Abstract
Little theoretical work has investigated how landscape structure affects invasive spread, even though broad-scale disturbances caused by habitat loss and fragmentation are believed to facilitate the spread of exotic species. Neutral landscape models (NLMs), derived from percolation theory in the field of landscape ecology, provide a tool for assessing the risk of invasive spread in fragmented landscapes. A percolation-based analysis of the potential for invasive spread in fragmented landscapes predicts that invasive spread may be enormously enhanced beyond some threshold level of habitat loss, which depends upon the species' dispersal abilities and the degree of habitat fragmentation. Assuming that invasive species spread primarily through disturbed areas of the landscape, poor dispersers may spread better in landscapes in which disturbances are concentrated in space, whereas good dispersers are predicted to spread better in landscapes where disturbances are small and dispersed (i.e., fragmented landscape). Assessing the risk of invasive spread in fragmented landscapes ultimately requires understanding the relative effects of landscape structure on processes that contribute to invasive spread--dispersal (successful colonization) and demography (successful establishment). Colonization success is predicted to be highest when >20% of the landscape has been disturbed, particularly if disturbances are large or aggregated in space, because propagules are more likely to encounter sites suitable for colonization and establishment. However, landscape pattern becomes less important for predicting colonization success if species are capable of occasional long-distance dispersal events. Invasive species are also more likely to persist and achieve positive population growth rates (successful establishment) in landscapes with clumped disturbance patterns, which can then function as population sources that produce immigrants that invade other landscapes. Finally, the invasibility of communities may be greatest in landscapes with a concentrated pattern of disturbance, especially below some critical threshold of biodiversity. Below the critical biodiversity threshold, the introduction of a single species can trigger a cascade of extinctions among indigenous species. The application of NLMs may thus offer new insights and opportunities for the management and restoration of landscapes so as to slow the spread of invasive species.