Which factors limit the etablishement of brachypodium retusum : a key species in ecological restoration of Mediterranean steppes ?

Ramose false brome (Brachypodium retusum) is a perennial herbaceous species that dominates dry grasslands of the Western Mediterranean. In our southern French study area, spontaneous re-colonisation is very low after soil disturbance. This observation does not correspond to the results of studies from other regions showing a high colonisation capacity. The major objective of this PhD thesis was to test different hypotheses potentially explaining the different colonisation patterns.In the first chapter, we tested whether these different colonisation patterns are the result of genetic differentiation among populations. AFLP markers were used to analyse genetic structure including neutral population differentiation in 17 Western Mediterranean populations. In a sub-sample of 13 French populations, differentiation in phenotypic traits under selection was tested in a common garden and compared to neutral differentiation. In the second chapter, we present a study on adaptive differentiation in phenotypic traits testing a potentially differential response to the manipulation of key environmental factors. The last two chapters of the PhD analysed environmental factors that limit re-colonisation in the field independent of genotype or population. In the third chapter, we tested the effect of grazing and fire on vegetative recovery as well as on sexual reproduction of established B. retusum and in the fourth chapter we measured the effect of initial watering and grazing on the establishment of transplanted seedlings pre-grown in a greenhouse and of field-sown seedlings.Our results showed that populations of B. retusum are genetically differentiated in neutral markers but also in phenotypic traits. This differentiation is superior to drift alone and suggests adaptation to environmental conditions, particularly to summer temperature and winter frost frequency. A differential response to experimental manipulation of environmental factors (soil, pasture, soil moisture) confirmed the adaptive character of genetic differentiation. The field experiments showed that fire has a positive effect on B. retusum reproduction and on the associated community whereas vegetative recovery was not higher than community average. Two seasons of grazing exclosure did not affect any of the measured parameters in adult populations. Initial watering affected seedling establishment in the first season. In the second season, the watering main effect was not significant but interestingly the effect remained positive on survival in grazed plots whereas no such effect was observed in exclosures. Grazing in early life cycle stages of B. retusum had a negative effect on seedling recruitment and growth.In conclusion, adaptive differentiation between populations may have contributed to regional differences in colonisation capacity and needs to be taken into account in targeting source populations for plant introduction in ecological restoration. The positive response of B. retusum indicated that fire was an important selective force in the past which may be used to favour the species and its associated plant community in current conservation and restoration management. Short-term grazing exclosure is tolerated by mature B. retusum populations but long-term abandonment results in a decrease of cover relative to high-growing perennial grasses. In early stages of seedling establishment grazing should be avoided to guarantee introduction success - or grazing stress needs to be compensated by watering.