Why some species become invasive while others do not remains an elusive question. It has been proposed that invasive species should depend less on mutualisms, because their spread would then be less constrained by the availability of mutualistic partners. We tested this idea with the genus Pinus, whose degree of invasiveness is known at the species level (being highly and negatively correlated with seed size), and which forms obligate mutualistic associations with ectomycorrhizal fungi (EMF). Mycorrhizal dependence is defined as the degree to which a plant needs the mycorrhizal fungi to show the maximum growth. In this regard, we use plant growth response to mycorrhizal fungi as a proxy for mycorrhizal dependence. We assessed the responsiveness of Pinus species to EMF using 1,206 contrasts published on 34 species, and matched these data with data on Pinus species invasiveness. Surprisingly, we found that species that are more invasive depend more on mutualisms (EMF). Seedling growth of species with smaller seeds benefited more from mutualisms, indicating a higher dependence. A higher reliance on EMF could be part of a strategy in which small‐seeded species produce more seeds that can disperse further, and these species are likely to establish only if facilitated by mycorrhizal fungi. On the contrary, big‐seeded species showed a lower dependence on EMF, which may be explained by their tolerance to stressful conditions during establishment. However, the limited dispersal of larger seeds may limit the spread of these species. We present strong evidence against a venerable belief in ecology that species that rely more on mutualisms are less prone to invade, and suggest that in certain circumstances greater reliance on mutualists can increase spread capacity.

Summary: The success of invasive plants is influenced by many interacting factors, but evaluating multiple possible mechanisms of invasion success and elucidating the relative importance of abiotic and biotic drivers is challenging, and therefore rarely achieved. We used live, sterile or inoculated soil from different soil origins (native range and introduced range plantation; and invaded plots spanning three different countries) in a fully factorial design to simultaneously examine the influence of soil origin and soil abiotic and biotic factors on the growth of invasive Pinus contorta. Our results displayed significant context dependency in that certain soil abiotic conditions in the introduced ranges (soil nitrogen, phosphorus or carbon content) influenced responses to inoculation treatments. Our findings do not support the enemy release hypothesis or the enhanced mutualism hypothesis, as biota from native and plantation ranges promoted growth similarly. Instead, our results support the missed mutualism hypothesis, as biota from invasive ranges were the least beneficial for seedling growth. Our study provides a novel perspective on how variation in soil abiotic factors can influence plant–soil feedbacks for an invasive tree across broad biogeographical contexts.

The naturalization of an introduced species is a key stage during the invasion process. Therefore, identifying the traits that favor the naturalization of non‐native species can help understand why some species are more successful when introduced to new regions. The ability and the requirement of a plant species to form a mutualism with mycorrhizal fungi, together with the types of associations formed may play a central role in the naturalization success of different plant species. To test the relationship between plant naturalization success and their mycorrhizal associations we analysed a database composed of mycorrhizal status and type for 1981 species, covering 155 families and 822 genera of plants from Europe and Asia, and matched it with the most comprehensive database of naturalized alien species across the world (GloNAF). In mainland regions, we found that the number of naturalized regions was highest for facultative mycorrhizal, followed by obligate mycorrhizal and lowest for non‐mycorrhizal plants, suggesting that the ability of forming mycorrhizas is an advantage for introduced plants. We considered the following mycorrhizal types: arbuscular, ectomycorrhizal, ericoid and orchid mycorrhizal plants. Further, dual mycorrhizal species were those that included observations of arbuscular mycorrhizas as well as observations of ectomycorrhizas. Naturalization success (based on the number of naturalized regions) was highest for arbuscular mycorrhizal and dual mycorrhizal plants, which may be related to the low host specificity of arbuscular mycorrhizal fungi and the consequent high availability of arbuscular mycorrhizal fungal partners. However, these patterns of naturalization success were erased in islands, suggesting that the ability to form mycorrhizas may not be an advantage for establishing self‐sustaining populations in isolated regions. Taken together our results show that mycorrhizal status and type play a central role in the naturalization process of introduced plants in many regions, but that their effect is modulated by other factors.