Combined effects of resource heterogeneity and simulated herbivory on plasticity of clonal integration in a rhizomatous perennial herb

Previous lines of investigation assuming potential advantage of clonal integration generally have neglected its plasticity in complex heterogeneous environments. Clonal plants adaptively respond to abiotic heterogeneity (patchy resource distribution) and herbivory-induced heterogeneity (within-clone heterogeneity in ramet performance), but to date little is known about how resource heterogeneity and simulated herbivory jointly affect the overall performance of clones. Partial damage within a clone caused by herbivory might create herbivory-induced heterogeneity in a resource-homogeneous environment, and might also decrease or increase the extent of heterogeneity under resource-heterogeneous conditions. We conducted a greenhouse experiment in which target-ramets of Leymus chinensis segments within homogeneous or heterogeneous nutrient treatments were subject to clipping (0% or 75% shoot removal). In homogeneous environments with high (9: 9) nutrient availability, ramet biomass of L. chinensis with intact or severed rhizomes is 0.70 or 0.69 g. Conversely, target-ramet biomass with intact rhizomes is obviously lower than that of the severed target-ramets in the homogeneous environments with medium (5: 5) and low (1: 1) nutrient availability. High resource availability and the presence of herbivory can alleviate negative effects of rhizome connection under homogeneous conditions, by providing copious resource or creating herbivory-induced heterogeneity respectively. Herbivory tolerance of clonal fragments with connected rhizomes was higher than that of fragments with severed rhizomes under heterogeneous conditions. These findings confirmed the unconditional advantage of clonal integration on reproduction under the combined influence of resource heterogeneity and simulated herbivory. Moreover, our results made clear the synergistically interactive effects of resource heterogeneity and simulated herbivory on costs and benefits of clonal integration. This will undoubtedly advance our understanding on the plasticity of clonal integration under complex environmental conditions.