This resilience could be caused by multiple processes: feedbacks between birds, seeds, and understory (seed limitation), understory and germination habitats (establishment limitation), or a combination of the two. A lack of food (i.e. fruit) and suitable habitat (i.e. structure) often discourages birds from visiting degraded sites. Because birds disperse seeds that are critical for tree and understory plant recruitment, this lack of bird visitation can lead to reinforcement of a degraded state. In addition, in degraded tropical forests, suitable germination sites are often limited by the presence of highly competitive exotic pasture grasses, which can stall natural forest regeneration even when seed dispersal occurs.
We explored the influence of both of these processes on the capacity for natural understory regeneration in Hawaiian forests. We quantified bird-mediated seed rain (deposition) under canopy trees along transects spanning intact, fragmented, and restoration forests. We also established plots around focal overstory trees to measure abundance of fruiting understory species and ground cover (e.g., exotic grass, moss), and we obtained estimates of bird density to evaluate the contribution of each of these factors to seedling abundance. We also used a seed addition/grass removal experiment to directly compare the influence of seed rain and ground cover. We found evidence of multiple feedbacks that reinforced the current state of each forest type.