Mixtures of macrophyte growth forms promote nitrogen cycling in wetlands.

The importance of aquatic plant diversity in regulating nutrient cycling in wetlands remains poorly understood. We investigated how variation in macrophyte growth form (emerging, submerged and bryophyte) combinations and species mixtures affect nitrogen (N) removal from the water and N accumulation in plant biomass. We conducted a wetland mesocosm experiment for 100 days during July-September 2015. Twelve species were grown in mono- and in two-species mixed cultures for a total of 32 single and two-growth form combinations. Nitrogen removal from the water was quantified on three occasions during the experiment, while N accumulation in plant biomass was determined following termination of the experiment. The number of species and growth forms present increased N removal and accumulation. The growth form combinations of emerging and bryophyte species showed the highest N accumulation and N removal from water, followed by combinations of emerging species. By contrast, submerged species growing in the presence of emerging or other submerged species showed the lowest levels of N accumulation and N removal. Temporal variation in N removal also differed among growth form combinations: N removal was highest for emerging-bryophyte combinations in July, but peaked for the emerging-submerged and emerging-bryophyte combinations in August. Indeed, the occurrence of complementarity among macrophyte species, particularly in combinations of bryophyte and emerging species, enhanced N removal and uptake during the entire growing season. Our study highlights the importance of bryophytes, which have been neglected in research on nutrient cycling in wetlands, for aquatic N cycling, especially given their worldwide distribution across biomes. Overall, our findings point towards the potential important role of the diversity of macrophyte growth forms in regulating key ecosystem processes related to N cycling in wetlands.