Does residence time affect responses of alien species richness to environmental and spatial processes?

One of the most robust emerging generalisations in invasion biology is that the probability of invasion increases with the time since introduction (residence time). We analysed the spatial distribution of alien vascular plant species in a region of north-eastern Italy to understand the influence of residence time on patterns of alien species richness. Neophytes were grouped according to three periods of arrival in the study region (1500–1800, 1800–1900, and > 1900). We applied multiple regression (spatial and nonspatial) with hierarchical partitioning to determine the influence of climate and human pressure on species richness within the groups. We also applied variation partitioning to evaluate the relative importance of environmental and spatial processes. Temperature mainly influenced groups with species having a longer residence time, while human pressure influenced the more recently introduced species, although its influence remained significant in all groups. Partial regression analyses showed that most of the variation explained by the models is attributable to spatially structured environmental variation, while environment and space had small independent effects. However, effects independent of environment decreased, and spatially independent effects increased, from older to the more recent neophytes. Our data illustrate that the distribution of alien species richness for species that arrived recently is related to propagule pressure, availability of novel niches created by human activity, and neutral-based (dispersal limitation) processes, while climate filtering plays a key role in the distribution of species that arrived earlier. This study highlights the importance of residence time, spatial structure, and environmental conditions in the patterns of alien species richness and for a better understanding of its geographical variation.