Enhanced biodiversity beyond marine reserve boundaries: the cup spillith over.

Overfishing can have detrimental effects on marine biodiversity and the structure of marine ecosystems. No-take marine reserves (NTMRs) are much advocated as a means of protecting biodiversity and ecosystem structure from overharvest. In contrast to terrestrial protected areas, NTMRs are not only expected to conserve or recover biodiversity and ecosystems within their boundaries, but also to enhance biodiversity beyond their boundaries by exporting species richness and more complex biological communities. Here we show that species richness of large predatory reef fish increased fourfold and 11-fold inside two Philippine no-take marine reserves over 14 and 25 years, respectively. Outside one reserve (Apo) the species richness also increased. This increase beyond the Apo reserve boundary was 78% higher closer to the boundary (200-250 m) than farther from it (250-500 m). The increase in richness beyond the boundary could not be explained by improvements over time in habitat or prey availability. Furthermore, community composition of predatory fish outside but close to (200-250 m) the Apo reserve became very similar to that inside the reserve over time, almost converging with it in multivariate space after 26 years of reserve protection. This is consistent with the suggestion that, as community composition inside Apo reserve increased in complexity, this complexity spilled over the boundary into nearby fished areas. Clearly, the spillover of species richness and community complexity is a direct consequence of the spillover of abundance of multiple species. However, this spillover of species richness and community complexity demonstrates an important benefit of biodiversity and ecosystem export from reserves, and it provides hope that reserves can help to reverse the decline of marine ecosystems and biodiversity.