore, a metacommunity landscape with intermediate connectivity increases autotrophs’ diversity and stability. However, it is yet unclear if these two factors are additive in increasing diversity and if the effects extend to the consumer level. With the goal of understanding how eutrophication impacts biodiversity in a metacommunity landscape, we hypothesized that pulsed nutrient addition will increase diversity among both autotrophs and heterotrophs, and this effect will be even greater in a metacommunity landscape. We simulated eutrophication and fragmentation in a microcosm experiment using phytoplankton as primary producers and microzooplankton as grazers. Four treatment combinations were tested including two different landscapes (metacommunity and isolated community) and two forms of nutrient supply (pulsed and continuous): metacommunity/continuous nutrient addition (MC); metacommunity/pulsed nutrient addition (MP); isolated community/continuous nutrient addition (IC); isolated community/pulsed nutrient addition (IP). As expected, pulsed nutrient addition had a persistent positive effect on phytoplankton diversity, with a weaker influence of landscape type. In contrast, the grazer community strongly benefited from a metacommunity landscape, with less significance of pulsed or continuous nutrient addition. Overall, the metacommunity landscape with pulsed nutrient supply supported higher diversity of primary producers and grazers. Electronic supplementary material The online version of this article (10.1007/s00442-018-4319-8) contains supplementary material, which is available to authorized users.

Cell size is a master trait in the functional ecology of phytoplankton correlating with numerous morphological, physiological, and life-cycle characteristics of species that constrain their nutrient use, growth, and edibility. In contrast to well-known spatial patterns in cell size at macroecological scales or temporal changes in experimental contexts, few data sets allow testing temporal changes in cell sizes within ecosystems. To analyze the temporal changes of intraspecific and community-wide cell size, we use the phytoplankton data derived from the Lower Saxony Wadden Sea monitoring program, which comprises sample-and species-specific measurements of cell volume from 1710 samples collected over 14 yr. We find significant reductions in both the cell volume of most species and the weighted mean cell size of communities. Mainly diatoms showed this decline, whereas the size of dinoflagellates seemed to be less responsive. The magnitude of the trend indicates that cell volumes are about 30% smaller now than a decade ago. This interannual trend is overlayed by seasonal cycles with smaller cells typically observed in summer. In the subset of samples including environmental conditions, small community cell size was strongly related to high temperatures and low total phosphorus concentration. We conclude that cell size captures ongoing changes in phytoplankton communities beyond the changes in species composition. In addition, based on the changes in species biovolumes revealed by our analysis, we warn that using standard cell size values in phytoplankton assessment will not only miss temporal changes in size, but also lead to systematic errors in biomass estimates over time.

The current policy and goals aimed to conserve biodiversity and manage biodiversity change are often formulated at the global scale. At smaller scales however, biodiversity change is more nuanced leading to a plethora of trends in different metrics of alpha diversity and temporal turnover. Therefore, largescale policy targets do not translate easily into local to regional management decisions for biodiversity. Using long-term monitoring data from the Wadden Sea (Southern North Sea), joining structural equation models and general dissimilarity models enabled a better overview of the drivers of biodiversity change. Few commonalities emerged as birds, fish, macroinvertebrates, and phytoplankton differed in their response to certain drivers of change. These differences were additionally dependent upon the biodiversity aspect in question and which environmental data were recorded in each monitoring program. No single biodiversity metric or model sufficed to capture all ongoing change, which requires an explicitly multivariate approaches to biodiversity assessment in local ecosystem management.

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One of the key challenges in managing eutrophication in coastal marine ecosystems is the harmonized cross-border assessment of phytoplankton. Some general understanding of the consequences of shifting nutrient regimes can be derived from the detailed investigation of the phytoplankton community and its biodiversity. Here, we combined long-term monitoring datasets of German and Dutch coastal stations and amended these with additional information on species biomass. Across the integrated and harmonized data set, we used multiple biodiversity descriptors to analyse temporal trends in the Wadden Sea phytoplankton. Biodiversity, measured as the number of species (S) and the effective number of species (ENS), has decreased in the Dutch stations over the last 20 years, while biomass has increased, indicating that fewer species are becoming more dominant in the system. However, biodiversity and biomass did not show substantial changes in the German stations. Although there were some differences in trends between countries, shifts in community composition and relative abundance were consistent across stations and time. Through a multi-metric approach to biodiversity and species biomass analysis, we have been able to gain a better understanding of changes in the Wadden Sea over the last 20 years. We emphasise the importance of continuous and harmonised monitoring programmes that can detect changes in the communities that are indicative of changes in the environment.

Temporal heterogeneity in nutrient availability is known to increase phytoplankton diversity by allowing more species to coexist under different resource niches. Spatial heterogeneity has also been positively correlated with species diversity. Here we investigated how temporal and spatial differences in nutrient addition together impact biodiversity in metacommunities varying in the degree of connectivity among the patches. We used a microcosm experimental design to test two spatiotemporal ways of supplying nutrients: synchronously (nutrients were added regionally—to all four patches at the same time) and asynchronously (nutrients were added locally—to a different patch each time), combined with two different degrees of connectivity among the patches (low or high connectivity). We used three species of algae and one species of cyanobacteria as the primary producers; and five ciliate and two rotifer species as the grazers. We expected higher diversity in metacommunities receiving an asynchronous nutrient supply, assuming stronger development of heterogeneous patches with this condition rather than with synchronous nutrient supply. This result was expected, however, to be dependent on the degree of connectivity among patches. We found significant effects of nutrient addition in both groups of organisms. Phytoplankton diversity increased until the fourth week (transiently) and zooplankton richness was persistently higher under asynchronous nutrient addition. Our results were consistent with our hypothesis that asynchronicity in nutrient supply would create a more favorable condition for species to co-occur. However, this effect was, in part, transient and was not influenced by the degree of connectivity.

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