Nutrient addition increases the capacity for division of labor and the benefits of clonal integration in an invasive plant

Many of the most invasive plants are clonal, and clonal integration has been proposed as an important mechanism promoting invasiveness. When the availabilities of two essential resources are negatively correlated in space, clonal integration may benefit clonal plants through division of labor. We hypothesized that environments with reciprocal patchiness of light and soil water may induce division of labor, and nutrient addition may increase both the division of labor and the benefits of clonal integration. To test this, we grew pairs of connected and disconnected ramets of the clonal invader Mikania micrantha under negative spatial covariance of light and soil water such that the proximal ramets were grown under high light and low soil water conditions and the distal ramets were grown under low light and high soil water conditions. In half of the ramet pairs, both ramets of a pair received a nutrient addition treatment. The results showed that connection decreased the root to shoot ratio in proximal ramets and increased it in distal ramets, indicating that division of labor was induced. In addition, connection increased the root to shoot ratio of distal ramets more under high soil nutrient conditions than under low soil nutrient conditions, indicating that nutrient addition increased the division of labor. Connection increased plant biomass at the whole clonal fragment level, and this increase was larger under high soil nutrient conditions than under low soil nutrient conditions. This study showed, for the first time, that in environments with reciprocal patchiness of two essential resources, the capacity for division of labor and its influence on plant performance may depend on the availability of a third essential resource. Because invasive plants often can acquire a larger amount of soil resources than native plants, our study may also contribute to the understanding why clonality is related to invasiveness. (C) 2018 Elsevier B.V. All rights reserved.