Colonization pressure: a second null model for invasion biology.
Understanding the causes of spatial variation in the distribution and richness of alien species is a key goal of invasion biology. Thanks to the increasing availability of geographical compendia of alien species it is also the subject of a burgeoning scientific literature. Here, we review elements of this literature to argue that understanding the causes of variation in alien species richness cannot be achieved without knowledge of the number of species introduced to an area-termed colonization pressure. The implications of a lack of information on colonization pressure are widespread and, we believe, poorly recognised. We start by discussing a recent general model for alien species richness that demonstrates why colonization pressure is a fundamental determinant of this. We then explore the literature on alien species richness, particularly on islands, to demonstrate how failing to account for colonization pressure affects our perception of richness drivers. We further show that ignoring colonization pressure can bias our understanding of patterns in the distributions of alien species. We finish by discussing situations when colonization pressure may be more or less important for our understanding of alien invasions.