Infoscience

Thesis

Resistance of plant communities to invasive species: disentangling invasiveness from invasibility

Invasive species are nowadays considered as one of the most important threat to biodiversity. By displacing native species, modifying ecosystem functioning and causing substantial losses to agricultural production, they represent a menace to natural and managed ecosystems. Although ecology of invasions has become an important research topic since the last decades, the mechanisms that determine why a given species may invade a given ecosystem and why some biomes are less resistant to invasion are still not clarified. Ecology of invasions is divided into 2 main topics: invasiveness and invasibility. While invasiveness refers to species ability to invade a community, invasibility focuses on the resistance of a community to invasion. Invasiveness may be the result of ecological processes, such as release from biotic constraints or human alteration of the environment (disturbance, stress…) or the consequence of evolutionary processes, such as hybridization or polyploidization that may increase genetic variation and therefore, enhance niche breadth. Invasibility has been said to be influenced by disturbance and biotic factors such as community diversity, dominant species identity, biotic interactions or community compositional stability. The invasion success is the consequence of the interaction between species invasiveness and community invasibility. Most studies in ecological invasions have focused on either invasiveness or invasibility, but hardly both together. By working at the same time and in the same conditions with native and introduced genotypes and by comparing their ecological performances, this thesis aims at a better understanding of both invasiveness and invasibility mechanisms. Two worldwide invasive species, Centaurea maculosa and Senecio inaequidens, were used in several experiments (pot, microcosm, field) to disentangle the importance of invasiveness and community invasibility in their invasion success. Both species encountered polyploidisation in their native range, leading to the presence of diploid and tetraploid populations, whereas only tetraploid populations have been found in the introduced range. Using native diploid, native tetraploid and introduced tetraploid genotypes of the two model-species, allows assessing the effects of genetic variation (diploid vs. tetraploid genotypes) and environmental variation (genotype from native vs. introduced range) on species phenotypic traits variations and consequently on fitness variation and invasiveness. In the community context, studying response of different genotypes to experimental factors and community change gives information on the interaction between invasiveness and invasibility. Plants were grown in pot, in field or in artificially built communities where (i) the management treatment, (ii) the community diversity and (iii) the spatial organisation of resident species were manipulated. In addition, (iv) community species composition, (v) community competitive ability and (vi) compositional and functional stability of the community were monitored. According to the experiments, data were gathered on survival, morphological traits (vegetative height, lateral spread, shoot and root biomass), leaf traits (specific leaf area, leaf dry matter content) and reproductive traits (probability of flowering, capitulum production) of the genotypes (native diploid, native tetraploid and introduced tetraploid) of both model species. Through the use of statistical models and multivariate approaches, the effects of management and biotic factors on survival, growth and reproduction of native and introduced genotypes of these two worldwide invasive species were assessed. Invasion strategy of the two model species was investigated through a growth experiment in optimal conditions in a pot experiment. High investment in seed production could explain invasive success of S. inaequidens through high propagule pressure, whereas C. maculosa's strategy seemed to be oriented towards interactions with belowground communities, as shown by the shift in rhizosphere bacterial communities between genotypes. For both species, polyploidisation in the native range could be linked to a specialisation towards higher competitive ability, which could have allowed the first step of invasion. Introduction in the new range could be related to a loss of specialisation through selection of traits allowing coping with various or changing environments, improving successful spread. Survival of both model species was highly affected by community spatial pattern, management and neighbouring competition. Responses of growth and reproductive output of native and introduced genotypes to management, community spatial pattern and community diversity were species-specific. Growth and reproductive output of both genotypes of S. inaequidens were affected by experimental factors whereas introduced genotypes of C. maculosa were less affected than native ones. Comparison of response of native and introduced genotypes to experimental factors allowed defining two strategies of invasion based on phenotypic plasticity. Centaurea maculosa was able to maintain fitness in all kinds of environments, either favourable or stressful ("Jack-of-all-trades" invader). Senecio inaequidens was able to deal with all kinds of environments and was also able to increase its fitness in favourable conditions ("Jack-and-Master" invader). The combination of the effects of management, community spatial pattern and community diversity on species genotypes allowed defining, for both invasive species, two invasion phases which were impacted by different factors. The introduction phase corresponds to the survival of seedlings and their ability to deal with neighbouring competition. If seedlings manage to survive despite neighbouring competition, they grow and reproduced in order to spread, which corresponds to the establishment phase. In terms of management perspectives, regular mowing or use of highly covering species could limit invasive success of S. inaequidens and C. maculosa respectively. The synthesis of all the experiments conducted in this thesis with C. maculosa and S. inaequidens highlights (1) the importance of polyploidisation in the invasion process as well as (2) the species-specific invasion strategies and consequently (3) the species-specific response of invasive species to abiotic and biotic factors. It also emphasizes on (4) the temporal evolution of the interaction between invasiveness and invasibility since the community factors that affected invasive species fitness changed according to the invasion stage (introduction vs. establishment phase) of the invader. The provided insights into the importance of the interaction between species invasiveness and community invasibility will contribute to improve knowledge in ecology of invasions, in addition to supply some clues for management efficiency.

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