Limited directed seed dispersal in the canopy as one of the determinants of the low hemi-epiphytic figs' recruitments in Bornean rainforests

Ficus species are keystone plants in tropical rainforests, and hemi-epiphytic figs play a notably important role in forest ecosystems. Because hemi-epiphytic figs have strict germination requirements, germination and establishment stages regulate their populations. Despite the ecological importance of hemi-epiphytic figs in the rainforests, seed dispersal systems by fig-eating animals under natural conditions remain unknown because of the difficulty in tracing the destiny of dispersed seeds in the canopy. Therefore, seed dispersal effectiveness (SDE) has never been evaluated for hemi-epiphytic figs. We evaluated the SDE of hemi-epiphytic figs using qualitative and quantitative components by three relatively large-sized (> 3 kg) arboreal and volant animals in Bornean rainforests that largely depend on fig fruits in their diets: binturongs Arctictis binturong, Mueller's gibbons Hylobates muelleri, and helmeted hornbills Rhinoplax vigil. The SDE values of binturongs was by far the highest among the three study animals. Meanwhile, successful seed dispersal of hemi-epiphytic figs by gibbons and helmeted hornbills is aleatory and rare. Given that seed deposition determines the fate of hemi-epiphytic figs, the defecatory habits of binturongs, depositing feces on specific microsites in the canopy, is the most reliable dispersal method, compared to scattering feces from the air or upper canopy. We showed that reliable directed dispersal of hemi-epiphytic figs occurs in high and uneven canopy of Bornean rainforests. This type of dispersal is limited to specific animal species, and therefore it may become one of the main factors regulating low-success hemi-epiphytic fig recruitment in Bornean rainforests.