Urbanization causes shifts in species' trait state frequencies

Urbanization is one of the most extreme forms of land transformation. It is supposed to change the frequencies of species trait states in species assemblages.We hypothesize that the flora of urban and rural areas differs in the frequency of trait states and ask which traits enable a plant to cope with the urban environment. We tested our hypothesis in Germany, which was divided into grid-cells of ca 130 km2. We distinguished urbanized (with more than 33% urban land use; n = 59), agricultural (with more than 50% agricultural land use; n = 1365) and semi-natural (with more than 50% forest and semi-natural land use; n = 312) grid-cells and calculated the proportions of plant species per trait state in each grid-cell. Multiple linear regressions explained the log-transformed ratio of one proportion to another with land use (urban, agricultural, semi-natural) and the environmental parameters (climate, topography, soils and geology). Additionally, linear mixed effect models accounted for the effects of land use and biogeography and differences in sample size of the three grid-cell types. Urbanized and rural areas showed clear differences in the proportion of trait states. Urbanized grid-cells had e.g., higher proportions of wind-pollinated plants, plants with scleromorphic leaves or plants dispersed by animals, and lower proportions of insect-pollinated plants, plants with hygromorphic leaves or plants dispersed by wind than other grid-cells. Our study shows that shifts in land use can change the trait state composition of plant assemblages. Far-reac