Trait-associated loss of frugivores in fragmented forest does not affect seed removal rates

1. Seed dispersal by frugivorous animals forms the basis for regeneration of numerous plant species. Habitat fragmentation has been found to be one major factor perturbing frugivore communities and dependent plant species. Yet, community-wide consequences of fragmentation for both frugivore and plant communities are still hardly understood. 2. Here, we studied the effects of habitat fragmentation on the seed removal by frugivorous birds and mammals from nine fleshy-fruited plant species in Bialowieza Forest (Eastern Poland). This last relict of old-growth lowland forest in Europe poses an exceptional reference site for studying the impact of habitat fragmentation on seed dispersal processes in temperate forest ecosystems. 3. In particular, (i) we tested for associations between forest fragmentation and response traits of frugivores, that is forest specialization and body mass; (ii) we studied the relationship between frugivore response and effect traits, that is centrality (number of consumed plant species) and interaction type (mutualistic vs. antagonistic); and (iii) we assessed the feedback of fragmentation-induced changes on plant-frugivore interactions and seed removal rates. 4. We found that fragmentation led to shifts in the frugivore community, associated with the response traits forest specialization and body mass, with fewer forest specialists and large-bodied frugivores in fragmented than in continuous forests. However, forest generalists and small-bodied frugivores were more central in the plant-frugivore associations than forest specialists and large-bodied frugivores. Therefore, the loss of vulnerable species did not result in reduced seed removal rates in fragmented compared with continuous forest. 5. Synthesis. These results indicate that seed removal may be relatively robust in spite of shifts in the frugivore community in forest fragments. The correlation between response and effect traits of frugivores highlights the importance of forest generalists and small-bodied frugivores for maintaining dispersal processes in fragmented forests in temperate regions. Yet, future studies should aim at quantifying the consequences of seed disperser loss on other aspects of dispersal, such as long-distance dispersal, spatial patterns of seed deposition, seed germination and plant regeneration.