Community regulation by herbivore parasitism and density: Trait-mediated indirect interactions in the intertidal

The abundant herbivorous mud-snail Hydrobia ulvae is an ecosystem engineer in soft-bottom intertidal habitats due to its grazing and bioturbation activity. However, mud snails are commonly infected by trematodes that reduce their overall activity, which in turn may affect their impact on the surrounding benthic community. To test this hypothesis, we performed field experiments manipulating both the abundance of uninfected snails (0, 7500 and 15.000 ind. m^- 2) and the level of snail parasitism (0, 33 and 100% trematode prevalence) on a Danish mud-flat. The results showed that increasing snail abundance and parasitism generally had opposite effects on the community of microphytobenthos and zoobenthos. Increasing snail density increased the chlorophyll-a concentration in the substrate (enhancement), whereas increasing parasitism decreased it. In accordance, the benthic primary producers were generally less nutrient limited at high snail density and mostly so at high levels of snail parasitism. Moreover, epipsammic diatoms were favoured over epipelic diatoms at increasing snail density, whereas the opposite was evident at increasing snail parasitism. At the community level, increasing snail density increased evenness among epipelic diatoms, whereas increasing snail parasitism decreased evenness and species diversity. Probably through the action of trophic cascades and varying levels of disturbance, the zoobenthic community was influenced by experimental treatments as well. The indirect effects of snail parasitism influenced significantly the abundance of more faunal species (seven) than did snail density (two). At the community level, increasing snail density decreased evenness and lowest species richness coincided with intermediate snail density. In contrast, increasing snail parasitism resulted in increasing evenness and peaking species richness at intermediate level of parasitism. Together, the results show that parasites solely through their impact on the behaviour of a single community member can be significant indirect determinants of community organisation and function.