Small Shallow Lakes (SSL) support exceptionally high and original biodiversity, providing numerous ecosystem services. Their small size makes them especially sensitive to anthropic activities, that causes a shift to dysfunctional turbid states and induces loss of services and biodiversity. In this study we investigated the relationships between environmental factors and macrophyte communities. Macrophytes play a crucial role in maintaining functional clear states. Better understanding factors determining the composition and richness of aquatic plant communities in least-impacted conditions may be useful to protect them. We inventoried macrophyte communities and collected chemical, climatic and morphological data from 89 least-impacted SSL widely distributed in France. SSL were sampled across four climatic ecoregions, various geologies and elevations. Hierarchical cluster analysis showed a clear separation of four macrophyte assemblages strongly associated with mineralisation. Determinant factors identified by db-RDA analysis are, in order of importance, geology, distance from source (DIS, a proxy for connectivity with river hydrosystems), surface area, climate and hydroperiod (water permanency). Surprisingly, at country-wide scale, climate and hydroperiod filter macrophyte composition weakly. Geology and DIS are the major determinants of community composition, whereas surface area determines floristic richness. DIS is identified as determinant in freshwater lentic ecosystems for the first time.

Biological invasions have reached an unprecedented level and the number of introduced species is still increasing worldwide. Despite major advances in invasion science, the determinants of success of introduced species, the magnitude and dimensions of their impact, and the mechanisms sustaining successful invasions are still debated. Empirical studies show divergent impacts of non-native populations on ecosystems and contrasting effects of biotic and abiotic factors on the dynamics of non-native populations; this is hindering the emergence of a unified theory of biological invasions. We propose a synthesis that merges perspectives from population, community, and ecosystem levels. Along a timeline of ecosystem transformation driven by non-native species, from historical to human-modified ecosystems, we order invasion concepts and theories to clarify their chaining and relevance during each step of the invasion process. This temporal sorting of invasion concepts shows that each concept is relevant at a specific stage of the invasion. Concepts and empirical findings on non-native species may appear contradictory. However, we suggest that, when mapped onto an invasion timeline, they may be combined in a complementary way. An overall scheme is proposed to summarise the theoretical dynamics of ecosystems subjected to invasions. For any given case study, this framework provides a guide through the maze of theories and should help choose the appropriate concepts according to the stage of invasion.