Aims: The stress‐gradient hypothesis (SGH) predicts how plant interactions change along environmental stress gradients. We tested the SGH in an aridity gradient, where support for the hypothesis and the specific shape of its response curve is controversial. Location: Almería, Cáceres and Coimbra, three sites in the Iberian Peninsula that encompass the most arid and wet habitats in the distribution range of a nurse shrub species –Retama sphaerocarpa L. (Boiss) – in Europe. Methods: We analysed the effect of Retama on its understorey plant community and measured the biomass and species richness beneath Retama and in gaps. We estimated the frequency (changes in species richness), importance and intensity of the Retama effects, and derived the severity–interaction relationship pattern, analysing how these interaction indices changed along this aridity gradient. Results and conclusions: The intensity and frequency of facilitation by Retama increased monotonically with increasing environmental severity, and the importance tended to have a similar pattern, overall supporting the SGH. Our data did not support predictions from recent revisions of the SGH, which may not apply to whole plant communities like those studied here or when interactions are highly asymmetrical. Facilitation by Retama influenced community composition and species richness to the point that a significant fraction of species found at the most arid end of the gradient were only able to survive beneath the nurse shrub, whereas some of these species were able to thrive in gaps at more mesic sites, highlighting the ecological relevance of facilitation by nurse species in mediterranean environments, especially in the driest sites.
Aim This study analysed the diversity and identity of the rhizobial symbionts of co‐existing exotic and native legumes in a coastal dune ecosystem invaded by Acacia longifolia. Location An invaded coastal dune ecosystem in Portugal and reference bradyrhizobial strains from the Iberian Peninsula and other locations. Methods Symbiotic nitrogen‐fixing bacteria were isolated from root nodules of plants of the Australian invasive Acacia longifolia and the European natives Cytisus grandiflorus, Cytisus scoparius and Ulex europaeus. Total DNA of each isolate was amplified by polymerase chain reaction (PCR) with the primer BOX A1R. Subsequent PCR‐sequencing and phylogenetic analyses of the internal transcribed spacer region and the nifD and nodA genes were performed for all different strains. Results The four plant species analysed were nodulated by bacteria from three different Bradyrhizobium lineages, although most of the isolates belonged to the Bradyrhizobium japonicum lineage sensu lato. Ninety‐five per cent of the bradyrhizobia isolated from A. longifolia, C. grandiflorus and U. europaeus in the invaded ecosystem had nifD and nodA genes of Australian origin. Seven isolates obtained in this study define a new distinctive nifD group of Bradyrhizobium from western and Mediterranean Europe. Main conclusions These results reveal the introduction of exotic bacteria with the invasive plant species, their persistence in the new geographical area and the nodulation of native legumes by rhizobia containing exotic symbiotic genes. The disruption of native mutualisms and the mutual facilitation of the invasive spread of the introduced plant and bradyrhizobia could constitute the first report of an invasional meltdown documented for a plant–bacteria mutualism.
The ability to form symbiotic associations with soil microorganisms and the consequences for plant growth were studied for three woody legumes grown in five different soils of a Portuguese coastal dune system. Seedlings of the invasive Acacia longifolia and the natives Ulex europaeus and Cytisus grandiflorus were planted in the five soil types in which at least one of these species appear in the studied coastal dune system. We found significant differences between the three woody legumes in the number of nodules produced, final plant biomass and shoot 15 N content. The number of nodules produced by A. longifolia was more than five times higher than the number of nodules produced by the native legumes. The obtained 15 N values suggest that both A. longifolia and U. europaeus incorporated more biologically-fixed nitrogen than C. grandiflorus which is also the species with the smallest distribution. Finally, differences were also found between the three species in the allocation of biomass in the different studied soils. Acacia longifolia displayed a lower phenotypic plasticity than the two native legumes which resulted in a greater allocation to aboveground biomass in the soils with lower nutrient content. We conclude that the invasive success of A. longifolia in the studied coastal sand dune system is correlated to its capacity to nodulate profusely and to use the biologically-fixed nitrogen to enhance aboveground growth in soils with low N content.
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