There is room for everyone: Invasion credit cannot be inferred from the species-area relationship in fragmented forests

Questions: Land use change, habitat fragmentation and biological invasion represent major drivers of global change that strongly interact to alter ecosystems. Following the breaking apart of forests into smaller fragments or the afforestation of former agricultural lands, biodiversity experiences drastic changes due to species loss and turnover over time. This leads to two important outcomes, namely extinction debt and invasion (colonization) credit, which both reflect the inertia of the system's response to environmental changes. Our study investigated the following questions: center dot Is it possible to infer invasion credit from species-area relationship (SAR) residuals both for native and alien plants? center dot Is there any trend linked with the degree of habitat fragmentation through time? Location: Somme, Oise and Aisne departments, northern France. Methods: We analyzed the pattern of SARs' residuals for native and alien vascular plant species separately across nine sets of forest fragments that differ by the landscape matrix they are embedded in (i.e., open field, bocage, forest), while considering plant richness, area and age of the 355 forest patches. Results: The relationship between alien and native SARs' residuals is positive across all landscapes, suggesting a lack of invasion credit. Instead, these results support the "rich get richer" hypothesis, that is a high environmental heterogeneity allows colonization by new species, be they native or alien. Interestingly, the relationship between alien and native residuals depends upon fragment age (i.e., time since patch creation) in the most intensively managed landscapes (i.e., open fields). In the latter, recent forest patches are more prone to alien invasion, as a likely consequence of increased alien propagule pressure (i.e., more sources and vectors for alien plants), increased forest invasibility (i.e., disturbance-induced environmental heterogeneity), and decreased matrix permeability (i.e., natives are more dispersal-limited than aliens). Conclusions: Our study provides new insights into alien species ecology, by showing that (i) it is not possible to infer "invasion credit" from the SAR's residuals; (ii) the invasion rate by alien species in forest fragments increases with their native species richness, and (iii) this relationship depends upon patch age in intensively managed landscapes.