Decadal-scale effects of benthic habitat and marine reserve protection on Philippine goatfish (F: Mullidae)

Reef fish populations can be affected by both fishing and changes in benthic habitat. Yet, partitioning these effects is often difficult, usually requiring an appropriate sampling design and long-term monitoring. Here we quantify, over a 30-yr period, the effects of benthic habitat change and no-take marine reserve (NTMR) protection on the density and species richness of a lightly harvested benthic-feeding reef fish family, the Mullidae (goatfish), at four Philippine islands. Boosted regression trees demonstrated that goatfish density and species richness had strong negative associations with hard coral cover and strong positive associations with cover of dead substratum. No-take marine reserve protection had no effect on the density or species richness of goatfish over 19 and 30 yr at Sumilon and Apo islands, respectively. However, environmental disturbances (e.g., typhoons, coral bleaching) that reduced hard coral cover subsequently led to increases in goatfish numbers for periods ranging from 2 to 8 yr. After initial increases due to benthic disturbance, goatfish populations decreased during coral recovery, occurring on timescales of 10-20 yr. This long-term, "natural experiment" demonstrated that changes to benthic habitat (bottom-up control) had a far greater effect on Philippine goatfish populations than protection from fishing (a top-down effect) in NTMRs. Given the strong positive response of goatfish populations to loss of live hard coral cover, this group of fishes may be a valuable indicator species for habitat degradation on coral reefs