RESEARCH ARTICLE Spatial spread of an alien tree species in a heterogeneous forest landscape: a spatially realistic simulation model

Abstract The effect of environmental heterogeneity on spatial spread of invasive species has received little attention in the literature. Altering landscape heteroge-neity may be a suitable strategy to control invaders in man-made landscapes. We use a population-based, spatially realistic matrix model to explore mechanisms underlying the observed invasion patterns of an alien tree species, Prunus serotina Ehrh., in a heterogeneous managed forest. By altering several parameters in the simulation, we test for various hypotheses regarding the role of several mechanisms on invasion dynamics, including spatial heterogeneity, seed dispersers, site of first introduction, large-scale natural disturbances, and forest management. We observe that landscape hetero-geneity makes the invasion highly directional resulting from two mechanisms: (1) irregular jumps, which occur rarely via long-distance dispersers and create new founder populations in distant suitable areas, and (2) regular, continuous diffusion toward adjacent cells via short- and mid-distance vectors. At the landscape scale, spatial heterogeneity increases the invasion speed but decreases the final invasion extent. Hence, natural disturbances (such as severe storms) appear to facilitate invasion spread, while forest management can have contrasting effects such as decreasing invasibility at the stand scale by increasing the proportion of light interception at the canopy level. The site of initial introduction influences the invasion process but without altering the final outcome. Our model represents the real landscape and incorporates the range of dispersal modes, making it a powerful tool to explore the interactions between environmental heterogeneity and invasion dynamics, as well as for managing plant invaders