Flying foxes create extensive seed shadows and enhance germination success of pioneer plant species in deforested Madagascan landscapes.

Seed dispersal plays a significant role in forest regeneration and maintenance. Flying foxes are often posited as effective long-distance seed dispersers due to their large home ranges and ability to disperse seeds when flying. We evaluate the importance of the Madagascan flying fox Pteropus rufus in the maintenance and regeneration of forests in one of the world's priority conservation areas. We tested germination success of over 20,000 seeds from the figs Ficus polita, F. grevei and F. lutea extracted from bat faeces and ripe fruits under progressively more natural conditions, ranging from petri-dishes to outdoor environments. Seeds from all fig species showed increased germination success after passing through the bats' digestive tracts. Outside, germination success in F. polita was highest in faecal seeds grown under semi-shaded conditions, and seeds that passed through bats showed increased seedling establishment success. We used data from feeding trials and GPS tracking to construct seed shadow maps to visualize seed dispersal patterns. The models use Gaussian probability density functions to predict the likelihood of defecation events occurring after feeding. In captivity, bats had short gut retention times (often < 30 mins), but were sometimes able to retain seeds for over 24h. In the wild, bats travelled 3-5 km within 24-280 min after feeding, when defecation of ingested seeds is very likely. They produced extensive seed shadows (11 bats potentially dispersing seeds over 58,000 ha over 45 total days of tracking) when feeding on figs within their large foraging areas and dispersed the seeds in habitats that were often partially shaded and hence would facilitate germination up to 20 km from the feeding tree. Because figs are important pioneer species, P. rufus is an important dispersal vector that makes a vital contribution to the regeneration and maintenance of highly fragmented forest patches in Madagascar.