Long-term nutrient load reductions and increasing lake TN : TP stoichiometry decrease phytoplankton biomass and diversity in a large shallow lake

Nutrient loading of freshwater and marine habitats has increased during the last century as a result of anthropogenic activities. From the 1980s onwards, following implementation of new policy targeting eutrophication, total phosphorus (TP) and total nitrogen (TN) loads were reduced in many European waters. Often, however, decreases in TP were stronger as compared to TN, leading to increased TN : TP ratios. Our analysis shows that the large and shallow lake IJsselmeer (the Netherlands) experienced a similar trend, whereas TN was reduced by 50%, TP was reduced by 89% between 1975 and 2018. Most of this nutrient load reduction was achieved before the year 2000, changes in nutrient concentrations in the lake became smaller afterwards, especially for TN, leading to a further increase in stoichiometric imbalance up to a yearly averaged TN : TP (molar) of 296 in 2018. The observed changes in nutrients were accompanied by a decline in total phytoplankton biomass, and slight declines in phytoplankton genus evenness and diversity. Although biomass decreases likely resulted from the overall decrease in nutrient availabilities, the reduced diversity may have resulted from the shift toward very high TN : TP ratios that indicate relatively low TP levels and enhanced competition for phosphorus. Overall, our findings demonstrate long-term trends with decreased phytoplankton biomass and diversity following reduced nutrient concentrations and enhanced stoichiometric imbalance. Ultimately, such changes at the food web base may alter the structure and functioning of the entire aquatic food- web in lake IJsselmeer.