Macroecology of macrophytes in the freshwater realm: Patterns, mechanisms and implications

Broad-scale studies of species distributions and diversity have contributed to the emergence of general macroecological rules. These rules are typically founded on research using well-known terrestrial taxa as models and it is thus uncertain whether aquatic macrophytes follow these macroecological rules. Our purpose is to draw together available information from broad-scale research on aquatic macrophytes growing in lakes, ponds, wetlands, rivers and streams. We summarize how different macroecological rules fit the patterns shown by freshwater plants at various spatial scales. Finally, we outline future actions which should be taken to advance macroecological research on freshwater plants. Our review suggested that some macroecological patterns are relatively well-evidenced for aquatic macrophytes, whereas little information exists for others. We found, for example, that the species richness-latitude relationship follows a unimodal pattern, and species turnover prevails over species nestedness, whereas higher nestedness-related richness differences are found in low beta diversity regions. Contrary to terrestrial plants, climate or history seem not to be dominant determinants explaining these broad-scale patterns; instead local explanatory variables (e.g., water quality, such as alkalinity and nutrients, and hydromorphology) are often important for freshwater plants. We identified several knowledge gaps related, for example, to a smaller number of studies in lotic habitats, compared with lentic habitats, lack of spatially-adequate aquatic plant studies, deficiency of comprehensive species traits databases for aquatic macrophytes, and absence of a true phylogeny comprising most freshwater plant lineages. We hope this review will encourage the undertaking of additional macroecological investigations on freshwater plants across broad spatial and temporal scales.