Going further on herbivore fishing: the removal of smaller fishes from algal-dominated reefs

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Abstract
Overfishing of large herbivorous fishes is connected to the rise of algal-dominated states on coral reefs. The recovery of their populations is challenging, and future herbivore assemblages may be composed of smaller fish. With fisheries now targeting these smaller-sized herbivore populations, coral reef benthic communities may face unknown outcomes. We performed caging experiments in algal-dominated reefs of Northeastern Brazil, that have been depleted of large herbivorous fishes, to appraise the effects of removing small herbivores on benthic community composition and succession. Full cages simulated herbivore removal, and partial cages and open plots functioned as controls. In total, 36 experimental plots were monitored for 1 yr, accounting for the influence of seasonal changes in local conditions of temperature and turbidity. Overall macroalgal cover did not change between experimental treatments, but filamentous algae increased 5-fold inside full cages by the end of the experiment, surpassing articulated coralline forms as the dominant group. Higher temperatures during the dry season promoted filamentous algae when the top-down control of the herbivores was removed, while a reverse pattern was observed when fishes were allowed to feed inside plots. Small herbivores accelerated benthic succession, facilitating the dominance of articulated coralline algae as the climax community. Our findings oppose previous studies performed at sites with high abundances of large-bodied fishes, where herbivory decreased overall macroalgal cover, promoted filamentous and crustose coralline algae and delayed community succession. The further depletion of smaller-bodied herbivores can trigger shifts in benthic community dynamics that interact with water temperature, which may have implications for reef resilience in an ocean-warming scenario.

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