Topography- and management-mediated resource gradients maintain rare and common plant diversity around paddy terraces

Examining the causes of interspecific differences in susceptibility to bidirectional land-use changes (land abandonment and use-intensification) is important for understanding the mechanisms of global biodiversity loss in agricultural landscapes. We tested the hypothesis that rare (endangered) plant species prefer wet and oligotrophic areas within topography- and management-mediated resource (soil water content, nutrient, and aboveground biomass) gradients, making them more susceptible to both abandonment and use-intensification of agricultural lands. We demonstrated that topography and management practices generated resource gradients in seminatural grasslands around traditional paddy terraces. Terraced topography and management practices produced a soil moisture gradient within levees and a nutrient gradient within paddy terraces. Both total and rare species diversity increased with soil water content. Total species diversity increased in more eutrophied areas with low aboveground biomass, whereas rare species diversity was high under oligotrophic conditions. Rare and common species were differentially distributed along the human-induced nutrient gradient, with rare species preferring wet, nutrient-poor environments in the agricultural landscapes studied. We suggest that conservation efforts should concentrate on wet, nutrient-poor areas within such landscapes, which can be located easily using land-use and topography maps. This strategy would reduce the costs of finding and conserving rare grassland species in a given agricultural landscape.