Niche complementarity and facilitation drive positive diversity effects on biomass production in experimental benthic diatom biofilms

Up to now, relatively few diversity-production experiments have been performed using microorganisms. Benthic diatom communities from estuarine intertidal mudflats are especially interesting for this purpose as they are relatively species poor and are thus more easy to simulate in laboratory conditions. We studied the effect of diversity on biomass production during microcosm experiments with diatoms assembled in combinations of up to eight species. Our results demonstrate a highly positive effect of biodiversity on production, with transgressive overyielding occurring in more than half of the combinations. These strong positive diversity effects could largely be attributed to positive complementarity effects (covering both niche complementarity and facilitation), although negative selection effects effects at higher diversities. We found a significant positive relation between functional diversity and the net biodiversity effects, indicating niche complementarity. In addition, we provide one of the first mechanistic evidences for facilitation by which biodiversity can enhance ecosystem functioning. This was demonstrated by the improved growth of Cylindrotheca closterium after addition of spent medium obtained from other diatom species. The stimulated growth of C. closterium was explained by a shift to mixotrophic growth with a down-regulation of the photosynthetic apparatus

Complementarity effects drive positive diversity effects on biomass production in experimental benthic diatom biofilms

1. Positive effects of species diversity on ecosystem functioning have often been demonstrated in ‘macrobial’ communities. This relation and the responsible mechanisms are far less clear for microbial communities. Most experimental studies on microorganisms have used randomly assembled communities that do not resemble natural communities. It is therefore difficult to predict the consequences of realistic, non-random diversity loss.2. In this study, we used naturally co-occurring diatom species from intertidal mudflats to assemble communities with realistically decreasing diversity and analysed the effect of non-random species loss on biomass production.3. Our results demonstrate a highly positive biodiversity effect on production, with mixtures outperforming the most productive component species in more than half of the combinations. These strong positive diversity effects could largely be attributed to positive complementarity effects (including both niche complementarity and facilitation), despite the occurrence of negative selection effects which partly counteracted the positive complementarity effects at higher diversities.4. Facilitative interactions were, at least in part, responsible for the higher biomass production. For one of the species, Cylindrotheca closterium, we show its ability to significantly increase its biomass production in response to substances leaked into the culture medium by other diatom species. In these conditions, the species drastically reduced its pigment concentration, which is typical for mixotrophic growth.5. Synthesis. We show that both species richness and identity have strong effects on the biomass production of benthic diatom biofilms and that transgressive overyielding is common in these communities. In addition, we show mechanistic evidence for facilitation which is partly responsible for enhanced production. Understanding the mechanisms by which diversity enhances the performance of ecosystems is crucial for predicting the consequences of species loss for ecosystem functioning