How to resolve the SLOSS debate: lessons from species-diversity models.

The SLOSS debate--whether a single large reserve will conserve more species than several small--of the 1970s and 1980s never came to a resolution. The first rule of reserve design states that one large reserve will conserve the most species, a rule which has been heavily contested. Empirical data seem to undermine the reliance on general rules, indicating that the best strategy varies from case to case. Modeling has also been deployed in this debate. We may divide the modeling approaches to the SLOSS enigma into dynamic and static approaches. Dynamic approaches, covered by the fields of island equilibrium theory of island biogeography and metapopulation theory, look at immigration, emigration, and extinction. Static approaches, such as the one in this paper, illustrate how several factors affect the number of reserves that will save the most species. This article approaches the effect of different factors by the application of species-diversity models. These models combine species-area curves for two or more reserves, correcting for the species overlap between them. Such models generate several predictions on how different factors affect the optimal number of reserves. The main predictions are: Fewer and larger reserves are favored by increased species overlap between reserves, by faster growth in number of species with reserve area increase, by higher minimum-area requirements, by spatial aggregation and by uneven species abundances. The effect of increased distance between smaller reserves depends on the two counteracting factors: decreased species density caused by isolation (which enhances minimum-area effect) and decreased overlap between isolates. The first decreases the optimal number of reserves; the second increases the optimal number. The effect of total reserve-system area depends both on the shape of the species-area curve and on whether overlap between reserves changes with scale. The approach to modeling presented here has several implications for conservational strategies. It illustrates well how the SLOSS enigma can be reduced to a question of the shape of the species-area curve that is expected or generated from reserves of different sizes and a question of overlap between isolates (or reserves).