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Invasive alien plants are species introduced and naturalized outside of their native distribution range and which have the capacity to maintain and expand their population. Some of these species are considered to be ecosystem transformers by altering their structure, functioning as well as resident animal and plant communities. These induced alterations make some of these species undesirable through their ecological and economical impacts. The work presented in this thesis aimed at a better understanding of the impact of biological invasions by alien plants. The soil fauna, native vegetation and their substrate, as well as ecosystem functioning, were studied at different spatial scales. Two exotic alien species, invasive in Europe, were considered as biological models for this work: the black locust (Robinia pseudoacacia) and the Japanese knotweed (Reynoutria japonica). Firstly, a global meta-analysis demonstrated the positive impact that plant invasions can exert on the abundance of some groups within the soil fauna, notably primary consumers, within different types of habitats (open or closed). Then, a large-scale study on the black locust revealed the differences that can can occur in the response of forest ecosystems to invasions along a latitudinal gradient. Study sites along this gradient, distributed amog four distinct regions in western Europe, exhibit differences in climate and dominant native vegetation which can alter the impact of the black locust. A detailed study on black locust impact in Normandy demonstrated the impact of R. pseudoacacia on native plant and soil fauna communities, as well as some ecosystem functions, in comparison to two native tree species. Finally, a laboraty experiment demonstrated the impact that allelopathic compounds extracted from Japanese knotweed rhizomes can have on some organisms within the soil fauna. This study showed that some invasive alient plants can influence the soil fauna, and soil food webs, through their secondary metabolism. This thesis illustrates that simultaneous study of both aboveground and belowground ecosystem compartments at different spatial scales is of interest in the context of biological invasions., Les espèces exotiques envahissantes végétales sont des plantes introduites et naturalisées hors de leur aire de répartition native et capables de maintenir et d’accroitre leur population. Certaines sont considérées comme transformatrices de par leur effet sur les écosystèmes : leur structure, leur fonctionnement ainsi que leur communauté végétale et animale. Ces transformations peuvent rendre certaines de ces espèces nuisibles de par leurs impacts écologiques et économiques important. Les travaux réalisés dans le cadre de cette thèse et présentés ici ont pour objectif d’approfondir les connaissances sur l’impact des invasions biologiques. La faune du sol, la végétation native et leur substrat ainsi que son fonctionnement ont été étudiés à différentes échelles spatiales. Deux espèces exotiques, envahissantes en Europe, ont été considérées comme modèles pour ces travaux : le robinier faux-acacia (Robinia pseudoacacia) et la renouée du japon (Reynoutria japonica). Premièrement, une méta-analyse globale a permis de démontrer l’effet positif des invasions biologiques végétales sur l’abondance de certains groupes de la faune du sol, notamment les consommateurs primaires, en fonction de la structure de l’habitat (ouvert ou fermé). Ensuite, une étude à large échelle sur le robinier faux-acacia a permis d’illustrer les différences qui peuvent exister dans la réponse des écosystèmes forestiers aux invasions le long d’un gradient latitudinal. Ce gradient, composé de quatre régions distinctes en Europe de l’Ouest présente des différences de climat et de végétation dominante, ces différences modifiant l’impact du robinier faux-acacia. Une étude approfondie sur le robinier faux-acacia en Normandie a permis de mieux comprendre l’effet du robinier faux-acacia sur les communautés animales et végétales ainsi que sur le fonctionnement des écosystèmes par comparaison avec deux essences natives dominantes. Finalement, une manipulation expérimentale en laboratoire a démontré l’impact des composés allélopathiques de la renouée du Japon sur une partie de la faune du sol. Cette étude a montré que certaines espèces exotiques envahissantes sont susceptibles d’influencer la faune, et les réseaux trophiques, du sol par leur métabolisme secondaire. Ces travaux illustrent l’intérêt, dans le contexte des invasions biologiques végétales, de l’étude simultanée des compartiments aériens et souterrains à différentes échelles spatiales.

The relevance of biodiversity to human health is an increasing international political issue as it causes concern for ethical and aesthetic reasons, but also has a strong impact on ecosystem properties and ecological goods and services utilised by humanity. However, humans have applied increasing pressure on worldwide biodiversity through pollution, land-use and climate change, over exploitation and the introduction of invasive species which can lead to major alterations of biological communities and a consequent decrease in biodiversity. When communities are assembled at random from a pool of species, more diverse mixtures have a higher probability to contain species or species-groups with high capacity to drive ecosystem processes. Therefore, it is relevant to ask "how may we classify species in a community in terms of functionality?", and "which species-group is important to maintain the productivity and stability of ecosystems?" In this thesis, I chose to classify species according their frequency and their cumulative relative cover in a plant community (i.e. plant hierarchy) and to differentiate three species-groups: dominant, subordinate and transient, which contribute towards biodiversity. Dominant species are clearly very important for ecosystem functioning, due to the large amount of biomass they produce, but there is growing evidence that subordinate species, which represent a low amount of plant biomass in grassland ecosystems, are also of considerable importance for ecosystem functioning. Given this recognition, a further objective was to explore the ecosystem level effects of these subordinate plant species, which is currently not well known. This research focused on two aspects: the persistence of subordinate species in the community, and their role in grassland ecosystems. The approach to satisfy these objectives encompassed four years of experimental field studies and glasshouse microcosm experiments. In order to explain the persistence of subordinate species in semi-natural grasslands, we performed two greenhouse experiments to test the effects of root competition exclusion and the inoculation of arbuscular mycorrhizal fungi (AMF) on the competitiveness of dominant and subordinate species, directly measured by their biomass production. The effects of subordinate species on ecosystem functioning were assessed through long-term field experiment in two sites in the Swiss Jura Mountains, where dominant, subordinate and transient species were preliminarily determined. At both site (Les Amburnex, La Frétaz), a removal experiment was carried out in randomly replicated plots with three different treatments: control without perturbations, removal of all subordinate species and partial removal of dominant biomass. In one site, a summer drought treatment was added using rainout shelters to simulate an extreme climatic event. Biomass production, litter decomposition and soil respiration were monitored during four years in each plot and complemented in some cases by measurements of soil dissolved inorganic nitrogen concentrations, soil microbial communities (bacteria and AMF) and carbon and nitrogen isotopes in plant leaves. The results of the different experiments conducted in this thesis highlight the importance of cattle activity and AMF on the persistence of subordinate plant species in semi-natural grasslands. Indeed, cattle activities (i.e. trampling) provide spatial heterogeneity, through gap creation (areas of root competition reduction), and thus favours the growth of less competitive subordinate species. While AMF acted as a parasite in the greenhouse experiment, the current findings suggest that the competitiveness of dominant species was reduced through the action of the fungus, which indirectly enhanced subordinate species. The interactions between subordinate species and soil organisms were confirmed in the field as bacterial and AMF communities shifted in plots where subordinate species were removed compared to the control treatments. Moreover, the absence of subordinate species had many negative effects on ecosystem functioning by reducing litter decomposition, soil respiration, nitrogen mineralization and community above-ground production. These findings suggest that plant-soil feedbacks explain the importance of subordinate species on ecosystem functioning despite only representing a very low quantity of biomass in plant communities. During the summer drought, subordinate species increased the resistance of the plant community and maintained productivity. Therefore, these outcomes confirm the role of drought-resistant subordinates in the functioning of grassland ecosystems threatened by climate change. The synthesis of this thesis has important implications for the way grasslands should be managed for biodiversity and confirms that extensive grazing must be maintained as it promotes species coexistence and the persistence of key subordinate species. Moreover, this work demonstrates the important role of subordinate species in ecosystem functioning, showing that not only dominant species determine ecosystem properties. This thesis serves as a contribution to the advances in our understanding of ecosystem functioning and above and below-ground linkages, and provides basis for future research in this domain.