Herbivory induced non-local responses of the clonal invader Carpobrotus edulis are not mediated by clonal integration

The anthropogenic displacement of species around the world results in new environmental situations where native and exotic species coexist. Exotic plants have to face native herbivores, and interactions between introduced plants and native herbivores seem to play an important role in the invasiveness of some exotic plant species. We studied the role of clonal integration in induce morphological, physiological, and biochemical responses in the clonal invader Carpobrotus edulis against the attack of the native snail Theba pisana. Our results demonstrated the presence of labour division mediated by physiological integration, with a significant increase of photosynthesis potential (both at morphological and physiological) in un-attacked integrated ramets. This response could be especially important under herbivory, as the negative impact of T. pisana on the photosynthetic structures of attacked C. edulis ramets could be buffered by transferring the dependence of photosynthetic activity to the un-attacked ramets. Our results also showed a constitutive resistance in un-attacked apical ramets, showing a similar amount of defence compounds to those exhibited in the basal branches attacked by snails. Results reported a non-local compensatory response, which there was an increase of total biomass in apical ramets when their basal ramets were attacked by the herbivore. We interpret this result as a compensatory response, with these apical ramets increasing shoot biomass to compensate for the biomass loss due to a potential attack from herbivores. However, this non-local response was not mediated by physiological integration but probably due to belowground communication, with the presence of alarm signals released by root exudates. We conclude that the attack of this snail is not enough to be a possible biological control due to the compensatory response to this snail by C. edulis, favouring their expansion. Future studies should focus on unravelling the role of belowground communication in the defensive responses of C. edulis.