Abstract
Range-expanding species, or neonatives, are native plants that spread beyond their original range due to recent climate or human-induced environmental changes. Retama monosperma was initially planted near the Guadalquivir estuary for dune stabilisation. However, changes in the sedimentary regime and animal-mediated dispersal have facilitated its exponential expansion, threatening endemic species and critical dune habitats. The main objective of this study was to identify the key functional traits that may explain the competitive advantage and rapid spread of R. monosperma in coastal dune ecosystems. We compared its seasonal responses with those of three co-occurring woody species, two native (Juniperus phoenicea and J. macrocarpa) and one naturalised (Pinus pinea), at two sites differing in groundwater availability within a coastal dune area (Doñana National Park, Spain). We measured water relations, leaf traits, stomatal conductance, photochemical efficiency, stable isotopes, and shoot elongation in 12 individuals per species. Repeated-measures ANOVA showed significant effects of species and species × season interaction for relative water content, shoot elongation, effective photochemical efficiency, and stable isotopes. R. monosperma showed significantly higher shoot elongation, relative water content, and photochemical efficiency in summer compared with the other species. Stable isotope data confirmed its nitrogen-fixing capacity. This characteristic, along with the higher seasonal plasticity, contributes to its competitive advantage. Given the ecological fragility of coastal dunes, understanding the functional traits favouring the success of neonatives such as R. monosperma is essential for biodiversity conservation and ecosystem management.