Seed dispersal of alien and native plants by vertebrate herbivores

Seed dispersal is crucial for the success and spread of alien plants. Herbivores often establish a dual relationship with plants: antagonist, through herbivory, and mutualist, through seed dispersal. By consuming plants, herbivores may disperse large amounts of seeds, and can facilitate the spread of alien plants. However, seed dispersal of alien plants by herbivores has been largely uninvestigated. I studied factors associated with dispersal of alien and native seeds by the three most important vertebrate herbivores in SW Australia: emus (Dromaius novaehollandia), western grey kangaroos (Macropus fuliginosus) and European rabbits (Oryctolagus cuniculus). Overall frequencies of alien and native seeds dispersed by these herbivores were determined by differences among them in (1) the plant groups they predominantly disperse, that differed in frequencies of aliens versus natives, and (2) the predominant dispersal of aliens or natives within those plant groups. Emus and kangaroos (natives) tended to disperse predominantly alien seeds within plant groups (defined by life forms, dispersal syndromes, and diaspore size), whereas rabbits (alien) tended to disperse predominantly natives. This agrees with the hypothesis that herbivores will use predominantly plants that have evolved in different areas, because of less effective defences against new enemies. Overall frequencies were consistent with this pattern in kangaroos and rabbits, but not in emus. Kangaroos dispersed mostly plant groups that were mainly aliens (herbaceous species and small and medium sized dispersal units and seeds), which together with their predominant use of aliens over natives within groups resulted in the highest overall frequency of alien seeds (73%). Rabbits were similar to kangaroos in the type of plants dispersed, but their predominant use of natives over aliens within groups contributed to an overall predominance of native seeds in their pellets (88%). Emus dispersed mostly plant groups that were mainly natives (e.g. woody species with big diaspores), resulting in low overall frequency of alien seeds (11%), despite their predominant use of aliens over natives within plant groups. Thus, the within-groups trend pointed to a facilitative role of native herbivores of plant invasions through seed dispersal, but was obscured by the different use by herbivores of plant groups with different frequency of aliens.