Although metacommunity ecology has been a major field of research in the last decades, with both conceptual and empirical outputs, the analysis of the temporal dynamics of metacommunities has only emerged recently and consists mostly of repeated static analyses. Here we propose a novel analytical framework to assess metacommunity processes using path analyses of spatial and temporal diversity turnovers. We detail the principles and practical aspects of this framework and apply it to simulated datasets to illustrate its ability to decipher the respective contributions of entangled drivers of metacommunity dynamics. We then apply it to four empirical datasets. Empirical results support the view that metacommunity dynamics may be generally shaped by multiple ecological processes acting in concert, with environmental filtering being variable across both space and time. These results reinforce our call to go beyond static analyses of metacommunities that are blind to the temporal part of environmental variability.

Summary1. Eutrophication remains a major stress for freshwater biodiversity. Its deleterious consequences on biodiversity are well documented for large waterbodies. However, the impact of eutrophication may differ in smaller waterbodies, such as ponds and small lakes, which generally support naturally high levels of nutrients in lowlands. Furthermore, this response could depend on the scale considered, from local (individual waterbody, alpha diversity) to regional (the network of waterbodies, gamma diversity). It is also unclear whether the richness of threatened species responds in the same way as the richness of the whole assemblage. 2. The present study investigates local-and regional-scale consequences of eutrophication on taxonomic richness (all taxa) and conservation value (threatened taxa) in temperate lowland small waterbodies. Five taxonomic groups were investigated: macrophytes, gastropods, water beetles, adult dragonflies and amphibians, in a set of natural waterbodies and a set of enriched waterbodies covering a large nutrient gradient from mesotrophic to hypertrophic conditions. 3. Globally, our study did not reveal consistent, systematic responses to eutrophication. For macrophytes, the richness and conservation value suffered from eutrophication at both local and regional scales. In contrast, for amphibians and gastropods, eutrophication did not impair biodiversity at the local nor the regional scale. Dragonflies and water beetles showed intermediate situations, with an impairment by eutrophication varying according to the type of waterbodies considered. At the regional scale, each trophic status, even the nutrient richest, brought an original contribution to biodiversity. 4. Synthesis and applications. The management of eutrophication for small lowland waterbodies has to be considered differently than for lakes. For an individual waterbody (the local scale), nutrient enrichment is not necessarily a major impairment and its impact depends on the taxonomic group considered. Conversely, at the landscape scale, eutrophication is a major pressure on small waterbody biodiversity, especially because nutrient-rich small waterbodies are dominant in the landscape. Therefore, conservation efforts should integrate the notion of pond regional networks or 'pondscapes', where the regional biodiversity is supported by a mosaic of trophic conditions, and promote the presence of less rich waterbodies.