This project predominantly investigated the implications of spatial heterogeneity in the ecological processes of competition and infection. Empirical analysis of spatial heterogeneity was carried out using the lepidopteran species Plodia interpunctella. Using differently viscous food media, it was possible to alter the movement rate of larvae. Soft Foods allow the movement rate of larvae to be high, so that individuals can disperse through the environment and avoid physical encounters with conspecifics. Harder foods lower the movement rate of larvae, restricting the ability of individuals to disperse away from birth sites and avoid conspecifics encounters. Increasing food viscosity and lowering movement rate therefore has the effect of making uniform distributed larval populations more aggregated and patchy. Different spatial structures changed the nature of intraspecific competition, with patchy populations characterised by individuals experiencing lower growth rates and greater mortality because of the reduced food and space available within densely packed aggregations. At the population scale, the increased competition for food individuals experience in aggregations emerges as longer generational cycles and reduced population densities. Aggregating individuals also altered the outcome of interspecific competition between Plodia and Ephestia cautella. In food media that allowed high movement rates, Plodia had a greater survival rate than Ephestia because the larger movement rate of Plodia allowed it to more effectively avoid intraspecific competition. Also the faster growth rate, and so larger size, of Plodia allowed it to dominate interspecific encounters by either predating or interfering with the feeding of Ephestia. In food that restricts movement, the resulting aggregations cause Plodia to experience more intraspecific encounters relative to interspecific, reducing its competitive advantage and levelling the survival of the two species. Spatial structure also affected the dynamics of a Plodia-granulosis virus interaction and the evolution of virus infectivity. Larval aggregation forced transmission to become limited to within host patches, making the overall prevalence of the virus low. However potentially high rates of cannibalism and multiple infections within overcrowded host aggregations caused virus-induced mortality to be high, as indicated by the low host population density when virus is presented. Also aggregated host populations cause the evolution of lower virus infectivity, where less infective virus strains maintain more susceptible hosts within the aggregation and so possess a greater transmission rate. The pattern of variation in resistance of Plodia interpunctella towards its granulosis virus was found using two forms of graphical analysis. There was a bimodal pattern of variation, with most individuals exhibiting either low or high levels of resistance. This pattern was related to a resistance mechanism that is decreasingly costly to host fitness.

Biodiversity is determined by a myriad of complex processes acting at different scales. Given the current rates of biodiversity loss and change, it is of paramount importance that we improve our understanding of the underlying structure of ecological communities. In this thesis, I focused on Species Abundance Distributions (SAD), as a synthetic measure of biodiversity and community structure, and on Beta (β) diversity patterns, as a description of the spatial variation of species composition. I systematically assessed the effect of scale on both these patterns, analysing a broad range of community data, including different taxa and habitats, from the terrestrial, marine and freshwater realms. Knowledge of the scaling properties of abundance and compositional patterns must be fully integrated in biodiversity research if we are to understand biodiversity and the processes underpinning it, from local to global scales. SADs depict the relative abundance of the species present in a community. Although typically described by unimodal logseries or lognormal distributions, empirical SADs can also exhibit multiple modes. However, the existence of multiple modes in SADs has largely been overlooked, assumed to be due to sampling errors or a rare pattern. Thus, we do not know how prevalent multimodality is, nor do we have an understanding of the factors leading to this pattern. Here, I provided the first global empirical assessment of the prevalence of multimodality across a wide range of taxa, habitats and spatial extents. I employed an improved method combining two model selection tools, and (conservatively) estimated that ~15% of the communities were multimodal with strong support. Furthermore, I showed that the pattern is more common for communities at broader spatial scales and with greater taxonomic diversity (i.e. more phylogenetically diverse communities, since taxonomic diversity was measured as number of families). This suggests a link between multimodality and ecological heterogeneity, broadly defined to incorporate the spatial, environmental, taxonomic and functional variability of ecological systems. Empirical understanding of how spatial scale affects SAD shape is still lacking. Here, I established a gradient in spatial scale spanning several orders of magnitude by decomposing the total extent of several datasets into smaller subsets. I performed an exploratory analysis of how SAD shape is affected by area sampled, species richness, total abundance and taxonomic diversity. Clear shifts in SAD shape can provide information about relevant ecological and spatial mechanisms affecting community structure. There was a clear effect of area, species richness and taxonomic diversity in determining SAD shape, while total abundance did not exhibit any directional effect. The results supported the findings of the previous analysis, with a higher prevalence of multimodal SADs for larger areas and for more taxonomically diverse communities, while also suggesting that species spatial aggregation patterns can be linked to SAD shape. On the other hand, there was a systematic departure from the predictions of two important macroecological theories for SAD across scales, specifically regarding logseries distributions being selected only for smaller scales and when species richness and number of families were proportionally much smaller than the total extent. β diversity quantifies the variation in species composition between sites. Although a fundamental component of biodiversity, its spatial scaling properties are still poorly understood. Here, I tested if two conceptual types of β diversity showed systematic variation with scale, while also explicitly accounting for the two β diversity components, turnover and nestedness (species replacement vs species richness differences). I provided the first empirical analysis of β diversity scaling patterns for different taxa, revealing remarkably consistent scaling curves. Total β diversity and turnover exhibit a power law decay with log area, while nestedness is largely insensitive to scale changes. For the distance decay of similarity analysis, while area sampled affected the overall dissimilarity values, rates of similarity were consistent across large variations in sampled area. Finally, in both these analyses, turnover was the main contributor to compositional change. These results suggest that species are spatially aggregated across spatial scales (from local to regional scales), while also illustrating that substantial change in community structure might occur, despite species richness remaining relatively stable. This systematic and comprehensive analysis of SAD and community similarity patterns highlighted spatial scale, ecological heterogeneity and species spatial aggregation patterns as critical components underlying the results found. This work expanded the range of scales at which both theories deriving SAD and community similarity studies have been developed and tested (from local plots to continents). The results here showed strong departures from two important macroecological theories for SAD at different scales. In addition, the overall findings in this thesis clearly indicate that unified theories of biodiversity (or assuming a set of synthetic minimal assumptions) are unable to accommodate the variability in SADs shape across spatial scales reported here, and cannot fully reproduce community similarity patterns across scales. Incorporating more realistic assumptions, or imposing scale dependent assumptions, may prove to be a fruitful avenue for ecological research regarding the scaling properties of SAD and community similarity patterns. This will allow deriving new predictions and improving the ability of theoretical models to incorporate the variability in abundance and similarity patterns across scales.

Les forêts tropicales renferment des peuplements arborescents dont la gestion et l’aménagement nécessitent des connaissances sur leur organisation spatiale et leur dynamique. Les analyses de la structure spatiale des espèces arborescentes peuvent être utilisées en forêts naturelles pour identifier les mécanismes sous-jacents qui structurent les peuplements forestiers afin d’améliorer la compréhension des relations entre les espèces. Cette étude a été menée dans la Réserve Forestière de Yoko (RFY) aux environs de Kisangani à l’est de la RD Congo (R.D.C). Elle consiste à analyser la structure spatiale horizontale des espèces les plus abondantes et à tenter d’identifier, à l’échelle locale, les facteurs et/ou processus écologiques potentiellement explicatifs pour en retirer des enseignements utiles à la gestion des massifs forestiers situés à proximité de Kisangani. <p>Pour ce faire, une parcelle d’échantillonnage de 25 ha (500m 500m) a été délimitée dans le bloc sud de la RFY constitué d’une végétation ligneuse mixte et semi-décidue. Un inventaire forestier a permis d’analyser la composition floristique et structurale de la zone. Trois techniques (ou modèles statistiques) relevant de l’écologie spatiale pour l’analyse de la structure horizontale des espèces ont été utilisées :la méthode du voisin le plus proche de Clark & Evans (1954), la méthode d’échantillonnage aléatoire de Hines & Hines (1979), ces deux méthodes reposant sur une analyse à échelle unique de la parcelle d’étude, et la méthode de Ripley (1977) permettant non seulement une analyse multi-échelle mais aussi l’étude des relations intra et interspécifiques. Concernant ce point précis, les arbres ont été catégorisés en trois stades de développement sur la base de leurs diamètres (les jeunes, les immatures et les adultes). Une analyse comparative et théorique des trois méthodes a été effectuée. <p>Un total de 169 espèces appartenant à 36 familles dont 114 genres ont été identifiées, et la famille des Fabaceae dont la majorité des espèces appartiennent à la sous famille des Caesalpinioideae est apparue prépondérante. Trois espèces se sont révélées les plus abondantes et les plus représentatives du peuplement, et ont pour cette raison fait l’objet de toutes les analyses :Gilbertiodendron dewevrei J. Léonard (De Wild), Scorodophloeus zenkeri Harms et Uapaca guineensis Mull. Arg. Elles ont toutes les trois montré une structure agrégée, et cela à toutes les échelles d’analyse. Les agrégats observés présentent un rayon d’environ 25 m de distance. L’analyse en fonction du diamètre a indiqué une structure agrégée pour les plus petits diamètres et régulière pour les plus grands, et que l’agrégation diminue avec le diamètre. Les résultats montrent également que les structures spatiales observées dépendent de l’échelle d’analyse considérée et de la méthode utilisée. La méthode de Clark & Evans est sensible à la variation de l’étendue. À cet égard, l’échantillonnage aléatoire de Hines & Hines est apparu plus adaptée que celui de Clark & Evans. Les fonctions de Ripley et dérivées sont des outils efficaces et apportent plus d’information. Les analyses ont montré des associations positives entre G. dewevrei et S. zenkeri, ainsi qu’entre G. dewevrei et U. guineensis. Par contre, une indépendance a été constatée entre S. zenkeri et U. guineensis. Il apparaît également que les jeunes sont associés positivement aux adultes supposés reproducteurs, et que les immatures sont indépendants par rapport aux adultes. La dispersion faible ou limitée des graines à proximité des arbres parents en est le principal facteur endogène responsable. Ce facteur explique également les associations positives entre les jeunes et les adultes de la même espèce. La compétition entre des individus pour les besoins en espace, en lumière ou en nutriments dans le sol, explique la structure régulière observée ainsi que l’indépendance des immatures envers les adultes. La dispersion limitée n’est cependant pas le seul facteur explicatif de l’agrégation spatiale des arbres, d’autres facteurs tels que l’hétérogénéité environnementale (sol, topographie,…) ou la perturbation sont vraisemblablement aussi impliqués. La perturbation anthropique ou naturelle est un processus écologique qui devrait avoir joué un rôle déterminant dans l’organisation spatiale des communautés de la forêt. Combinée au phénomène de masting, aux effets de Janzen-Connell et aux ectomycorhizes, elle est très probablement à la base des structures spatiales et des relations spatiales observées entre les espèces de la RFY. / Tropical forests contain tree-stands with management and planning requires knowledge of their spatial organization and dynamics. Analyses of the spatial structure of tree species can be used in natural forests to identify the underlying mechanisms that structure of forest stands to improve the understanding of the relationships between species. This study was conducted in the Yoko Forest Reserve (YFR) around Kisangani in eastern DR Congo (DRC). It is to analyze the horizontal spatial structure of the most abundant species and to try to identify, on a local scale, factors and / or potentially explicative ecological processes to draw valuable lessons for the management of forest areas nearby Kisangani. <p>To do this, a sample plot of 25 ha (500m x 500m) was delineated in the southern block of the YFR consists of a mixed woody vegetation and semi-deciduous. A forest inventory was used to analyze the floristic and structural composition of the area. Three techniques (or statistical models) under spatial ecology for the analysis of the horizontal structure of the species were used: the nearest neighbor method of Clark & Evans (1954), the random sampling method Hines & Hines (1979), these two methods based on a single scale analysis of the study plot, and the method of Ripley (1977) allows not only a multi-scale analysis, but also the study of intra- and inter-relationships. Regarding this point, the trees were categorized into three stages of development on the basis of their diameters (young, immature and adult). A comparative and theoretical analysis of the three methods was performed. <p>A total of 169 species belonging to 36 families with 114 genera have been identified and the family Fabaceae which the majority of species belong to the subfamily Caesalpinioideae appeared decisive. Three species have proved the most abundant and the most representative of the stand, and for this reason the subject of all analyzes Gilbertiodendron dewevrei J. Léonard (De Wild) Scorodophloeus zenkeri Harms and Uapaca guineensis Mull. Arg. They all three showed an aggregated structure, and that all scales of analysis. Observed aggregates have a radius of about 25 m distance. The analysis based on the diameter indicated an aggregated structure for smaller diameters and regular for larger and that aggregation decreases with diameter. The results also show that the observed spatial structures depend on the considered analysis of scale and the method used. The method of Clark & Evans is sensitive to the variation in the extent. In this regard, the random sampling of Hines & Hines appeared more suitable than that of Clark & Evans. Ripley's functions and derivatives are effective tools and provide more information. Analyses showed positive associations between G. dewevrei and S. zenkeri, and between G. dewevrei and U. guineensis. By against, independence was found between S. zenkeri and U. guineensis. It also appears that young people are positively associated with the supposed breeding adults and immatures are independent compared to adults. The low or limited seed dispersal near parent trees is the main endogenous factor responsible. This factor also explains the positive associations between youth and adults of the same species. The competition between individuals to space requirements, light or nutrients in the soil, explains the observed regular structure and the independence of immature towards adults. Limited dispersal, however, is not the only factor explaining the spatial aggregation trees, other factors such as environmental heterogeneity (soil, topography, ) or disturbance are probably also involved. Anthropogenic or natural disturbance is an environmentally friendly process that should have played a decisive role in the spatial organization of forest communities. Combined with masting phenomenon, the effects of Janzen-Connell and Ectomycorrhizae, it is very probably the basis of spatial structures and spatial relationships observed between species of the YFR.<p><p> / Doctorat en Sciences agronomiques et ingénierie biologique / info:eu-repo/semantics/nonPublished