The boundary-quality penalty: a quantitative method for approximating species responses to fragmentation in reserve selection.

Aggregation of reserve networks is generally considered desirable for biological and economic reasons: aggregation reduces negative edge effects and facilitates metapopulation dynamics, which plausibly leads to improved persistence of species. Economically, aggregated networks are less expensive to manage than fragmented ones. Therefore, many reserve-design methods use qualitative heuristics, such as distance-based criteria or boundary-length penalties to induce reserve aggregation. We devised a quantitative method that introduces aggregation into reserve networks. We call the method the boundary-quality penalty (BQP) because the biological value of a land unit (grid cell) is penalized when the unit occurs close enough to the edge of a reserve such that a fragmentation or edge effect would reduce population densities in the reserved cell. The BQP can be estimated for any habitat model that includes neighborhood (connectivity) effects, and it can be introduced into reserve selection software in a standardized manner. We used the BQP in a reserve-design case study of the Hunter Valley of southeastern Australia. The BQP resulted in a more highly aggregated reserve network structure. The degree of aggregation required was specified by observed (albeit modeled) biological responses to fragmentation. Estimating the effects of fragmentation on individual species and incorporating estimated effects in the objective function of reserve-selection algorithms is a coherent and defensible way to select aggregated reserves. We implemented the BQP in the context of the Zonation method, but it could as well be implemented into any other spatially explicit reserve-planning framework.