Macroecological drivers of alien conifer naturalizations worldwide

Understanding the factors that drive the global distribution of alien species is a pivotal issue in invasion biology. Here, we used data on naturalized conifers (Pinaceae, Cupressaceae) from sixty temperate and subtropical regions and five continents to test how environmental and socio-economic conditions of recipient areas as well as introduction efforts affect naturalization probabilities. We collated 18 predictor variables for each region describing environmental, biogeographic and socio-economic conditions as well as a measure of the macro-climatic match with the species' native ranges, and the extent to which alien conifers are used in commercial forestry. Naturalization probabilities across all species and regions were then related to these predictor variables by means of generalized linear mixed models. For both Pinaceae and Cupressaceae, naturalization probabilities were generally higher in the Southern Hemisphere, and increased with indicators of habitat diversity of the recipient region. The match in macro-climatic conditions between the native and introduced regions was a significant predictor of conifer naturalization, but socio-economic variables were less powerful predictors. Only for Cupressaceae did a socio-economic variable (human population density) affect naturalization probabilities. Key attributes facilitating naturalization were related to introduction effort. Moreover, usage in commercial forestry generally fostered naturalization, although the actual size of alien conifer plantations in a region was only correlated with the naturalization of Pinaceae. Our results suggest that climate matching, habitat diversity and introduction effort co-determine the probability of naturalization, which additionally, is modulated by biogeographic features of the recipient area, such as incidence of natural enemies or competitors. To date, the most widely used tools for invasive plant risk assessment only account for climate match and rarely factor in other attributes of the recipient environment. Future tools should additionally consider biotic environment and introduction effort if risk assessment is to be effective.