Unraveling the multiple facilitative effects of consumers on marine primary producers

The loss of consumers threatens the integrity of ecological systems, but the mechanisms underlying the effects on communities and ecosystems remain difficult to predict. This is, in part, due to the complex roles that consumers play in those systems. Here, we highlight this complexity by quantifying two mechanisms by which molluscan grazers-typically thought of as consumers of their algal resources-facilitate algae on rocky shores. Initial observations in high-zone tide pools revealed that both water-column ammonium concentrations and photosynthetic biomass were higher in pools containing higher densities of grazers, suggesting that local-scale nutrient recycling by the grazers could be enhancing algal biomass. We assessed this possibility by experimentally manipulating grazer abundances at the level of whole tide pools but controlling access of those grazers to experimental plots within each pool. Contrary to predictions that algal biomass inside grazer exclusions would increase as grazer abundances in the pools increased, we found that algal biomass inside grazer-exclusion fences was unaffected by grazer abundances. Instead, the consumptive effects of grazers that were evident at low grazer abundances transitioned to facilitative effects as experimentally manipulated grazer abundances increased. This finding suggested that these positive interactions were associated with the physical presence of grazers and not just grazers' effects on nutrient availability. Subsequent experiments highlighted the potential role of "slime"-the pedal mucous trails left behind as the mollusks crawl on the substratum-in promoting the recruitment of algae and thereby mediating a spatial subsidy of new organic matter into the system. Furthermore, different grazer groups contributed disproportionately to ammonium excretion (i.e., turban snails) versus slime production (i.e., littorine snails), suggesting a potential role for grazer diversity. Our work highlights the complex ways in which consumers affect their resources, including multiple, complementary mechanisms by which these grazers facilitate the algae they consume.