Effects of a recalcitrant understory fern layer in an enclosed tropical restoration experiment

Abstract

Establishing mixed-species tree plantings and fencing them to protect seedlings from herbivory is a valuable strategy for reconnecting forest fragments separated by agropastoral lands. However, fencing may provide exotic plants with the escape from herbivory required to invade the understory of planted communities. Here we take advantage of such a situation to ask how the identity of planted species and the resulting canopy cover influenced invasion success by the Asian swordtail fern (Nephrolepis brownii Desv. Nephrolepidaceae) in a 13-year-old tropical restoration experiment. Through a seed addition experiment, we also evaluated the effects the ferns had on recruiting seedlings. We found that the invasion was most acute in the unplanted control plots where canopy cover was consistently scarce. Frond density correlated negatively with canopy cover, though most of the variance in the model is explained by the design of our experiment (r2m = .161, r2c = .460). Between planting treatments that differed in the dispersal mode of the planted trees, the wind-dispersed treatment had higher fern density and longer fronds than the animal-dispersed treatment. The animal-dispersed treatment had the highest recruiting species richness, which was negatively correlated with fern density (r2 = .748). The seed addition experiment confirmed that mortality rates increased where frond density was higher (F1,41 = 7.159, p = .011) and germination rates were lowered for the smaller-seeded species (F1,42 = 13.2, p = .002). To prevent recalcitrant understory layers from establishing in plantings in the future, we recommend: (1) establishing larger plantings or expanding existing ones to minimize edge effects (particularly light filtration), (2) supplementing young plantings with additional seedlings to prevent canopy gaps from forming, and (3) planting an assemblage of species that cover the full forest strata and have consistently full tree-canopies.