Trophic interactions and feedbacks maintain intact and degraded states of Hawaiian tropical forests

Feedbacks within ecosystems can lead to internal reinforcement of the current state providing ecosystem resilience. Often, multiple biotic interactions across trophic levels play a role in such feedbacks, yet these are generally studied independently, obscuring the relative importance of interactions among different factors. We look at various potential feedbacks in intact and degraded mesic forests on Hawaiʻi Island where managers have planted native Acacia koa (koa) trees in an attempt to jumpstart succession in former cattle pastures. These restoration forests, however, have not undergone secondary succession, instead maintaining a koa overstory with an exotic pasture grass understory. We contrasted different trophic level processes that influence the capacity for natural understory regeneration: feedbacks between bird‐mediated seed rain and fruiting understory (“top‐down”), as well as links between understory composition and microhabitats for native seed germination (“bottom‐up”). We quantified bird‐mediated seed rain under canopy trees along transects spanning intact, fragmented, and restoration forests. Along these transects, we established plots around focal overstory trees to measure abundance of fruiting understory species, ground cover (e.g., exotic grass, bryophyte), and obtained estimates of bird density to evaluate the contribution of each of these factors to seedling abundance. We also used a factorial seed addition/grass removal experiment to directly compare the influence of seed rain and germination substrate. We found evidence of both top‐down and bottom‐up feedbacks that reinforced the current state of each forest type. In the intact and fragmented forests, the combination of comparatively high seed rain and ample germination substrate is likely critical for maintaining a diverse forest system. In contrast, exotic grasses exhibit priority effects in restoration forests, inhibiting seed germination and effectively negating any benefits that could be derived from bird‐mediated seed rain. Such internal reinforcement suggests that active, rather than passive, restoration would be beneficial to increase forest diversity in restoration areas.