Effects of reef attributes on fish assemblage similarity between artificial and natural reefs

Artificial reefs are used to enhance populations of marine organisms, but relatively few studies have quantitatively evaluated which attributes of reef structure are most critical in determining whether assemblages of organisms on artificial reefs are similar to those on natural reefs. Using five pairs of artificial and natural reefs that spanned 225 km in the Southern California Bight, we evaluated how well fish assemblages on artificial reefs mimicked those on natural reefs and which attributes of reefs best predicted assemblage structure. Along underwater visual transects, we quantified fish species richness, density, and size structure, as well as substrate structure (rugosity and cover of substrate types), giant kelp density, and invertebrate density. Artificial reefs that were more similar in physical structure to natural reefs (low relief, low rugosity, and composed of small-to medium-sized boulders) supported fish assemblages that were similar to those on natural reefs. Fish species richness was not significantly different between artificial and natural reefs, but density and biomass tended to be higher on average on artificial reefs, body size was slightly smaller, and assemblage structure differed between the two reef types. Generally, artificial reefs extended higher off the seabed, were made of larger boulders, had higher rugosity, harboured more invertebrates, and supported less giant kelp. At both the within-reef (transect) and whole-reef scales, fish density and biomass were positively correlated with complex substrate structure, positively correlated with invertebrate density, and negatively correlated with giant kelp abundance, which was sparse or absent on most artificial reefs. Our results indicate that artificial reefs can support fish assemblages that are similar to those found on natural reefs if they are constructed to match the physical characteristics of natural reefs, or they can be made to exceed natural reefs in some regards at the expense of other biological attributes.