Benthic micro- and macro-community succession and coral recruitment under overfishing and nutrient enrichment

Herbivory and nutrient availability are fundamental drivers of benthic community succession in shallow marine systems, including coral reefs. Despite the importance of early community succession for coral recruitment and recovery, studies characterizing the impact of top-down and bottom-up drivers on micro- and macrobenthic communities at scales relevant to coral recruitment are lacking. Here, a combination of tank and field experiments were used to assess the effects of herbivore exclusion and nutrient enrichment on micro- to macrobenthic community succession and subsequent coral recruitment success. Herbivore exclusion had the strongest effect on micro- and macrobenthic community succession, including a community shift toward copiotrophic and potentially opportunistic/pathogenic microorganisms, an increased cover of turf and macroalgae, and decreased cover of crustose coralline algae. Yet, when corals settled prior to the development of a macrobenthic community, rates of post-settlement survival increased when herbivores were excluded, benefiting from the predation refugia provided by cages during their vulnerable early post-settlement stage. Interestingly, survival on open tiles was negatively correlated with the relative abundance of the bacterial order Rhodobacterales, an opportunistic microbial group previously associated with stressed and diseased corals. Development of micro- and macrobenthic communities in the absence of herbivory, however, led to reduced coral settlement. In turn, there were no differences in post-settlement survival between open and caged treatments for corals settled on tiles with established benthic communities. As a result, open tiles experienced marginally higher recruitment rates, driven primarily by the higher initial number of settlers on open tiles compared to caged tiles. Overall, we reveal that the primary interaction driving coral recruitment is the positive effect of herbivory in creating crustose coralline algae (CCA)-dominated habitats, free of fleshy algae and associated opportunistic microbes, to enhance coral settlement. The negative direct and indirect impact of fish predation on newly settled corals was outweighed by the positive effect of herbivory on the initial rate of coral settlement. In turn, the addition of nutrients further altered benthic community succession in the absence of herbivory, reducing coral post-settlement survival. However, the overall impact of nutrients on coral recruitment dynamics was