The Farther the Better: Effects of Multiple Environmental Variables on Reef Fish Assemblages along a Distance Gradient from River Influences

The conservation and management of site-attached assemblages of coastal reefs are particularly challenging because of the tremendous environmental variation that exists at small spatial scales. In this sense, understanding the primary sources of variation in spatial patterns of the biota is fundamental for designing effective conservation policies. We investigated spatial variation in fish assemblages around the windward and leeward sides of coastal islands situated across a gradient of riverine influence (13 km in length). Specifically, relationships between rocky reef fish assemblages and benthic, topographic and physical predictors were assessed. We hypothesized that river induced disturbances may overcome local habitat features in modeling spatial patterns of fish distribution. Fish assemblages varied primarily due to the strong directional gradient of riverine influence (22.6% of the estimated components of variation), followed by topographic complexity (15%), wave exposure (9.9%), and benthic cover (8%). The trophic structure of fish assemblages changed from having a high abundance of invertebrate feeders in macroalgae-dominated reefs close to river mouths to a high proportion of herbivores, planktivores and invertebrate feeder species in reefs with large boulders covered by epilithic algal matrices, as the distance from rivers increased. This gradient led to an increase of 4.5-fold in fish richness and fish trophic group diversity, 11-fold in fish biomass and 10-fold in fish abundance. Our results have implications for the conservation and monitoring of assemblages patchily distributed at small spatial scales. The major role of distance from river influences on fish assemblages rather than benthic cover and topographic complexity suggest that managing land-based activities should be a conservation priority toward reef restoration.