Trophic rewilding benefits a tropical community through direct and indirect network effects

Species reintroductions can be used as a conservation strategy to restore ecological interactions and the functionality of impoverished ecosystems. The ecological effects of reintroductions go beyond restoring pairwise interactions, because reintroductions can change how extant species are indirectly linked to each other in an ecological community. These indirect pathways, in turn, may shape a myriad of ecological and evolutionary processes operating in ecological systems. Here, we investigated how reintroductions may affect the direct and indirect pathways connecting species in ecological networks. We modeled the potential effects of the reintroduction of four frugivore species (channel-billed toucans, red-humped agoutis, brown howler monkeys and yellow-footed tortoises) to the local seed dispersal network in an Atlantic Forest site, the Tijuca National Park (Tijuca, Brazil). We used a seed dispersal interaction dataset together with data on species occurrences in Tijuca to build network models. Then, we calculated how network structure and the total amount of indirect effects varied across simulated networks with and without the reintroduced species. Using random reintroduction simulations, we tested if the observed network changes were expected merely from the increase in species richness. The reintroduction of the frugivore species increased network connectance, nestedness, robustness, number of pathways and total amount of indirect effects in all simulated networks. The increase in number of pathways with the addition of the four reintroduced species was greater than the sum of isolated effects for each species, as some interaction pathways contained several reintroduced species. These changes in network metrics were significantly greater than if a randomly chosen set of four species were reintroduced. Furthermore, our results indicate that multiple reintroductions in the same area, known as refaunation, may have an even greater restoration effect than single species reintroductions through increased indirect connections in the network.