Phosphorus addition ameliorates soil micro-food web simplification due to nitrogen enrichment but does not restore nematode community composition

Nitrogen (N) inputs to terrestrial ecosystems are increasing faster than phosphorus (P) inputs, causing N:P imbalances and substantial changes in the diversity and structure of plant, soil microbial, and fauna communities. The responses of the soil micro-food web to N:P imbalances remain unclear but are crucial to understanding the impacts on ecosystem functioning. We investigated the effect of N and P additions on plant communities, edaphic properties, and nematode species diversity and composition, and assessed shifts in soil micro-food web complexity using nematode-based indices and structural equation modeling (SEM) in a nine-year nutrient addition experiment in an alpine meadow on the Tibetan Plateau. We found that N addition decreased the structure index (SI) and enrichment index (EI), indicating that N enrichment led to a simplification of the soil micro-food web. The addition of P alleviated the negative effect of N on soil micro-food web complexity but did not restore nematode community composition. The SEM supported the hypothesis that N addition simplified soil micro-food webs through increases in the N:P ratio and associated reductions in plant species richness, while P addition ameliorated the effect of N by decreasing the soil N:P ratio and increasing plant species richness. Our results highlight the positive effects of plant diversity in maintaining the complexity of the soil micro-food web. Our findings provide information that can be used to guide sustainable management and restoration of grasslands to alleviate the projected effects of future N:P imbalances on the soil micro-food web. However, efforts should be made to avoid nutrient deposition given the observed effects on both plant and nematode communities that could not be restored by alleviating the N:P imbalance through P addition.