Re-oligotrophication and warming stabilize phytoplankton networks.

Phytoplankton taxa are strongly interconnected as a network, which could show temporal dynamics and non-linear responses to changes in drivers at both seasonal and long-term scale. Using a high quality dataset of 20 Danish lakes (1989-2008), we applied extended Local Similarity Analysis to construct temporal network of phytoplankton communities for each lake, obtained sub-network for each sampling month, and then measured indices of network complexity and stability for each sub-network. We assessed how lake re-oligotrophication, climate warming and grazers influenced the temporal dynamics on network complexity and stability of phytoplankton community covering three aspects: seasonal trends, long-term trends and detrended variability. We found strong seasonality for the complexity and stability of phytoplankton network, an increasing trend for the average degree, modularity, nestedness, persistence and robustness, and a decreasing trend for connectance, negative:positive interactions and vulnerability. Our study revealed a cascading effect of lake re-oligotrophication, climate warming and zooplankton grazers on phytoplankton network stability through changes in network complexity characterizing diversity, interactions and topography. Network stability of phytoplankton increased with average degree, modularity, nestedness and decreased with connectance and negative:positive interactions. Oligotrophication and warming stabilized the phytoplankton network (enhanced robustness, persistence and decreased vulnerability) by enhancing its average degree, modularity, nestedness and by reducing its connectance, while zooplankton richness promoted stability of phytoplankton network through increases in average degree and decreases in negative interactions. Our results further indicate that the stabilization effects might lead to more closed, compartmentalized and nested interconnections especially in the deeper lakes, in the warmer seasons and during bloom periods. From a temporal dynamic network view, our findings highlight stabilization of the phytoplankton community as an adaptive response to lake re-oligotrophication, climate warming and grazers.

Abiotic and biotic drivers of temporal dynamics in the spatial heterogeneity of zooplankton communities across lakes in recovery from eutrophication.

Seasonal and annual dynamics of the zooplankton community in lakes are affected by changes in abiotic drivers, trophic interactions (e.g., changes in phytoplankton and fish communities and abundances) and habitat characteristics (e.g. macrophyte abundance and composition). However, little is known about the temporal responses of the zooplankton community to abiotic and biotic drivers across lakes at the regional scale. Using a comprehensive 20-year dataset from 20 Danish lakes in recovery from eutrophication, we assessed the seasonal and annual trends in the spatial heterogeneity of zooplankton community across lakes and related it to abiotic and biotic drivers. We found significant seasonality and inter-annual decreases in spatial zooplankton heterogeneity in both shallow and deep lakes, with the decrease in the spatial turnover dominating the temporal dynamics of the beta diversity. For the inter-annual changes, decreased spatial heterogeneity of phytoplankton, macrophytes and fish were important biotic drivers at the regional scale. Using a series of ordinary least squares regressions and model selection with model averaging approaches, we revealed that both local (e.g., total phosphorus, total nitrogen, pH, Secchi depth, alkalinity, Schmidt stability, water temperature) and regional drivers (e.g., air temperature, solar irradiance) were important variables influencing the spatial zooplankton heterogeneity, although the directions depended on the beta diversity measures and water depth. Our results highlight an important role of bottom-up forces through phytoplankton community as well as macrophytes and top-down forces via fishes in driving the temporal changes in zooplankton community composition patterns at the regional scale.

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.