Effects of Soil Characteristics, Allelopathy and Frugivory on Establishment of the Invasive Plant Carpobrotus edulis and a Co-Occuring Native, Malcolmia littorea

Background: The species Carpobrotus edulis, native to South Africa, is one of the major plant invaders of Mediterranean coastal ecosystems around the world. Invasion by C. edulis exerts a great impact on coastal habitats. The low number of native species in invaded communities points to the possible existence of mechanisms suppressing their germination. In this study we assessed whether soil factors, endozoochory, competition and allelopathic effects of the invader affect its own early establishment and that of the native species Malcolmia littorea. We used laboratory solutions representing different chemical composition and moisture of the soil, herbivore feeding assays to simulate seed scarification and rainwater solutions to account for the effect of differently aged C. edulis litter. Principal Findings: We show that unlike that of the native species, germination and early growth of C. edulis was not constrained by low moisture. The establishment of C. edulis, in terms of germination and early growth, was increased by scarification of seeds following passage through the European rabbit intestines; the rabbits therefore may have potential implications for plant establishment. There was no competition between C. edulis and M. littorea. The litter of the invasive C. edulis, which remains on the soil surface for several years, releases allelopathic substances that suppress the native plant germination process and early root growth. Conclusions: The invasive species exhibits features that likely make it a better colonizer of sand dunes than the co-occurring native species. Allelopathic effects, ability to establish in drier microsites and efficient scarification by rabbits are among the mechanisms allowing C. edulis to invade. The results help to explain the failure of removal projects that have been carried out in order to restore dunes invaded by C. edulis, and the long-lasting effects of C. edulis litter need to be taken into account in future restoration projects.