Seasonal and long-term trends in the spatial heterogeneity of lake phytoplankton communities over two decades of restoration and climate change.

World-wide, reducing the external nutrient loading to lakes has been the primary priority of lake management in the restoration of eutrophic lakes over the past decades, and as expected this has resulted in an increase in the local environmental heterogeneity, and thus biotic heterogeneity, within lakes. However, little is known about how the regional spatial heterogeneity of lake biotic communities changes with restoration across a landscape. Using a long-term monitoring dataset from 20 Danish lakes, we elucidated the seasonal and long-term trends in the spatial heterogeneity of climate, local abiotic variables and phytoplankton communities over two decades of restoration and climate change at landscape level. We found significant seasonality in the spatial heterogeneity of most climatic and local drivers as well as in the total beta diversity (Sørensen coefficient) and its turnover components (Simpson coefficient) of phytoplankton communities among the lakes. The seasonality tended to be less marked in deep than in shallow lakes. We found significant spatial homogenisation of most local drivers (except for alkalinity) and phytoplankton communities after two decades of restoration and that turnover dominated the temporal responses of the total beta diversity of phytoplankton communities. Path analyses showed that the homogenisation of phytoplankton communities was mainly due to a decrease in spatial heterogeneity of total phosphorus and Schmidt stability in shallow lakes and to a decrease in spatial total phosphorus and total nitrogen heterogeneity in deep lakes. However, albeit weakly, the spatial heterogeneity of the phytoplankton communities was affected indirectly by climatic warming in both shallow and deep lakes and directly by wind speed in shallow lakes. We conclude that restoration of eutrophic lakes may lead to an increase in the local heterogeneity of phytoplankton communities at lake scale and an increase in homogeneity at landscape scale.