Forest understorey plant responses to long-term experimental warming, light and nitrogen addition.

Climate change, eutrophication and intensified forest management are affecting forest understorey plants, a major component of forest biodiversity. The main impacts of these drivers have often been studied, but we lack a good understanding of how key understorey species are affected by potential interactive effects of these drivers and which species drive community changes. Here we assessed the responses of 15 species occurring in the understorey of a deciduous temperate forest to experimental warming, light addition and enhanced nitrogen inputs in permanent plots surveyed for 9 years. We analysed vegetation cover and key functional traits (plant height, specific leaf area and reproductive traits) at the species level and identified the species driving community change with principal response curves (PRC). Light addition and warming, and to a lesser extent also nitrogen addition, had profound effects on cover and functional traits. Many species showed directional change over time, and this change can either be strengthened or weakened by treatments, indicating the importance of long-term monitoring. Against expectations, we observed few interactions between treatments. Species responses to treatments were related to ecological strategies (generalists versus forest specialist). Generalists, such as Rubus fruticosus, benefitted from the warming and light treatments and outcompeted forest specialists. This might ultimately lead to biotic homogenization. Since the treatment effects of light and warming were additive, keeping the canopy closed will only mitigate, but not stop, the effects of global warming on the forest understorey plants.