Nutrient pollution and fishing are the primary local causes of coral reef decline but their ecosystem-scale effects are poorly understood. Results from small-scale manipulative experiments of herbivores and nutrients suggest prioritizing management of fishing over nutrient pollution because herbivores can control macroalgae and turf in the presence of nutrients. However, ecological theory suggests that the opposite occurs at large scales. Moreover, it is unclear whether fishing decreases herbivores because fishing of predators may result in an increase in herbivores. To investigate this paradox, data on the fish and benthic communities, fishing, and nutrients were collected on Kiritimati, Kiribati. Oceanographic conditions and a population resettlement program created a natural experiment to compare sites with different levels of fishing and nutrients. Contrary to theory, herbivores controlled macroalgae in the presence of nutrients at large spatial scales, and herbivores had greater effects on macroalgae when nutrients were higher. In addition, fishing did not increase herbivores. These results suggest that protecting herbivores may have greater relative benefits than reducing nutrient pollution, especially on polluted reefs. Reallocating fishing effort from herbivores to invertivores or planktivores may be one way to protect herbivores and indirectly maintain coral dominance on reefs impacted by fishing and nutrient pollution. 1. Introduction Fishing [1–7] and nutrient pollution [8, 9] or both [10–14] are cited as the most important local causes of coral reef decline. It is difficult, however, to evaluate local fishing and nutrient effects independently because these factors are almost always confounded and large-scale experiments are infeasible. Results from theoretical and small-scale experimental studies (typically <1？m2, four from 50 to 250？m2 [14]) suggest prioritizing management of herbivore populations because herbivores can control the effect of nutrients on macroalgal and turf abundance and nutrient enrichment alone is not sufficient to cause a phase shift from coral to macroalgal and/or turf algal dominance [11, 14, 15]. In order to apply these results and implement ecosystem-based management, information is needed on (1) how fishing and nutrients interactively affect the fish and benthic communities, (2) the mechanisms by which fishing (rather than cages that exclude fish) and nutrients are linked to shifts to macroalgal and turf algal dominance, and (3) whether herbivores can control macroalgae and turf algae when nutrient enrichment
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