Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.

Abstract. The degree of dietary generalism among consumers has important consequences for population, community, and ecosystem processes, yet the effects on consumer fitness of mixing food types have not been examined comprehensively. We conducted a meta-analysis of 161 peer-reviewed studies reporting 493 experimental manipulations of prey diversity to test whether diet mixing enhances consumer fitness based on the intrinsic nutritional quality of foods and consumer physiology. Averaged across studies, mixed diets conferred significantly higher fitness than the average of single-species diets, but not the best single prey species. More than half of individual experiments, however, showed maximal growth and reproduction on mixed diets, consistent with the predicted benefits of a balanced diet. Mixed diets including chemically defended prey were no better than the average prey type, opposing the prediction that a diverse diet dilutes toxins. Finally, mixed-model analysis showed that the effect of diet mixing was stronger for herbivores than for higher trophic levels. The generally weak evidence for the nutritional benefits of diet mixing in these primarily laboratory experiments suggests that diet generalism is not strongly favored by the inherent physiological benefits of mixing food types, but is more likely driven by ecological and environmental influences on consumer foraging.

In coastal marine food webs, small invertebrate herbivores (mesograzers) have long been hypothesized to occupy an important position facilitating dominance of habitat-forming macrophytes by grazing competitively superior epiphytic algae. Because of the difficulty of manipulating mesograzers in the field, however, their impacts on community organization have rarely been rigorously documented. Understanding mesograzer impacts has taken on increased urgency in seagrass systems due to declines in seagrasses globally, caused in part by widespread eutrophication favoring seagrass overgrowth by faster-growing algae. Using cage-free field experiments in two seasons (fall and summer), we present experimental confirmation that mesograzer reduction and nutrients can promote blooms of epiphytic algae growing on eelgrass (Zostera marina). In this study, nutrient additions increased epiphytes only in the fall following natural decline of mesograzers. In the summer, experimental mesograzer reduction stimulated a 447% increase in epiphytes, appearing to exacerbate seasonal dieback of eelgrass. Using structural equation modeling, we illuminate the temporal dynamics of complex interactions between macrophytes, mesograzers, and epiphytes in the summer experiment. An unexpected result emerged from investigating the interaction network: drift macroalgae indirectly reduced epiphytes by providing structure for mesograzers, suggesting that the net effect of macroalgae on seagrass depends on macroalgal density. Our results show that mesograzers can control proliferation of epiphytic algae, that top-down and bottom-up forcing are temporally variable, and that the presence of macroalgae can strengthen top-down control of epiphytic algae, potentially contributing to eelgrass persistence.

Abstract. Latitudinal gradients in species interactions are widely cited as potential causes or consequences of global patterns of biodiversity. However, mechanistic studies documenting changes in interactions across broad geographic ranges are limited. We surveyed predation intensity on common prey (live amphipods and gastropods) in communities of eelgrass (Zostera marina) at 48 sites across its Northern Hemisphere range, encompassing over 37°of latitude and four continental coastlines. Predation on amphipods declined with latitude on all coasts but declined more strongly along western ocean margins where temperature gradients are steeper. Whereas in situ water temperature at the time of the experiments was uncorrelated with predation, mean annual temperature strongly positively predicted predation, suggesting a more complex mechanism than simply increased metabolic activity at the time of predation. This large-scale biogeographic pattern was modified by local habitat characteristics; predation declined with higher shoot density both among and within sites. Predation rates on gastropods, by contrast, were uniformly low and varied little among sites. The high replication and geographic extent of our study not only provides additional evidence to support biogeographic variation in predation intensity, but also insight into the mechanisms that relate temperature and biogeographic gradients in species interactions.