Stratification along tropical forest succession enhances pollinator diversity via functionally unique canopies

Vertical stratification is a prominent driver of biodiversity in forests. As the proportion of successional forests increases worldwide and canopies are lost at a fast pace, it is crucial to understand the role of stratification as a driver of community recovery and ecological processes during succession, particularly for important plant mutualist groups essential for forest recovery. Within a well-resolved chronosequence in the northwestern Ecuadorian rainforest, we compiled an extensive database of over 20,000 diurnal and nocturnal pollinators and 2,000 interactions with plants and examined the interacting effects of recovery age and stratification on pollinator community and interaction network reassembly. In late successional and old-growth forests, stratification was a stronger predictor of pollinator abundance, alpha-diversity, functional richness and beta-diversity than forest legacy. While most groups were strongly associated with canopies (moths, social bees, and nocturnal bees), orchid bees exhibited an inverse pattern, also presenting the strongest functional response to stratification and recovery status. Within the entire chronosequence (0-38 years of recovery, plus old-growth forests), recovery status and stratification, alongside landscape features, acted synergistically as predictors of pollinator community parameters. Although structurally stable across the chronosequence and between strata, Interaction networks were richer and most distinct in canopies, while the highest pollinator and interaction diversity were found in old-growth canopies. In successional forests, networks were only comparable in size to active disturbances and early recovery with both strata combined, whereas complete networks in old-growth forests were more than twice as large as those in other recovery statuses. Our study underlines the importance of stratification in leveraging pollinator diversity during forest succession. By serving as havens for canopy-adapted species, tropical canopies may thus help safeguard pollination processes in disturbed landscapes.