Climate warming and land-use changes drive broad-scale floristic changes in Southern Sweden.

Land-use changes, pollution and climate warming during the 20th century have caused changes in biodiversity across the world. However, in many cases, the environmental drivers are poorly understood. To identify and rank the drivers currently causing broad-scale floristic changes in N Europe, we analysed data from two vascular plant surveys of 200 randomly selected 2.5 × 2.5 km grid-squares in Scania, southernmost Sweden, conducted 1989-2006 and 2008-2015, respectively, and related the change in frequency (performance) of the species to a wide range of species-specific plant traits. We chose traits representing all plausible drivers of recent floristic changes: climatic change (northern distribution limit, flowering time), land-use change (light requirement, response to grazing/mowing, response to soil disturbance), drainage (water requirement), acidification (pH optimum), nitrogen deposition and eutrophication (N requirement, N fixation ability, carnivory, parasitism, mycorrhizal associations), pollinator decline (mode of reproduction) and changes in CO2 levels (photosynthetic pathway). Our results suggest that climate warming and changes in land-use were the main drivers of changes in the flora during the last decades. Climate warming appeared as the most influential driver, with northern distribution limit explaining 30%-60% of the variance in the GLMM models. However, the relative importance of the drivers differed among habitat types, with grassland species being affected the most by cessation of grazing/mowing and species of ruderal habitats by on-going concentration of both agriculture and human population to the most productive soils. For wetland species, only pH optimum was significantly related to species performance, possibly an effect of the increasing humification of acidic water bodies. An observed relative decline of mycorrhizal species may possibly be explained by decreasing nitrogen deposition resulting in less competition for phosphorus. We found no effect of shortage or decline of pollinating lepidopterans and bees.