Secondary invasion by an invasive shrub limits native tree seedling growth in a size-dependent manner following the eradication of a different invasive species in an island ecosystem

Following the control of an invasive tree species, native forest ecosystem restoration can be prevented by a secondary invasion by another invasive species that inhibits the reestablishment of native trees. In this study, we evaluated the effects of the invasive shrub Leucaena leucocephala on the initial growth of planted seedlings of four native tree species (Pandanus boninensis, Rhaphiolepis indica, Schima mertensiana, and Terminalia catappa) after eradication of the invasive tree Casuarina equisetifolia in the Ogasawara Islands in the northwestern Pacific Ocean. In a 24-month field experiment, we compared the sizes of planted seedlings (diameter at ground level, height, and leaf number) among four treatments in a 2 x 2 design of initial seedling size (large vs. small) x L. leucocephala invasion (sowing L. leucocephala seeds vs. control). More than 75% of the seedlings (excluding S. mertensiana) survived regardless of the treatments. For all four species, initially large seedlings tended to become larger plants at the end of the experiment than initially small seedlings. We detected significant effects of L. leucocephala invasion on one or more plant size parameters in all four species. Seedlings grown in L. leucocephala invasion quadrats were smaller than those in control quadrats at the end of the experiment. We detected significant interactions between initial seedling sizes and L. leucocephala invasion for three species. Together, these results indicate that secondary invasion by L. leucocephala after controlling C. equisetifolia inhibits the initial growth of native tree seedlings, depending on their size at the time of invasion.