Since Darwin first observed the establishment of exotic or alien species in areas outside their native distribution ranges, leading him to postulate what is now known as Darwin's naturalization conundrum (Darwin, 1859), understanding the drivers of alien species establishment, and in some cases their biological invasion, has become a major area of investigation (Fristoe et al., 2021;Mack et al., 2000;Richardson et al., 2000;Vilà et al., 2011). Darwin's naturalization conundrum proposes two alternative hypotheses that may favour the establishment of non-native species (Figure 1). The first is the preadaptation hypothesis (Figure 1), where alien species that are