Contexte & Objectifs. Le feu est une perturbation récurrente dans les écosystèmes méditerranéens et est souvent considéré comme une composante intrinsèque pour le fonctionnement de ces écosystèmes. Cependant, le rôle des incendies de forêt sur la biodiversité reste controversé. Ainsi, bien que la perception de cette perturbation par la communauté scientifique prenne en compte les effets négatifs et positifs, le feu reste pour une grande part de l’opinion publique une catastrophe humaine et environnementale. Mes recherches ont eu pour objectif de quantifier les effets du feu sur la biodiversité végétale dans les écosystèmes à climat méditerranéen en abordant la relation feu-biodiversité à différentes échelles et en particulier en considérant les caractéristiques spatiales et temporelles des mosaïques générées par les feux au niveau du paysage. L’étude a été dupliquée dans deux biomes méditerranéens distincts (France méditerranéenne et Sud-ouest Australien) de sorte à rechercher à travers les différences les patrons de réponse communs.Méthodologie. Généralement, les feux ont été étudiés comme des évènements singuliers dont il convenait de mesurer l’impact sur les écosystèmes. Dans cette étude, les feux ont été considérés comme une succession d’évènements formant au niveau du paysage des mosaïques en s’additionnant dans le temps et en se superposant dans l’espace. La diversité de ces mosaïques a été quantifiée et a permis de définir un nouveau paramètre, la pyrodiversité. Mes recherches proposent une méthodologie objective et adaptée à la caractérisation de la pyrodiversité à l’échelle du paysage. Les patrons de feux, compilés sur 50 ans, ont été analysés et la diversité de la végétation (habitats, espèces et traits fonctionnels) a été échantillonnée en suivant le protocole développé dans le cadre du projet Européen EBONE. En utilisant la même échelle géographique pour croiser patrons de feu et patrons de végétation, il a été possible de relier pyrodiversité et biodiversité et d’examiner cette relation simultanément au niveau de la communauté végétale (habitat) et du paysage (mosaïque d’habitats).Résultats. Cette étude démontre qu’une pyrodiversité élevée (grande variabilité des patrons spatio-temporels des feux à l’échelle du paysage) est associée à des stades de succession de végétation plus variés et de ce fait maximise la diversité des habitats au niveau du paysage. Les résultats acquis permettent de montrer que le feu exerce une influence significative, avec d’autres facteurs comme les conditions environnementales ou l’utilisation du sol, sur la composition des habitats ainsi que sur l’hétérogénéité spatiale de la mosaïque d’habitats au sein du paysage. Que l’on considère les effets du feu sur les espèces végétales ou les traits fonctionnels de ces taxons, mes travaux montrent également que la diversité des plantes vasculaires au niveau de l’habitat varie avec la fréquence des feux suivant l’hypothèse de perturbation intermédiaire. À l’échelle du paysage, la relation pyrodiversité-biodiversité vérifie l’hypothèse de perturbation hétérogène : les maxima de diversité alpha et beta ont été observés lorsque les mosaïques de feux étaient caractérisées par un niveau élevé de pyrodiversité. À l’échelle régionale, la pyrodiversité favorise la diversité des habitats (diversité gamma) jusqu’à un certain seuil. Les deux aires d’étude affichent des profils de pyrodiversité contrastés, du fait de leurs gestions opposées du feu, mais égalemet des flores très contrastées. Il en résulte des différences de résilience et résistance au feu au sein de leurs écosystèmes respectifs. Toutefois, les deux biomes méditerranéens et leur végétation présentent des convergences évidentes dans leurs réponses écologiques au gradient de pyrodiversité.ConclusionsCette étude permet de resituer le rôle du feu et sa contribution à la richesse floristique des écosystèmes méditerranéens. / Aim and Background. Fire is an integral component of Mediterranean type ecosystems and other fire-prone systems worldwide but the perception of fire as an ecological disaster remains a widely-held view among human societies. Understanding whether and to what extent fire is beneficial or detrimental to biodiversity is a current research priority for conservation management in fire-prone systems. My research has attempted to quantify the multi-scale effects of fire on various facets of vegetation diversity, including landscape diversity, taxonomic diversity and functional diversity from a landscape ecology perspective. We used two contrasted Mediterranean study cases i.e., south-eastern France and south-western Australia in order to distinguish from their differences common ecological patterns.Methods. Fires and their effects on vegetation are normally studied as single and localized events. Here, we considered landscape-scale fire mosaics resulting from the compilation of fire events that occurred among time and space in a given landscape. The diversity of those fire mosaics has been quantified and permitted to define a new parameter i.e., pyrodiversity. This research proposes a conceptual and practical methodology for the objective characterization of pyrodiversity based on the comparison of two Mediterranean environments: south-west of Western Australia and southern France. Fire patterns were analyzed retrospectively over a 50-year period while vegetation, habitats and species diversity were quantified at the landscape scale using a monitoring protocol developed within the EBONE EU-project. Using the same geographical scale for fire and vegetation patterns permitted to cross the information on biodiversity and pyrodiversity. The relationship between pyrodiversity and biodiversity was examined at the community level (i.e., habitat) and landscape level (mosaic of habitats) with considering successively habitat diversity, taxonomic diversity and functional diversity.ResultsThis study provides evidence that high pyrodiversity (i.e., spatio-temporal diversity of fire patterns) is associated to various successional trajectories of vegetation and thereby maximizes the diversity of habitats at landscape scale. The results indicate that fire significantly contributes with environmental factors and land management to determine the composition of habitats across a landscape and spatial heterogeneity of landscape mosaics. Whether considering the effects of a single fire event on plant species or on plant functional traits, I found that plant diversity varies with fire frequency according to the intermediate disturbance hypothesis at the habitat level. At the landscape level, the pyrodiversity-biodiversity relationship verified the heterogeneous disturbance hypothesis i.e., maxima of alpha and beta diversity are associated to fire mosaics of high spatiotemporal heterogeneity. At the regional scale, pyrodiversity enhances habitat diversity (gamma diversity) within the landscape until a certain threshold. The two studies areas display contrasting profiles of pyrodiversity due to their different fire management strategies but also exhibit taxonomically unrelated floras. Hence, the two Mediterranean biomes were characterized by different ecosystem resilience and resistance to fire. Despite these differences, the two Mediterranean biomes show convergence trends in their ecological response to the gradient of pyrodiversity.Conclusion. This study tackled the Pyrodiversity-Biodiversity paradigm and demonstrated that Mediterranean-type ecosystems need pyrodiversity to maintain the variety of habitats and species at both habitat and landscape levels. Fire effects on vegetation cannot be interpreted directly from a local perspective with considering a single fire event.

I. An original methodological discussion is proposed on the problem of the typology of tropical rain forest’s plant communities, based on the study of forest types across gradients of continentality and elevation, within Atlantic central Africa. These investigations were based on the statement that the main problems in forest typology are related to the non-zonal or zonal character of the different vegetation types and to non considering the relations and differences between forest strata. II. Field data consisted in phytosociological homogeneous sample plots localized within different recognized phytogeographical entities, in a region of tropical Africa where these entities are known to be well conserved. A total of 37 such plots were inventoried in the region extending from the littoral forests of Ndoté, Equatorial Guinea, which are wet evergreen forests, to the continental forests of the Dja, Cameroon, known as evergreen seasonal forests. The studied region also included the oriental Atlantic forests of Equatorial Guinea, known as moist evergreen forests or caesalp forests. In various parts of this continentality gradient, some plots were localized within climax non-zonal formations, namely the submontane rain forests. The emphasis was put on the vegetation of the Monte Alén National Park. The sampling methodology was willing to be as "complete ", including all strata, "quantitative ", enumerating all individuals, and "representative ", within each stratum, as possible. These multi-layers plots were realised using nested sub-plots, with a sampling size of 100 individuals for every ligneous stratum recognized (dominant trees, dominated trees and shrubs) and a sampling size of 200m² for the herbaceous and suffrutex stratum. Forest types were defined independently for each stratum and the differences were analysed. A method was proposed for the simultaneous analysis of all floristic data, converting and standardizing the values from ligneous strata, on the one hand, and from understorey strata, on the other hand. III. Ten forest types were described using IndVal and discussed in the general context of the guineo-congolian region, from a syntaxonomic view point (agglomerative classification) and from a phytogeographical view point (divisive classification). Homologies between these two approaches are described. The proposed phytogeographical system is based on an "open " conception of hierarchical classifications, combining advantages of agglomerative and divisive classifications. In concrete terms, the non-zonal criteria, for example the submontane variants, are categorised separately and in analogy with the zonal criteria, related to the usual phytochoria. Analysis of ecological relationships for the 10 communities showed that the main variables related to the floristic variability in our mainland rain forests are elevation, rainfall, hygrometry (estimated using bryophytes cover levels) and distance to the ocean. The two extremes on the vertical microclimatic gradient, dominant trees stratum and herbaceous stratum, give similar typologies, however canonical analysis showed that for the herbaceous layer, non-zonal variables (hygrometry and elevation) were gaining more importance when the influence of the two zonal variables was attenuated. In every case, spatial autocorrelation was less important than the environment in explaining floristic variability but its role increased in the spatial arrangement of understorey species, whose dispersal capacity is generally lower than canopy trees. The phytosociological, phytogeographical and ecological description of forest types is accompanied by a physiognomical description using biological types spectrum, as well as architectural models, leaf sizes, etc. With regard to diversity, we have demonstrated that species richness was higher from upper to lower strata because of the accumulation in lower strata of species from various strata. On the other hand, the proper stratum diversity, i.e. the structural set, decreased from dominant trees to shrubs. The proper diversity of the herb layer showed relatively high figures mainly due to the higher individual density in relation to the existence of microstrata. Within the 37 sample plots, 1,050 taxa have been identified to species or morpho-species levels, for a total of 25,750 individuals. These taxa represent 442 genus among 104 families. The richest forest type is found on the foothills of the Niefang range, on the windward side. This forest type is also characterised by a high number of oligotypic genus and by species belonging to functional types indicators of glacial refuges. These functional types are defined on the basis of the dispersal capacity and on kind of stand needed for effective germination. We formulated the hypothesis that this kind of "foothills refuge ", characterised by his zonal nature, could have been one of the rare refuges for species from mainland rain forests, while montane and fluvial refuges would mainly have preserved species from non-zonal forest types: (sub)montane and riverine. Based on indicator species of submontane forests, a potential distribution map of this forest type has been realised at the Atlantic central African scale. More than 400 submontane forest localities have been mapped. These forests begin at 400m of altitude near the ocean, and progressively at higher altitude for increasing distance to the ocean. Many lowland localities also comprised submontane species, which could indicate the existence of ecological transgressions. These transgressions would allow migratory tracks for submontane species between isolated mountain ranges, not only during glacial periods, through heights at the northern and southern borders of the congo basin, but also contemporarily through the lowland riverine forest network, in the centre of this basin. Finally, a special attention has been attributed to littoral forests and to some cases of choroecological transgressions, coupled to the ecological equalization phenomenon.

I. An original methodological discussion is proposed on the problem of the typology of tropical rain forest’s plant communities, based on the study of forest types across gradients of continentality and elevation, within Atlantic central Africa. These investigations were based on the statement that the main problems in forest typology are related to the non-zonal or zonal character of the different vegetation types and to non considering the relations and differences between forest strata.<p><p>II. Field data consisted in phytosociological homogeneous sample plots localized within different recognized phytogeographical entities, in a region of tropical Africa where these entities are known to be well conserved. A total of 37 such plots were inventoried in the region extending from the littoral forests of Ndoté, Equatorial Guinea, which are wet evergreen forests, to the continental forests of the Dja, Cameroon, known as evergreen seasonal forests. The studied region also included the oriental Atlantic forests of Equatorial Guinea, known as moist evergreen forests or caesalp forests. In various parts of this continentality gradient, some plots were localized within climax non-zonal formations, namely the submontane rain forests. The emphasis was put on the vegetation of the Monte Alén National Park.<p><p>The sampling methodology was willing to be as "complete ", including all strata, "quantitative ", enumerating all individuals, and "representative ", within each stratum, as possible. These multi-layers plots were realised using nested sub-plots, with a sampling size of 100 individuals for every ligneous stratum recognized (dominant trees, dominated trees and shrubs) and a sampling size of 200m² for the herbaceous and suffrutex stratum.<p><p>Forest types were defined independently for each stratum and the differences were analysed. A method was proposed for the simultaneous analysis of all floristic data, converting and standardizing the values from ligneous strata, on the one hand, and from understorey strata, on the other hand.<p><p>III. Ten forest types were described using IndVal and discussed in the general context of the guineo-congolian region, from a syntaxonomic view point (agglomerative classification) and from a phytogeographical view point (divisive classification). Homologies between these two approaches are described. The proposed phytogeographical system is based on an "open " conception of hierarchical classifications, combining advantages of agglomerative and divisive classifications. In concrete terms, the non-zonal criteria, for example the submontane variants, are categorised separately and in analogy with the zonal criteria, related to the usual phytochoria.<p><p>Analysis of ecological relationships for the 10 communities showed that the main variables related to the floristic variability in our mainland rain forests are elevation, rainfall, hygrometry (estimated using bryophytes cover levels) and distance to the ocean. The two extremes on the vertical microclimatic gradient, dominant trees stratum and herbaceous stratum, give similar typologies, however canonical analysis showed that for the herbaceous layer, non-zonal variables (hygrometry and elevation) were gaining more importance when the influence of the two zonal variables was attenuated. In every case, spatial autocorrelation was less important than the environment in explaining floristic variability but its role increased in the spatial arrangement of understorey species, whose dispersal capacity is generally lower than canopy trees. The phytosociological, phytogeographical and ecological description of forest types is accompanied by a physiognomical description using biological types spectrum, as well as architectural models, leaf sizes, etc.<p><p>With regard to diversity, we have demonstrated that species richness was higher from upper to lower strata because of the accumulation in lower strata of species from various strata. On the other hand, the proper stratum diversity, i.e. the structural set, decreased from dominant trees to shrubs. The proper diversity of the herb layer showed relatively high figures mainly due to the higher individual density in relation to the existence of microstrata. Within the 37 sample plots, 1,050 taxa have been identified to species or morpho-species levels, for a total of 25,750 individuals. These taxa represent 442 genus among 104 families. The richest forest type is found on the foothills of the Niefang range, on the windward side. This forest type is also characterised by a high number of oligotypic genus and by species belonging to functional types indicators of glacial refuges. These functional types are defined on the basis of the dispersal capacity and on kind of stand needed for effective germination. We formulated the hypothesis that this kind of "foothills refuge ", characterised by his zonal nature, could have been one of the rare refuges for species from mainland rain forests, while montane and fluvial refuges would mainly have preserved species from non-zonal forest types: (sub)montane and riverine.<p><p>Based on indicator species of submontane forests, a potential distribution map of this forest type has been realised at the Atlantic central African scale. More than 400 submontane forest localities have been mapped. These forests begin at 400m of altitude near the ocean, and progressively at higher altitude for increasing distance to the ocean. Many lowland localities also comprised submontane species, which could indicate the existence of ecological transgressions. These transgressions would allow migratory tracks for submontane species between isolated mountain ranges, not only during glacial periods, through heights at the northern and southern borders of the congo basin, but also contemporarily through the lowland riverine forest network, in the centre of this basin. Finally, a special attention has been attributed to littoral forests and to some cases of choroecological transgressions, coupled to the ecological equalization phenomenon.<p> / Doctorat en sciences agronomiques et ingénierie biologique / info:eu-repo/semantics/nonPublished