Larger direct than indirect effects of multiple environmental changes on leaf nitrogen of forest herbs

Aims Disentangling direct and indirect effects of global change drivers on plant nitrogen (N) uptake in leaves is important for understanding species and community responses in a changing world. Methods We created understorey herb communities on forest soils with and without recent agricultural history. We traced pulse additions of (NH4NO3)-N-15-N-15 within these mesocosms while applying two-level factorial treatments of N enrichment, warming and illumination. We modelled direct and indirect effects of treatments on leaf N content and N-15 uptake in leaves. Results Warming and illumination had three times larger direct negative effects on leaf N content per dry mass and percentage leaf N derived from label (Ndfl%) than their indirect negative effects via an increasing community cover. These results imply a tissue dilution of N with increasing growth, in response to environmental change directly and indirectly exacerbated by community cover. We additionally found that interspecific differences in Ndfl% correlated with a species' colonisation capacity and resource acquisition strategy. Conclusions Global change can directly affect allocation of N into foliage, with simultaneous indirect effects via altered community properties that influence individual plant responses. Predicting the future of plant communities in a changing world requires accounting for such understudied pathways.