Small scale environmental variation modulates plant defence syndromes of understorey plants in deciduous forests of Europe

Aim: Variation in plant defence traits has been frequently assessed along largescale macroclimatic clines. In contrast, local-scale changes in the environment have recently been proposed to also modulate plant defence traits. Yet, the relative importance of drivers at both scales has never been tested. We aimed to quantify the relative importance of environmental drivers inherent to large and small spatial scales on the physical and chemical defence and tolerance to herbivory in understorey plant species of deciduous forests of Europe. Location: Deciduous forests in Europe. Time period: Present. Major taxa studied: Forest understorey plants. Methods: We sampled four typical ancient forest herbs (Anemone nemorosa, Oxalis acetosella, Deschampsia cespitosa, Milium effusum) along small and large spatial scale gradients (those driven by latitude, elevation, forest management and distance to the forest edge), and analysed a suite of nine constitutively expressed traits associated with overall resistance to herbivory, and their multivariate response to environmental clines. Results: Although our study included a large gradient in macroclimate, we found variation in the local environment at small spatial scales (i.e. soil nutrient concentration and forest structural complexity) to be more important in predicting plant resistance to herbivory. Main conclusions: In addition to macroclimatic conditions, subtle differences in forest microclimate and soil characteristics also played a major role in modulating plant defence phenotypes. These findings highlight the importance of the local habitat structure and environmental conditions in modulating plant resistance to herbivory.