Declining calcium concentration drives shifts toward smaller and less nutritious zooplankton in northern lakes.

Zooplankton community composition of northern lakes is changing due to the interactive effects of climate change and recovery from acidification, yet limited data are available to assess these changes combined. Here, we built a database using archives of temperature, water chemistry and zooplankton data from 60 Scandinavian lakes that represent broad spatial and temporal gradients in key parameters: temperature, calcium (Ca), total phosphorus (TP), total organic carbon (TOC), and pH. Using machine learning techniques, we found that Ca was the most important determinant of the relative abundance of all zooplankton groups studied, while pH was second, and TOC third in importance. Further, we found that Ca is declining in almost all lakes, and we detected a critical Ca threshold in lake water of 1.3 mg L-1 , below which the relative abundance of zooplankton shifts toward dominance of Holopedium gibberum and small cladocerans at the expense of Daphnia and copepods. Our findings suggest that low Ca concentrations may shape zooplankton communities, and that current trajectories of Ca decline could promote widespread changes in pelagic food webs as zooplankton are important trophic links from phytoplankton to fish and different zooplankton species play different roles in this context.

Mixed effects of a national protected area network on terrestrial and freshwater biodiversity.

Protected areas are considered fundamental to counter biodiversity loss. However, evidence for their effectiveness in averting local extinctions remains scarce and taxonomically biased. We employ a robust counterfactual multi-taxon approach to compare occupancy patterns of 638 species, including birds (150), mammals (23), plants (39) and phytoplankton (426) between protected and unprotected sites across four decades in Finland. We find mixed impacts of protected areas, with only a small proportion of species explicitly benefiting from protection-mainly through slower rates of decline inside protected areas. The benefits of protection are enhanced for larger protected areas and are traceable to when the sites were protected, but are mostly unrelated to species conservation status or traits (size, climatic niche and threat status). Our results suggest that the current protected area network can partly contribute to slow down declines in occupancy rates, but alone will not suffice to halt the biodiversity crisis. Efforts aimed at improving coverage, connectivity and management will be key to enhance the effectiveness of protected areas towards bending the curve of biodiversity loss.

Eutrophication effect on production and transfer of omega-3 fatty acids in boreal lake food webs.

Eutrophication, i.e. increasing level of nutrients and primary production, is a central environmental change of lakes globally with wide effects on food webs. However, how eutrophication affects the synthesis of physiologically essential biomolecules (omega-3 fatty acids) and their transfer to higher trophic levels at the whole food web level is not well understood. We assessed food web (phytoplankton, zooplankton, and fish) biomass, community structure and fatty acid content (eicosapentaenoic acid [EPA], and docosahexaenoic acid [DHA]), together with fatty acid specific primary production in 12 Finnish boreal lakes covering the total nutrient gradient from oligotrophic to highly eutrophic lakes (4-140 µg TP l-1; 413-1814 µg TN l-1). Production was measured as the incorporation of 13C-NaHCO3 into phytoplankton fatty acids and differentiated into volumetric production (production per litre of water) and productivity (production per phytoplankton biomass). Increases in nutrients led to higher biomass of phytoplankton, zooplankton and fish communities while also affecting community composition. Eutrophication negatively influenced the contribution of phytoplankton biomass preferentially grazed by zooplankton (<35 µm). Total volumetric production saturated at high phytoplankton biomass while EPA volumetric production presented a logarithmic relationship with nutrient increase. Meanwhile, total and EPA productivity had unimodal responses to this change in nutrients. DHA volumetric production and productivity presented large variation with increases in total phosphorus, but a unimodal model best described DHA changes with eutrophication. Results showed that eutrophication impaired the transfer of EPA and DHA into zooplankton and fish, showing a clear negative impact in some species (e.g. perch) while having no effect in other species (e.g. roach, ruffe). Results show non-linear trends in fatty acid production and productivity peaking at nutrient concentrations 22-35 µg l-1 TP followed by a gradual decrease.

Macrosystem community change in lake phytoplankton and its implications for diversity and function.

Aim: We use lake phytoplankton community data to quantify the spatio-temporal and scale-dependent impacts of eutrophication, land-use and climate change on species niches and community assembly processes while accounting for species traits and phylogenetic constraints. Location: Finland. Time period: 1977-2017. Major taxa: Phytoplankton. Methods: We use hierarchical modelling of species communities (HMSC) to model metacommunity trajectories at 853 lakes over four decades of environmental change, including a hierarchical spatial structure to account for scale-dependent processes. Using a "region of common profile" approach, we evaluate compositional changes of species communities and trait profiles and investigate their temporal development. Results: We demonstrate the emergence of novel and widespread community composition clusters in previously more compositionally homogeneous communities, with cluster-specific community trait profiles, indicating functional differences. A strong phylogenetic signal of species responses to the environment implies similar responses among closely related taxa. Community cluster-specific species prevalence indicates lower taxonomic dispersion within the current dominant clusters compared with the historically dominant cluster and an overall higher prevalence of smaller species sizes within communities. Our findings denote profound spatio-temporal structuring of species co-occurrence patterns and highlight functional differences of lake phytoplankton communities. Main conclusions: Diverging community trajectories have led to a nationwide reshuffling of lake phytoplankton communities. At regional and national scales, lakes are not single entities but metacommunity hubs in an interconnected waterscape. The assembly mechanisms of phytoplankton communities are strongly structured by spatio-temporal dynamics, which have led to novel community types, but only a minor part of this reshuffling could be linked to temporal environmental change.

Consistency of Targeted Metatranscriptomics and Morphological Characterization of Phytoplankton Communities.

The composition of phytoplankton community is the basis for environmental monitoring and assessment of the ecological status of aquatic ecosystems. Community composition studies of phytoplankton have been based on time-consuming and expertise-demanding light microscopy analyses. Molecular methods have the potential to replace microscopy, but the high copy number variation of ribosomal genes and the lack of universal primers for simultaneous amplification of prokaryotic and eukaryotic genes complicate data interpretation. In this study, we used our previously developed directional primer-independent high-throughput sequencing (HTS) approach to analyze 16S and 18S rRNA community structures. Comparison of 83 boreal lake samples showed that the relative abundances of eukaryotic phytoplankton at class level and prokaryotic cyanobacteria at order level were consistent between HTS and microscopy results. At the genus level, the results had low correspondence, mainly due to lack of sequences in the reference library. HTS was superior to identify genera that are extensively represented in the reference databases but lack specific morphological characteristics. Targeted metatranscriptomics proved to be a feasible method to complement the microscopy analysis. The metatranscriptomics can also be applied without linking the sequences to taxonomy. However, direct indexing of the sequences to their environmental indicator values needs collections of more comprehensive sample sets, as long as the coverage of molecular barcodes of eukaryotic species remains insufficient.

Eutrophication reduces the nutritional value of phytoplankton in boreal lakes.

Eutrophication (as an increase in total phosphorus [TP]) increases harmful algal blooms and reduces the proportion of high-quality phytoplankton in seston and the content of ω-3 long-chain polyunsaturated fatty acids (eicosapentaenoic acid [EPA] and docosahexaenoic acid [DHA]) in fish. However, it is not well-known how eutrophication affects the overall nutritional value of phytoplankton. Therefore, we studied the impact of eutrophication on the production (as concentration; µg L-1) and content (µg mg C-1) of amino acids, EPA, DHA, and sterols, i.e., the nutritional value of phytoplankton in 107 boreal lakes. The lakes were categorized in seven TP concentration categories ranging from ultra-oligotrophic (<5 µg L-1) to highly eutrophic (>50 µg L-1). Phytoplankton total biomass increased with TP as expected, but in contrast to previous studies, the contribution of high-quality phytoplankton did not decrease with TP. However, the high variation reflected instability in the phytoplankton community structure in eutrophic lakes. We found that the concentration of amino acids increased in the epilimnion whereas the concentration of sterols decreased with increasing TP. In terms of phytoplankton nutritional value, amino acids, EPA, DHA, and sterols showed a significant quadratic relationship with the lake trophic status. More specifically, the amino acid contents were the same in the oligo- and mesotrophic lakes, but substantially lower in the eutrophic lakes (TP > 35 µg L-1/1.13 µmol L-1). The highest EPA and DHA content in phytoplankton was found in the mesotrophic lakes, whereas the sterol content was highest in the oligotrophic lakes. Based on these results, the nutritional value of phytoplankton reduces with eutrophication, although the contribution of high-quality algae does not decrease. Therefore, the results emphasize that eutrophication, as excess TP, reduces the nutritional value of phytoplankton, which may have a significant impact on the nutritional value of zooplankton, fish, and other aquatic animals at higher food web levels.

Identifying multiple stressors that influence eutrophication in a Finnish agricultural river.

In Finland, a recent ecological classification of surface waters showed that the rivers and coastal waters need attention to improve their ecological state. We combined eco-hydrological and empirical models to study chlorophyll-a concentration as an indicator of eutrophication in a small agricultural river. We used a modified story-and-simulation method to build three storylines for possible changes in future land use due to climate change and political change. The main objective in the first storyline is to stimulate economic activity but also to promote the sustainable and efficient use of resources. The second storyline is based on the high awareness but poor regulation of environmental protection, and the third is to survive as individual countries instead of being part of a unified Europe. We assumed trade of agricultural products to increase to countries outside Europe. We found that chlorophyll-a concentration in the river depended on total phosphorus concentration. In addition, there was a positive synergistic interaction between total phosphorus and water temperature. In future storylines, chlorophyll-a concentration increased due to land use and climate change. Climate change mainly had an indirect influence via increasing nutrient losses from intensified agriculture. We found that well-designed agri-environmental measures had the potential to decrease nutrient loading from fields, as long as the predicted increase in temperature remained under 2 °C. However, we were not able to achieve the nutrient reduction stated in current water protection targets. In addition, the ecological status of the river deteriorated. The influence of temperature on chlorophyll-a growth indicates that novel measures for shading rivers to decrease water temperature may be needed in the future.

Suitability of Phytosterols Alongside Fatty Acids as Chemotaxonomic Biomarkers for Phytoplankton.

The composition and abundance of phytoplankton is an important factor defining ecological status of marine and freshwater ecosystems. Chemotaxonomic markers (e.g., pigments and fatty acids) are needed for monitoring changes in a phytoplankton community and to know the nutritional quality of seston for herbivorous zooplankton. Here we investigated the suitability of sterols along with fatty acids as chemotaxonomic markers using multivariate statistics, by analyzing the sterol and fatty acid composition of 10 different phytoplankton classes including altogether 37 strains isolated from freshwater lakes. We were able to detect a total of 47 fatty acids and 29 sterols in our phytoplankton samples, which both differed statistically significantly between phytoplankton classes. Due to the high variation of fatty acid composition among Cyanophyceae, taxonomical differentiation increased when Cyanophyceae were excluded from statistical analysis. Sterol composition was more heterogeneous within class than fatty acids and did not improve separation of phytoplankton classes when used alongside fatty acids. However, we conclude that sterols can provide additional information on the abundance of specific genera within a class which can be generated by using fatty acids. For example, whereas high C16 ω-3 PUFA (polyunsaturated fatty acid) indicates the presence of Chlorophyceae, a simultaneous high amount of ergosterol could specify the presence of Chlamydomonas spp. (Chlorophyceae). Additionally, we found specific 4α-methyl sterols for distinct Dinophyceae genera, suggesting that 4α-methyl sterols can potentially separate freshwater dinoflagellates from each other.