Dissertations / Theses on the topic 'Facilitation and competition'

Plants established next to other plants may have their performance positively or negatively affected by their neighbors, which characterizes interactions of facilitation and competiton, respectively. Facilitation and competition, however, may occur simultaneously and understanding which one predominates in pairwise interactions under different ecological contexts is important to comprehend the structure of plant communities. According to the stress-gradient hypothesis (SGH), facilitation is expected to prevail in more severe environments, but the outcome of interactions may change depending on features of interacting individuals, such as life form and ontogenetic stages. As harsh conditions and high diversity of plant life-forms are characteristic of coastal dunes, the amount of studies about plant interactions has been rapidly increasing in these ecosystems, with apparent divergent conclusions. However, until now, there is not a systematic and quantitative synthesis about the factors affecting the net outcome of facilitation and competition in these ecosystems. We conducted a meta-analysis to investigate the effects of environmental stress and the simultaneous influence of plant life form and ontogenetic stage on the outcome of facilitation and competition in coastal dunes around the world. We used four performance measures to estimate the outcome of interactions: abundance, survival, growth, and reproduction. Contrary to what was expected by SGH, we found that negative impacts of neighbors on plant reproduction increase towards more arid conditions, but this effect was not observed for other performance measures. Our results also indicate that woody neighbors facilitate the survival of woody seedlings and the reproduction of herbs, while herbaceous neighbors facilitate the growth of other herbaceous plants. Overall, the outcome of plant interactions in coastal dunes depends on the performance variable measured and on both environmental conditions and plant features, indicating an interaction between these factors. Such interaction and different mechanisms underlying facilitation and competition should be more investigated in the future. The global scale of our meta-analysis supports generalization of important processes of succession and conservation in coastal dunes. Benefits of woody neighbors to the survival of woody seedlings corroborate the concept of successional feedbacks in the beach-inland physiognomic gradient, and our results reinforce the use of nurse plants in coastal dunes as a valuable tool to restoration of these endangered ecoystems
Plantas estabelecidas nas proximidades de outras plantas podem ter sua performance afetada positiva ou negativamente por seus vizinhos, caracterizando, respectivamente, interações de facilitação e competição. Considerando que ambas as interações podem ocorrer de forma simultânea, compreender o predomínio de cada uma delas em diferentes contextos ecológicos é fundamental para o entendimento da estrutura de comunidades vegetais. De acordo com a hipótese do gradiente de estresse (HGE), a facilitação tende a predominar em ambientes mais severos, mas o balanço das interações depende ainda das características dos indivíduos envolvidos, como forma de vida e estágio ontogenético. Como condições ambientais severas e alta diversidade de formas de vida vegetais são características de planícies costeiras, o número de estudos investigando interações entre plantas tem aumentado rapidamente nesses ambientes, com conclusões aparentemente divergentes. No entanto, ainda não há uma síntese sistemática e quantitativa dos fatores que afetam o balanço entre facilitação e competição nesses ecossistemas. Nós realizamos uma meta-análise em escala global para investigar os efeitos do estresse ambiental e a influência simultânea da forma de vida e do estágio ontogenético das plantas sobre o balanço entre facilitação e competição em planícies costeiras. Utilizamos quatro variáveis de performance para estimar o balanço de interações: abundância, sobrevivência, crescimento e reprodução. Ao contrário do predito pela HGE, encontramos que impactos negativos de plantas vizinhas sobre a reprodução de outras plantas aumentam com a aridez do ambiente, mas que esse efeito não é observado para outras variáveis de performance. Nossos resultados também mostram que vizinhos lenhosos facilitam a sobrevivência de plântulas lenhosas e a reprodução de plantas herbáceas, enquanto vizinhos herbáceos facilitam o crescimento de outras herbáceas. De modo geral, o balanço das interações depende da variável de performance medida e tanto de condições ambientais quanto de características das plantas, indicando que esses fatores interagem. Tal interação e os diferentes mecanismos subjacentes à facilitação e à competição devem ser melhor investigados no futuro. A maior sobrevivência de plântulas lenhosas na presença de vizinhos lenhosos corrobora a ideia de retroalimentação positiva no processo sucessional que caracteriza o gradiente fisionômico da praia ao interior. Nossos resultados também reforçam o potencial do uso de plantas-berçários como ferramenta para restauração de planícies costeiras degradadas

Increased knowledge of plant interactions is important for our understanding of how ecosystems will respond to climate changes. Using four common low-herb and three tall- herb species as phytometers I measured the net outcome of plant interactions in an alpine environment by a neighbour removal experiment. Grazing and climate were tested as explanatory factors for differences in the outcome of plant interactions, with two altitudes representing different climates. The most important finding in this experiment is that competition is the dominating interaction among plants in this habitat, regardless of plant size, climate and grazing. Climatic exposure and grazing only influenced tall-herb species while low-herbs were mainly limited by competition, presumably for light. These results are important since facilitative interactions and net facilitation in plant communities are often reported to become more common in severe climates.

AN ABSTRACT OF THE THESIS OF LEWIS KENNEDY REED, for the Master of Science degree in PLANT BIOLOGY presented on October 30, 2009, at Southern Illinois University Carbondale TITLE: COMPETATIVE EVQUIVALENCY OF CULTIVAR AND NON-CULTIVAR DOMINANT GRASSES IN AN EXPERIMENTAL RETORATION MAJOR PROFESSORS: Dr. Sara G. Baer and Dr. David J. Gibson Multiple population sources of species for use in prairie restoration exist, including cultivars and non-cultivars of dominant native grasses. However, little is known about the competitive equivalency of different population sources of dominant C4 grasses and whether intraspecific variation in their competitive effect on the community scales to affect ecosystem assembly. In 2006, an experimental restoration was established in a former agricultural field using cultivars and non-cultivars of the dominant grasses (Andropogon gerardii, Sorghastrum nutans, and Schizachyrium scoparium) and two different species pools of non-dominant species containing equal richness and distribution of species among functional groups. I evaluated inter - and intra-specific variation in the competitive equivalency of A. gerardii, S. nutans, and S. scoparium in terms of community response to their individual and complete (all three species) removal. Removals were maintained for two growing seasons, though little maintenance was needed after the initial treatment. The competitive effect of each grass species and population source was determined by calculating a response ratio for percent cover and above ground primary productivity between removal and control plots for each dominant grass individually, forbs, non-dominant grasses, and legumes which were each analyzed using a mixed model procedure for a split-split-plot randomized block design. Effects of removals on overall community composition were assessed using non-metric multi-dimensional scaling (NMDS) and analysis of similarity (ANOSIM). In addition I monitored changes in photosynthetically active radiation (PAR) at the soil surface and inorganic soil nitrogen. Based on the 2008 data (after two years of maintaining removals), several important differences were detected between the focal species and in some cases the two sources of the same species in terms of their effects on neighbors, inorganic soil nitrogen, and PAR. Many of these differences depended on which species pool the comparison took place in. A 3-way interaction was detected between species pool, source, and removal treatment in the volunteer forb group (F3, 60 =3.28, p = 0.0268). Volunteer forbs showed a positive response to removal of cultivars of the dominant grass functional group in one species pool but not the other. A three-way interaction was detected between species pool, source, and removal treatment in terms of Bray-Curtis similarity (F3, 60 = 2.91, p = 0.0417). Within one of the species pools, similarity values of communities were higher between removals and controls in cultivar plots than in non-cultivar plots where A. gerardii was removed. While NMDS ordination showed separation of some plots by dominant grass source, within group variation was higher than among group variation and ANOSIM deemed this separation insignificant. Planted forb ANPP exhibited a significantly negative response to removal of S. nutans that was not observed in the other removal treatments (F3,41.1 = 3.09, p = 0.038) suggesting facilitation by the dominant grass on planted forbs. The aboveground net primary productivity (ANPP) of the subordinate community (i.e. all species except the dominant grasses) exhibited a 3-way interaction among species pool, dominant grass source, and species removed (F3,61 = 2.76, p=0.0499). This interaction resulted from a stronger negative response of subordinate community ANPP to cultivars of the dominant grass functional group removal than non-cultivars of this group that only occurred in one of the species pools. There was a significant main effect of species removed on %PAR at ground level (F3, 60 = 4.84, p = 0.0044). All removal treatments allowed higher penetration of PAR to ground level, with S. nutans and all dominant grass removal having the strongest effect. Inorganic N availability was lower in response to removal of A. gerardii cultivars compared to the removal of non-cultivars of this species in one species pool but not the other. Removal of cultivar S. scoparium lead to a positive response in total inorganic nitrogen while removal of non-cultivars of this species lead to a negative response in total inorganic nitrogen in this same species pool (F3, 51 = 3.61, p = 0.018). Results demonstrate that inter- and intra-specific variation among these dominant species affect some aspects of community structure and ecosystem properties, but these effects are not consistent among dominant species and among subordinate species pools. These complex interactions may have important implications for restoration and land management.

Understanding forest resilience to novel disturbances and how tree interactions will be affected by global change is critical for predicting future forest composition. The widespread decline of the endangered whitebark pine (Pinus albicaulis) in the Canadian Rockies due to non-native white pine blister rust (Cronartium ribicola) and native mountain pine beetle (Dendroctonus ponderosae) and Ips sp. permitted examination of interactions between disturbances and tree responses in high-elevation forests. Disturbance severity was high with 20-90% whitebark pine mortality over 50 years in 16 stands. Basal annual increment (BAI) of whitebark pines prior to mortality from mountain pine beetles declined 46%, but only by 25% for those subsequently killed by blister rust and Ips sp.. Climate-growth relationships suggest blister rust increased sensitivity of whitebark pines to variation in summer precipitation, reducing resistance to beetles. The mortality of whitebark pine was used as an in-situ experiment simulating neighbour removal to test the stress-gradient hypothesis of tree interactions. Facilitation intensity, determined by comparing subalpine fir (Abies lasiocarpa) regeneration around live, top-killed, or dead adult whitebark pines, increased with elevation but depended on benefactor size and neighbourhood density. Large-diameter, top-killed whitebark pines were more facilitative than live trees, indicating thresholds in benefactor size, below which live, healthy trees were facilitators and above which they were competitors. Size thresholds were also found in interactions between adult trees where competition intensity increased between trees of greater diameter differences as indicated in BAI releases of subalpine fir after the death of neighbouring trees. Conversely, the importance of competition relative to other factors influencing growth increased between trees more similar in size and with abiotic stress. My results refine the stress-gradient hypothesis by demonstrating hierarchical influences on tree interactions. The predominant release from competition doubled subalpine fir’s BAI from the landscape average pre- disturbances compensating for the decline predicted by climate-growth relationships. Lack of regeneration and growth release in surviving whitebark pines and an abrupt shift in key variables suggest a regime shift to fir dominance and whitebark pine extirpation. Whitebark pine resilience was higher at sites of low abiotic stress and disturbance severity, relationships useful for conservation.

Diss. (sammanfattning) Stockholm : Stockholms universitet, 2010.
At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Manuscript. Paper 4: Manuscript. Paper 5: In press.

Community composition results from an integrated combination of random processes, regional habitat spatial structure, local environmental conditions, and species interactions. For example, the outcome of plant interactions can change depending on local environmental conditions such as nutrient availability, land management, or herbivory intensity. In particular, plant interactions may vary between facilitation and competition depending on ecological context, with facilitation expected to be prevalent under stressful conditions. I present the results of four studies that address different aspects of the community assemblage and dynamics emphasizing the synergistic effect of different processes. In the first, I investigated the importance of habitat isolation in determining species richness of wetlands with contrasting land use. The second describes an experiment to test the hypothesis that plant interactions with an unpalatable plant (Juncus effusus) would range from competition in ungrazed areas to facilitation in grazed areas and predicted that facilitative effects of Juncus would differ among functional groups of beneficiary species and be strongest when grazing was intense. In the third, I examine the community composition impacts of Juncus and predicted that Juncus would preserve functional diversity in grazed wetlands but that the effects of Juncus would vary along a grazing gradient. The fourth study investigated the relative importance of competition and nutrients in determining wetland invasion in two different land use types. Broadly, I demonstrate that the importance of different processes (habitat isolation, nutrient availability, competition/facilitation) to community composition is dependent on ecological conditions. This integrated view of community dynamics is interesting from a purely ecological perspective but also can be applied to understanding ecological problems such as exotic invasions and restoration of disturbed habitats.
Ph.D.
Department of Biology
Sciences
Conservation Biology PhD

The low-lying topographic nature of salt marshes makes plants in these communities particularly vulnerable to increased salinity and inundation exposure associated with sea level rise. Both increased salinity and inundation have been cited as major causes of reduced plant performance and survival in marsh and areas fringing marsh. In addition to limitations imposed by physical stress, interspecific interactions have also been shown to mediate the performance and survival of salt marsh and salt marsh fringing species. The Stress Gradient Hypothesis (SGH) postulates that species interactions shift from competitive to facilitative as stress levels increase and predicts that (a) the frequency and intensity of facilitative interactions increase as conditions become more stressful for plants and (b) the strength of competitive interactions increases as abiotic stress levels diminish. The SGH has been rigorously tested to examine how both the frequency and intensity of species interactions change under varying physical stress levels. Studies conducted in salt marsh systems have shown facilitation to be as strong of a driving force as competition in influencing plant performance and survival and have shown that while competition appears to be the pervasive force in the less physically stressful terrestrial zones fringing salt marshes, facilitation influences the performance and survival of species in harsher marsh areas. Under conditions of sea level rise, it remains unclear if the nature of interspecific interactions would shift as stress levels change. This research endeavors to examine the interplay between abiotic stresses and biotic interactions under conditions of increased salinity and inundation exposure.

The first study presented here investigated the effects of increased inundation and soil salinity associated with sea level rise on four salt marsh fringing species, and assesses how competition and facilitation impact survival of salt marsh fringing plant survival under these changing conditions. All plant species experienced reduced growth and photosynthetic inhibition below their current distributional positions, both in the presence and absence of neighboring above ground vegetation. The findings also signal a potential shift in the nature of interspecific interactions from competition to facilitation to neutral as plants begin to experience increased salt and inundation exposure.

The second study aimed to disentangle the effects of increased soil salinity and increased soil moisture on four salt marsh fringing species, and to examine the effects of plant neighbors. The results showed that fringe plants exposed to increased inundation experienced a two-fold reduction in performance and survival over 750 g pure salt addition, suggesting that inundation may be a more important limiting factor than salinity with rising sea levels. Landward transplants at the forest-fringe margin exposed to lower soil salinity and decreased inundation exhibited a three-fold increase in performance and survival when compared to controls. Neighbor manipulation studies, which consisted of trimming neighboring vegetation to ground level, again suggested that interspecific interactions in salt marsh fringing species may shift from competitive to facilitative with climate-induced sea level rise. Overall, our findings suggest that salt marsh fringing species may not be able to tolerate changing conditions associated with sea level rise and their survival may hinge on their ability to migrate towards higher elevations.

The final experiment tested the Stress Gradient Hypothesis and investigated the relative importance of facilitation and competition in a salt marsh system under varying stress levels. This study also ascertained whether salt or inundation exposure is the primary influence on salt marsh plant performance and survival. As in previous studies, our findings suggest that many salt marsh plants don't require, but merely tolerate harsher abiotic conditions. The results showed that plants at higher elevations were depressed by strong competitive pressure from neighboring fringe species while plants at lower elevations benefited from the presence of neighbors. Collectively, the results of these studies indicate that species interactions are an integral driver of plant distribution in salt marsh communities. Furthermore, our findings indicate that changing stress levels may not always result in a shift in the nature of interspecific interactions. These studies have endeavored to show that the interplay between competition and facilitation interacts with physical processes to determine the growth and performance of both fringe and marsh plant species. The paucity of studies examining the roles of species interactions and changing abiotic stress levels on multiple salt marsh and salt marsh fringing species warrants the need for additional research. The responses of salt marsh and salt marsh fringing species to sea level rise can not only serve as very valuable and sensitive indictors of climate change, but will also aid in predicting the future location of the marsh-fringe-forest ecotone, which is predicted to shift inland as sea levels continue to rise.

Pollinator sharing in mixed species communities is expected to significantly contribute to mating patterns in contemporary populations but may also affect the evolutionary trajectory of traits associated with plant mating. In this thesis, I considered how the spring environment and pollinator sharing may contribute to the widespread convergence in traits among spring flowering species using comparative biology. The proposed correlation between a spring flowering phenology and white or light floral colour, fleshy fruits, woody growth forms and understory occupation is confirmed. In addition, I examined the effects of pollinator responses to community and population traits to determine the relative importance of inter- and intraspecific interactions in pollinator mediated reproductive success of a spring flowering species, Trillium grandiflorum. In this study, the reproductive success of T. grandiflorum was pollen limited. However, the magnitude of pollen limitation was influenced only by intraspecific density and varied independently of community diversity. The results of this thesis contribute significantly to our understanding of pollinator-mediated interactions in spring flowering communities but also highlight future avenues of investigation.

Most animals grow substantially during their lifetime and change in competitive ability, predatory capacity and their susceptibility to predation as they grow. This thesis addresses the implications of this on regulation and dynamics within populations as well as between population interactions. In size-structured populations either reproduction or maturation may be more limiting. If juveniles are competitively superior, the competitive bottleneck will be in the adults and reproduction will be limiting. Mortality will in this case result in overcompensation in juvenile biomass through increased reproduction. Compensation in biomass was demonstrated in Daphnia pulex populations subjected to size-independent mortality, where juvenile biomass did not decrease when a substantial harvest was imposed due to increase per capita fecundity. This supported that juveniles were superior competitors and that population cycles seen in Daphnia are juvenile-driven. Compensatory responses in biomass may lead to that predators facilitate eachothers existence by feeding on a common prey, a phenomenon coined emergent facilitation. In an experimental test of the mechanism behind emergent facilitation it was demonstrated that the invertebrate predator Bythotrephes longimanus was favoured by thinning of its prey Holopedium gibberum. The thinning mimicked fish predation and targeted large individuals while Bythotrephes preferrs small prey. Size dependent predation also occurs within populations, i.e. cannibalism, were large individuals feed on smaller conspecifics. Two populations of the common guppy (Poecilia reticulata) originating from different environments were demonstrated to differ in cannibalistic degree. Cannibalism was also affected by the presence of refuges and females and juveniles from one population were better adapted to structural complexity than the other. The effects of these differences in cannibalism on population regulation and dynamics were studied in long term population experiments. Both populations were regulated by cannibalism in the absence of refuges, and displayed cannibal-driven cycles with suppression of recruitment and high population variability. The presence of refuges decreased density dependence and population variability and harvesting of large females in the absence of refuges led to population extinctions in the more cannibalistic population. The less cannibalistic population had higher population biomass and stronger density-dependence in the presence of refuges. When refuges were present, cohort competition increased and cycles with short periodicity were seen. Large individuals were not only cannibals, but could successfully prey on other species. Small and large guppies were allowed to invade resident populations of Heterandria formosa. Small invaders failed while large invaders succeeded as predation from large invaders broke up the competitive bottleneck that the resident population imposed on juveniles of the invader.

This thesis investigated aspects of plant recruitment across an altitudinal gradient of mountain tops in the Victorian Alps, Australia, and provides a baseline for the patterns and processes of alpine plant recruitment in the absence of large-scale disturbance. The patterns in alpine vegetation across the study sites were described in relation to abiotic environmental factors. Temperatures were lower and precipitation was higher at the high altitude sites. The vegetation did not differ significantly between the sites, although sites at low altitudes were shrubbier than those at high altitudes. Analysis of the soil seed bank revealed high mean densities of germinable seed (80 to 1400 seeds m-2) across the gradient of sites. The similarity between the seed bank samples and the standing vegetation was low (qualitative similarity: 0.08 to 0.2; quantitative similarity: 0.03 to 0.19). In laboratory germination experiments, I found rapid and substantial germination. Final percent germination was above 90% for most species. One species, Aciphylla glacialis, showed evidence of dormancy mechanisms. In subsequent experiments, I found that innate primary seed dormancy in this species could be broken with cold-wet stratification. There were no significant patterns in natural seedling recruitment across the altitudinal gradient. Similarities between the seedling flora and the standing vegetation were low (qualitative similarity: 0.18 to 0.45: quantitative similarity 0.04 to 0.09). Mean seedling density was best predicted by a combination of soil wilting point, altitude and plant litter. In some cases, seedling density was greater than 80 seedlings m-2. The relative importance of either negative (competitive) or positive (facilitative) interactions between seedlings with adjacent vegetation were investigated in relation to seed germination, seedling growth and seedling survival. Facilitative interactions were common at the higher altitude sites. At lower altitudes, facilitative and competitive interactions were common. Without close neighbours at high altitudes, seedlings were unlikely to survive into their second year. An understanding of plant recruitment can provide a useful basis for predicting species responses to large-scale disturbance and climate change.

Invasive plants can displace native species, deteriorate native forest, and change plant communities and ecosystem functions. Native plant populations are fundamentally impacted by invasive species because of the interactions between invasive species and native plants. This study focuses on understanding the extent, mechanisms and consequences of interaction between a non-indigenous invader Ailanthus altissima and its functionally similar native species Robinia pseudoacacia in the Mid-Appalachian region, from an individual scale to a regional scale. These two subject species are common and coexist in early-successional eastern deciduous forest. The interactions between these two common species are important to community structure and canopy tree regeneration. To address the type and extent of interactions of these two species, a greenhouse experiment utilizing various species proportions, nutrient levels and seed sources was performed. In addition, a common-garden experiment with various species densities and proportions over three consecutive growing seasons was performed in a more natural condition than that of the greenhouse experiment. We found at the seedling stage, the dominant interaction was competition, and R. pseudoacacia was the winner both above- and belowground. The allelopathic compounds of A. altissima may have inhibited nodulation of R. pseudoacacia. Ailanthus altissima seedlings from its native region had slightly stronger competitive abilities compared with the seedlings from its invaded range. In the common garden experiment, R. pseudoacacia plants grew quicker than A. altissima, but A. altissima inhibited the growth of R. pseudoacacia by interspecific competition. The negative impact of A. altissima on R. pseudoacacia became larger as time progressed. To assess the community-level consequences of the two species, we conducted a forest mapping and a complete target-tree-based forest survey, and analyzed regional-scale data from the Forest Inventory Analysis Data Base. The two target species were significantly associated with themselves and with each other. Community species composition and diversity were significantly different across sites. A negative impact of both species on the understory community diversity and tree regeneration at the neighborhood scale was detected; while at a regional level, tree diversity in the FIA plots with either A. altissima or R. pseudoacacia was higher than the reference plots.
Ph. D.

Morella cerifera is a rapidly expanding native shrub on the Virginia barrier islands which displaces other native coastal species and may interrupt normal sediment dynamics. Barrier islands are considered stressful environments with low nutrients, high solar load, and frequent drought and salt exposure; facilitation often dominates in stressful environments according to the Stress Gradient Hypothesis. The objective of this project was to understand the importance of species interactions with grasses on the growth and physiology of M. cerifera at the seedling life stage through both field and lab experiments. Grasses provided ~1.3°C insulation to shrubs during winter freeze events and a freezing threshold for M. cerifera seedlings was experimentally found between -6°C and -11°C. Seedlings competed for light with grasses during warm months and grew more where grasses were clipped, revealing a tradeoff between winter insulation and summer light competition. M. cerifera shows evidence of ecosystem engineering at the seedling stage by significantly reducing summer maximum temperatures. This enables rapid expansion of M. cerifera across the landscape. As M. cerifera expands, island migration is altered, leading to decreased island stability and increased erosion. Although seedlings are small and relatively vulnerable, this life stage appears to have significant implications for the ecosystem trajectory and stability of the Virginia barrier islands.

Cette thèse visait à caractériser et à modéliser la dynamique des trois ressources principales que sont l'eau, l'azote et le rayonnement dans une association en relais maïs-plante de couverture en semis direct en région tropicale subhumide et ses conséquences sur la productivité de la culture principale et de l'ensemble du système et sur les flux d'eau et d'azote dans le sol. Le travail repose sur un dispositif expérimental mis en place dans les Cerrados, à Unaí (Minas Géras, Brésil) en 2007 pour deux années agricoles, d'une part en station expérimentale permettant un effort de mesure intense et d'autre part dans un réseau de parcelles d'agriculteurs offrant une large gamme de situations pédoclimatiques et de modalités de mise en œuvre des systèmes de culture étudiés. Dans ces derniers, la culture principale était un maïs, et la plante de couverture associée en relais était une culture fourragère tropicale, soit la légumineuse, Cajanus cajan soit la poacée, Brachiaria ruziziensis. Deux dates de semis, précoce et tardive étaient considérées pour le semis de la plante de couverture au sein du maïs, dans le cas de l'essai en station. Ce travail montre que rendement du maïs n'a pas été significativement affecté par la présence de la plante de couverture semée précocement au sein du maïs, en comparaison avec le rendement du maïs en culture pure. En revanche, la production en biomasse de la plante de couverture a été significativement réduite dans l'association. Le semis précoce de la plante de couverture au sein du maïs permet une production en biomasse significativement plus importante qu'un semis tardif (20 jours après floraison du maïs), et une production totale en biomasse de l'ensemble de l'association significativement plus importante par rapport aux cultures pures. Cette production totale a été plus du double comparée à celle de la culture pure de maïs dans le cas de l'association maïs-Cajanus. Les valeurs du Land Equivalent Ratio (LER) du rendement en grain de maïs et de la production de biomasse ont été supérieures à 1, atteignant jusqu'à 2.03 témoignant d'une meilleure valorisation des ressources par les espèces associées et l'avantage de tels systèmes pour produire à la fois du grain et du fourrage. La thèse montre aussi la forte variabilité du potentiel de production de ces systèmes en parcelles de producteurs et suggère que lorsque les ressources en eau et en azote sont moins disponibles que dans l'essai de station, des compétitions entre maïs et plantes de couverture peuvent réduire les performances agronomiques du maïs et/ou de la plante de couverture comparativement par rapport à la culture pure équivalente. La biomasse totale produite reste cependant supérieure à celle de la culture pure de maïs, sauf pour de rares exceptions où de plus faibles disponibilités en ressources, parfois accentué par des difficultés de gestion technique, le maïs semble souffrir de compétition plus forte et/ou la plante de couverture ne produit que très peu de biomasse. La thèse évalue la capacité d'un modèle de simulation de culture associées, STICS-CA, à fournir une analyse plus fine de la dynamique du partage des ressources dans ces systèmes et à en estimer les performances agronomiques et certains impacts environnementaux en fonction de conditions techniques et édapho-climatiques différentes de celles de notre dispositif. Des modifications mineures et le calage du modèle ont conduit à une simulation satisfaisante des cultures pures de maïs et Brachiaria et de l'association maïs-Brachiaria en semis direct sur mulch pailleux. En revanche il n'a pas été possible d'obtenir une simulation satisfaisante du maïs en semis conventionnel sur sol labouré, ni du Cajanus en culture pure et de l'association maïs-Cajanus en semis direct. . La thèse fournit des informations sur les imperfections du modèle et propose des améliorations aussi bien en termes de formalismes pour la prise en compte de la minéralisation du mulch, qu'en termes de dispo exp
This thesis dealt with characterizing and modeling the dynamics of the three principal resources which are water, nitrogen and radiation in a no-tillage relay intercropping maize-cover crop systems, under sub-humid tropical area climate. Moreover, productivity of the main crop and of the whole system and nitrogen and water flows in the soil are assessed.The study was based on an on-field approach/experimental design carried out during two crop growing seasons in Cerrados region, in Unaí (Minas Géras, Brazil) since 2007. Firstly, an experimental station was used allowing an intense effort of measurement and secondly, a whole of several farmer's fields offering a wide range of soil and climate conditions and modalities of implementation of the studied cropping systems. In the latter, main crop was maize, and intercropped cover crop was a tropical fodder crop, either leguminous, Cajanus cajan or gramineous, Brachiaria ruziziensis. Two dates of sowing, early and late were considered for the sowing of the cover crop in established maize, in the case of the experimental station. This study showed that maize yield was not significantly affected by the presence of the relay cover crops in comparison with maize as the sole crop, even when the cover crop was sown soon after maize emergence in comparison with maize sole crop. In contrast, the production of biomass by the cover crop was significantly lower when grown with maize than when it was grown as a sole crop. In the intercropped systems, when sown early, the cover crop produced higher total biomass than when sown late (20 days after maize flowering). Moreover, total aboveground biomass production of maize intercropped with a cover crop was much higher than that of any of the crops sown alone. Total biomass produced by maize and pigeon pea (Cajanus cajan) was more than double that maize grown alone. The Land Equivalent Ratio (LER) values of both maize grain yield and biomass production were higher than one, whatever the intercrop system, reaching up to 2.03 providing better available resources use efficiency by the intercropped plants. Thus, such systems permit to produce both maize grains and forage.The thesis shows also the strong variability of the production potential of these systems in farmer's fields and suggests that when water and nitrogen resources are less available than in the experimental station, competitions between maize and cover crops can reduce the agronomic performances of maize and/or the cover crop when compared to the equivalent pure sole crop. Total biomass produced by intercropping remained however higher than that of maize sole crop, except for rare exceptions where lower resources availability, sometimes accentuated by difficulties of technical practice, maize seems to suffer from stronger competition and/or the cover crop produced only very little biomass. The thesis evaluates the capacity of the STICS intercrop model, STICS-CA, to provide a finer analysis of the dynamics of the resource sharing in these systems and to estimate their agronomic performances and certain environmental impacts according to technical practices and soil and climate conditions that were different from those of our experimental station. Minor modifications and model calibration led to obtain satisfying simulations of the sole crops maize and Brachiaria and intercrop maize-Brachiaria in no-tillage under mulch. On the other hand, it was not possible to obtain satisfying simulations of sole crop maize in conventional tillage, nor of sole crop pigeon pea and intercrop maize-pigeon pea in no-tillage under mulch. The thesis provides information on the model imperfections and as well proposes improvements in terms of formalisms to take into account the mineralization of the mulch, as in terms of experimental design

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O estudo dos efeitos que a diversidade de esp?cies pode causar nos processos ecossist?micos tem crescido vertiginosamente nas ?ltimas duas d?cadas. Diversos trabalhos experimentais realizados no mundo todo t?m demonstrado que uma maior diversidade de plantas contribui para o aumento da produtividade de ecossistemas terrestres. Al?m disso, esse efeito pode influenciar processos em diversos n?veis tr?ficos, contribuindo assim para a estabilidade dos processos ecossist?micos a longo prazo. Paralelamente com os estudos do efeito da diversidade, muita aten??o tem sido dada para desvendar o papel das caracter?sticas funcionais das esp?cies no funcionamento dos ecossistemas. Isto porque as caracter?sticas funcionais das esp?cies t?m se mostrado importantes "pe?as" no entendimento dos efeitos que esp?cies individuais podem exercer nos ecossistemas e suas respostas ao ambiente. Nesta tese de doutorado eu explorei algumas lacunas de conhecimento dentro dessa ?rea em crescente desenvolvimento conhecida na literatura ecol?gica como "biodiversidade e funcionamento dos ecossistemas". No primeiro cap?tulo, eu busquei evid?ncias para mecanismos que podem explicar a rela??o positiva entre diversidade e funcionamento com foco em cinco mecanismos relacionados ?s intera??es entre plantas, tendo como par?metro de funcionamento a produtividade prim?ria. No segundo cap?tulo, eu utilizei t?cnicas para a estimativa de padr?es de diversidade em escalas biogeogr?ficas e bases de dados de sat?lites com longa dura??o para desvendar se a biodiversidade em escalas macroecol?gicas promove a estabilidade da produtividade dos ambientes terrestres no semi?rido brasileiro. Por fim, o objetivo do terceiro cap?tulo foi entender como a perda da cobertura vegetal origin?ria do uso da terra por comunidades tradicionais no semi?rido brasileiro influenciaria os processos de intera??es entre plantas e o papel das caracter?sticas funcionais das esp?cies nessas intera??es. Acredito que a contribui??o individual de cada cap?tulo preenche lacunas de conhecimento importantes dessa ?rea da Ecologia que ainda se encontra em expans?o.

Alpine and arctic tree line expansion depends on the establishment of tree seedlings above the current tree line, which is expected to occur with climate warming. However, tree lines often fail to respond to higher temperatures. Other environmental factors are therefore likely important for tree seedling establishment. Above the tree line, establishing seedlings encounter existing vegetation such as bryophytes, which often dominate in arctic and alpine tundra. Bryophytes modify their environment in various ways and may mediate climate change effects on establishing tree seedlings, and with that tree line expansion. The aim of this thesis was to understand if and how the environment, in particular bryophytes, mediates the impact of climate change on tree seedling establishment at the alpine and arctic tree line. This was explored by reviewing literature on tree seedling establishment at alpine and arctic tree lines globally. In addition, tree seedling survival and growth of Betula pubescens and Pinus sylvestris were assessed experimentally. Here, individuals were planted into mono-specific mats of different bryophytes species and exposed to warming and different precipitation regimes. The literature review revealed that besides from temperature, tree seedling establishment is affected by a wide range of abiotic and biotic factors including water, snow, nutrients, light, disturbance and surrounding vegetation. Furthermore the review revealed that for example vegetation can change tree seedling responses to climate change. The experiments showed that especially tree seedling survival was adversely affected by the presence of bryophytes and that the impacts of bryophytes were larger than those of the climate treatments. Seedling growth, on the other hand, was not hampered by the presence of bryophytes, which is in line with earlier findings that seedling survival, growth and seed germination do not respond similarly to changes in environmental conditions. Moreover, we found several indications that vegetation above the tree line, including bryophytes, mediated tree seedling responses to warming and precipitation or snow cover. This thesis shows that temperature alone should not be used to predict future tree seedling establishment above the alpine and arctic tree line and that extrapolations from climate envelope models could strongly over or under estimate tree line responses to warming. This underlines the value of multi-factorial studies for understanding the interplay between warming and other environmental factors and their effects on tree seedling establishment across current tree lines.

O objetivo geral dessa tese foi compreender melhor o funcionamento das raízes finas (diâmetro <2 mm) de Eucalyptus. Mais especificamente, nosso trabalho teve como objetivo obter informações sobre as mudanças no padrão de absorção de nutriente pelas raízes finas em função da fertilização e profundidade do solo. Foi também, avaliar a dinâmica de crescimento das raízes finas em plantações mistas com uma espécie fixadora de nitrôgenio para testar a hípotese do gradiente de estresse. O capítulo 1 teve como objetivo estudar o efeito da fertilização mineral na especialização funcional das raízes finas de Eucalyptus grandis jovens em camadas profundas do solo (Itatinga-SP). Foram injetados macadores de NO3-15N, Rb+ (K+) e Sr2+ (Ca2+) simultaneamente em uma solução a 10, 50, 150 e 300 cm de profundidade. A determinação das concentrações foliares de Rb+, Sr2+ e a porcentagem de átomos de 15N permitiu estimar o potencial de absorção relativo (PAR) e o PAR específico, definido como PARE, obtido pela razão entre o PAR e a unidade de densidade do comprimento de raízes finas por camada de solo correspondente. O PAR de NO3-15N diminuiu rapidamente com a profundidade e os valores mais altos do PARE de NO3-15N foram encontrados a uma profundidade de 50 cm. O PARE de Rb+ e Sr2+ foi maior a 300 cm de profundidade em relação à camada superficial do solo, com um aumento do diâmetro da raiz e uma diminuição da densidade do tecido radicular com a profundidade. O PARE de Rb+ e Sr2+ a 300 cm de profundidade foi, em média, 88% maior para árvores fertilizadas quando comparado com as árvores não fertilizadas. Os resultados sugerem que a especialização funcional das raízes finas para a absorção de nutrientes é uma característica estável do eucalipto e que pode ser reforçada pela aplicação de fertilizantes. O capítulo 2 focou nos processos ecológicos entre Acacia mangium e Eucalyptus em um gradiente de estresse nutricional. Raízes finas foram amostradas aos 16 e 34 meses após o plantio em blocos casualisados com dois tratamentos: uma mistura com 50% de cada espécie (50A:50E) com e sem fertilização. Durante este período, dois tubos de minirhizotron, perto de eucalipto e acacia, em cada tratamento e bloco, foram utilizados para monitorar o crescimento e o tempo de vida das raízes finas. Aos 16 e 34 meses após o plantio, a DRF de Eucalyptus foi maior em relação à Acacia e maior em F+ do que em F- na camada superior do solo. Este resultado mostrou que, provavelmente, há uma maior competição das raízes de eucalipto nas raízes de acacia em F+ do que em F-. Na camada superficial, a DRF de Eucalyptus em F- foi maior aos 34 meses e perto de árvores de Acacia comparado aos eucaliptos, o que seria consistente com uma maior facilitação de N da Acacia para os Eucalyptus em ambiente com maior deficiência de N (hipótese de gradiente de estresse). A mesma concentração de N nas folhas de Eucalyptus em F + e F- também está de acordo com essa hipótese. A produção de raízes finas de eucalipto entre as duas datas de amostragem foi maior em F- que em F+ em paralelo ao aumento de DRF de eucaliptos perto das acacias. Não foi possível estimar o tempo de vida da raiz, pois não houve mortalidade radicular durante o período de estudo para ambas as espécies. Futuros estudos poderiam ser realizados para uma melhor compreensão dos mecanismos de captação de nutrientes pelas árvores visando um manejo mais sustentável das plantações florestais.
The main objective of this thesis was to better understand the functioning of Eucalyptus fine roots (diameter < 2 mm). More specifically, our work aimed at getting insights into the changes in fines roots nutrient uptake changes depending on soil depth and fertilization. Root growth dynamics was also evaluated in mixed plantations with a nitrogen fixing species to test the stress gradient hypothesis. The first chapter aimed at studying the effect of fertilization on the functional specialization of young Eucalyptus fine roots in deep soil layers (Itatinga-SP). We injected NO3-15N, Rb+ (K+) and Sr2+ (Ca2+) tracers simultaneously in a solution at 10, 50, 150 and 300 cm in depth. Determination of foliar Rb+, Sr2+ concentrations and 15N atom % made it possible to estimate the relative uptake potential (RUP), and the Specific RUP, defined as SRUP, per unit of fine root length density in the corresponding soil layer. The RUPs of NO3-15N decreased sharply with depth and the highest values of the SRUPs of NO3-15N were found at a depth of 50 cm. The SRUP of Rb+ and Sr2+ were higher at 300 cm in depth than in the topsoil with an increase in root diameter and a decrease in root tissue density with depth. The SRUP of Rb+ and Sr2+ at a depth of 300 cm was on average 88% higher for fertilized trees than for unfertilized trees. The results suggest that functional specialization of fine roots for nutrient uptake is a stable characteristic of Eucalyptus that can be enhanced by fertilization application. Chapter 2 focused on the ecological processes between Acacia mangium and Eucalyptus under a gradient of nutrient stress. Fine roots were sampled at 16 and 34 months after planting in a randomized block design with two treatments: a mixture with 50% of each species (50A:50E), with and without fertilization. For each treatment, soil samples were collected in 3 blocks at 0-15; 15-30; 30-50 and 50-100 cm at various distances of Eucalyptus and Acacia trees. During this period, two tubes of minirhizotron near eucalypt and Acacia trees in each treatment and block were used to monitoring fine roots growth and turnover. At 16 and 34 months after planting, fine root mass density (RMD) of Eucalyptus was 30% higher than of Acacia and higher in F+ than in F- in the top soil layer (0-15 cm). This result likely showed higher competition of eucalypt roots on Acacia roots in F+ than in F-. In the superficial layer, RMD of Eucalyptus in F- were higher at 34 months near Acacia trees than Eucalyptus trees, that would be consistent with greater N facilitation of Acacia on Eucalyptus in higher N-deficient environment (stress gradient hypothesis). The same N concentration in Eucalyptus leaves in both F+ and F- was also consistent with this hypothesis. Production of Eucalyptus fine roots between the two sampling dates was higher in F- than in F+ in parallel to the increase of RMD of Eucalyptus near Acacia trees. It was not possible to estimate root life span as no root mortality occurred during the study period for both species. Furthers studies should be conducted to better understand the mechanisms of tree nutrient uptake promoting a better sustainable management of forest plantations.

Stability of coastal systems are threatened by oceanic and atmospheric drivers of climate change. Sea-level rise compounded with increased frequency and intensity of storms emphasizes need for protection of inner island systems by dune formations. Dune building processes are affected by interactions between growth of ecosystem engineering dune grasses and environmental factors associated with disturbance such as sand burial and salt spray. Climate change may also cause latitudinal expansion of some species, resulting in emergence of competitive interactions that were previously absent. Topographic structure of coastlines, traditionally influenced by sand burial, could change as a result of competition emergence. My goal was to determine if species functional trait responses to common abiotic factors are altered by novel and current biotic interactions. I performed a multi-factorial greenhouse experiment by planting three common dune grasses (Ammophila breviligulata, Uniola paniculata, and Spartina patens) in different biotic combinations, using sand burial and salt spray as abiotic stressors. I hypothesized that biotic interactions will cause these dune grasses to shift functional trait responses to abiotic factors that are associated with dune building. I found that plants consistently decreased in biomass when buried. I also found that competition between A. breviligulata and U. paniculata negatively affected dune building function traits of A. breviligulata. This indicates that competition with U. paniculata could alter dune structure. In comparison A. breviligulata had a positive interaction with S. patens, which increased functional trait responses to abiotic stress. Last, we found that competitive intransitivity could occur between these species. My results can be used to make predictions on cross-scale consequences of novel competitive events. This experiment also provides evidence that consideration of local biotic interactions is important in understanding connections between plant level dynamics and large-scale landscape patterns in high stress environment.

Une meilleure connaissance des interactions entre plantes permettrait d’améliorer la gestion de la régénération forestière naturelle et artificielle. Il est bien établi que la molinie (Molinia caerulea), poacée monopoliste du sous-bois, a un effet particulièrement dépressif sur l’installation et la croissance du jeune chêne sessile (Quercus petraea). Son importante capacité à absorber l’eau et les nutriments du sol entraine une forte compétition par exploitation des ressources. Cependant, il semblerait que d’autres interactions soient impliquées dans les relations entre le jeune chêne et la molinie. Ainsi, les propriétés allélopathiques de la molinie ont été suggérées, mais aucune étude n’a mis en évidence un tel effet sur le chêne. À l’inverse, le jeune chêne semble faciliter la croissance de la molinie. L’interaction chêne-molinie présente donc la singularité d’être antagoniste : la molinie inhibe la croissance du chêne tandis que celui-ci favorise la croissance de la poacée, mais les mécanismes impliqués ne sont pas entièrement connus. L’objectif de cette thèse a été d’analyser les mécanismes fonctionnels et écophysiologiques impliqués dans les interactions entre le jeune chêne sessile et la molinie bleue. Les expérimentations réalisées dans la thèse démontrent que (i) le chêne rhizodépose et transfère rapidement de l’azote à la molinie par les voies souterraines (rhizodéposition de composés azotés dans le sol) favorisant la croissance de cette dernière, (ii) la présence de molinie dans le même pot entraine une diminution du taux de racines latérales et du taux d’ectomycorhization du chêne et que (iii) la molinie rhizodépose des métabolites secondaires différents de ceux du chêne et ayant un potentiel allélopathique. Dans la lignée de travaux actuels, la thèse met en perspective que les composés chimiques que les plantes émettent dans le sol agissent comme de véritables signaux directs ou indirects (via les communautés biologiques du sol) et induisent chez les voisins des modifications dans leur métabolisme et/ou leur croissance
A better understanding of plant interactions would improve the management of natural and artificial forest regeneration. It is well established that Molinia (Molinia caerulea), a monopolist species of understory, has a particularly depressive effect on young sessile oak (Quercus petraea) establishment and growth. Large capacity of Molinia in water and soil nutrients uptake leads to strong competition by resources exploitation. However, it seems that other interactions are involved in the relationship between young oak and Molinia. Thus, the allelopathic properties of Molinia have been suggested, but no study has shown such an effect on oak. Inversely, young oak seems to facilitate Molinia growth. Oak-Molinia interaction thus has the singularity of being antagonistic: the Molinia inhibits oak growth while this one favors grass growth, but the mechanisms involved are not entirely known. The aim of this thesis was to analyze the functional and ecophysiological mechanisms involved in the interactions between young sessile oak and Molinia. The experiments carried out in the thesis show that (i) oak quickly transfers nitrogen to Molinia by the underground pathways (rhizodeposition of nitrogen compounds in the soil) favoring grass growth, (ii) Molinia presence in the same pot leads to a decrease in the lateral root rate and ectomycorhization rate of oak and (iii) Molinia rhizodeposes secondary metabolites different from those of oak and have an allelopathic potential. In line with current work, the thesis puts into perspective that the chemical compounds emitted by plants in the soil act as real direct or indirect signals (via the biological communities of the soil) and induce neighbors modifications in their metabolism and / or their growth

Cette thèse avait pour objectif de tester l’effet croisé de la gestion et de sécheresses extrêmes liées au changement climatique sur les interactions biotiques entre plantes, la biodiversité, la composition floristique et le fonctionnement écosystémique de pelouses caractéristiques du sud de l’Europe, en bordure occidentale calcaire du Massif Central français. Des sècheresses printanières et une fauche régulière ont été appliquées expérimentalement suivant un plan factoriel sur deux types de communautés situées, en Dordogne, dans des conditions de sols contrastées: les Mesobromion (pente nulle) et les Xerobromion (pente forte convexe et exposition sud). Les résultats ont validé des modèles théoriques prédisant un effondrement des interactions biotiques à l’extrémité sévère d’un gradient de stress, surtout si le système est soumis à de la perturbation. La réponse des pelouses était très dépendante de la communauté considérée. La sècheresse a diminué la biodiversité et altéré la composition floristique des Xerobromion. Ceci s’explique par la combinaison au sein de cette communauté d’un grand nombre de contraintes (sècheresse expérimentale, stress hydrique dû à l’exposition sud et forte perturbation physique induisant notamment un effondrement de la facilitation et un stress édaphique important). Les Mesobromion étaient plutôt affectés par la fauche qui a augmenté leur biodiversité via une diminution du couvert de graminoides compétitrices. En Europe du sud, et dans un contexte de changement climatique, nous recommandons de maintenir des pratiques de gestion basées sur la perturbation dans les Mesobromion et de se focaliser plutôt sur la conservation d’espèces peu tolérantes à la sècheresse dans les Xerobromion
This PhD aimed at testing the crossed effects of management and extreme droughts linked to climate change on plant biotic interactions, biodiversity, composition and ecosystem functions of extensive grasslands from Southern Europe, at the Western calcareous boundary of the French Central Massif. Extreme spring droughts and a mowing-based management were experimentally applied in a factorial design in the Dordogne department on two communities with contrasted soil conditions: the Mesobromion (null slope) and the Xerobromion (steep convex slope and southern exposure). Our results support theoretical models predicting a collapse of biotic interactions at the severe end of a stress gradient, especially if the system is subject to disturbance. Grassland responses were strongly dependent on local environmental conditions. Drought decreased biodiversity and altered composition in the Xerobromion community. This is explained by the combination of several constraints (experimental drought, water stress due to southern exposition and physical disturbance which induces a collapse of facilitation and an important edaphic stress) in this community. The Mesobromion community was rather affected by mowing, which increased biodiversity through a reduction of competitive graminoids cover. In Southern Europe, and in a climate change context, we recommend maintaining disturbance-based management in the Mesobromion communities and focusing on the conservation of species with a poor tolerance to drought in the Xerobromion communities

Abstract Deciduous shrubs form the tallest type of vegetation in arctic-alpine areas and are important for ecosystem function. In the southern part of the Eurasian tundra zone, willows (Salix spp.) are the most common species in the shrub layer. In the alpine areas of Northern Fennoscandia, willow shrubs are characteristic to areas between tree line and treeless tundra heaths. Vertical structure and composition of willow-characterized tundra vegetation is affected by a variety of ecological factors including climate and herbivory. In turn, the abundance of the willow canopy affects understory species in several ways that still remain inadequately understood. In this PhD work I describe compositional differentiation of willow-characterized vegetation by using a large data set spanning from north-western Fennoscandia to the Yamal Peninsula in north-western Siberia. I studied environmental factors affecting willow-characterized vegetation and willow growth by using correlative analyses. The factors under investigation were latitude, distance from the sea, depth of thaw, position in the slope, industrial disturbance and reindeer grazing. In addition, I examined the relationships between the shrub biomass estimate and composition and species richness of understory vegetation. The effects of reindeer grazing on vegetation in an alpine forest-tundra ecotone were studied experimentally using reindeer-proof exclosures. I found that willow-characterized vegetation is floristically variable and comprises at least eight vegetation types. The most abundant willow thickets typically have a forb-rich understory. The growth of willow increased along with increasing summer temperatures. However the height of willow was more determined by distance from the sea, thaw depth and slope position. Reindeer grazing decreased the abundance of willow and changed the composition of understory vegetation. In addition, industrial activities were detected to have destructed shrub vegetation and turned it into graminoid-dominated vegetation. Shrub canopies facilitated forbs but decreased the cover of all the other groups including dwarf shrubs, bryophytes and lichens. The species richness of vegetation decreased along with increasing shrub abundance. My study shows that arctic-alpine willow vegetation is more diverse than previously thought. There is a predictable relationship between summer temperatures and willow growth. However, the results also show that there are many factors, both physical and anthropogenic, that are likely to complicate this pattern. Most important of these counteracting effects are industrial activities and reindeer grazing. In the areas where shrubs grow in abundance, the species richness of understory vegetation is likely to decrease and forbs are likely to replace other tundra species.

As plantações de eucalipto ocupam 20 milhões de hectares em solos altamente intemperizados. Plantações consorciadas de eucalipto com acácia têm potencial de melhorar a produtividade do povoamento, em relação às respectivas plantações homogêneas, através do efeito facilitador da fixação biológica de N pelas árvores de acácia. O objetivo foi avaliar o efeito do consórcio de genótipos de Eucalyptus com Acacia mangium sobre a produção da biomassa aérea de cada espécie e sobre a ciclagem de N. Quatro experimentos foram estabelecidos em áreas manejadas com eucalipto durante várias rotações, em blocos casualizados, contendo três tratamentos: plantação homogênea de acácia (100A), de eucalipto (100E) e plantação consorciada na proporção 1:1 (50A:50E). As áreas experimentais localizam-se em condições ecológicas diferentes na região sudeste do Brasil, nos Estados de São Paulo e de Minas Gerais. O crescimento das árvores foi medido durante toda a rotação de cultivo e equações alométricas de biomassa e de mineralomassa de N foram estabelecidas em cada área, no final da rotação, para os compartimentos aéreos das árvores. A deposição de serapilheira e a mineralização de N in situ foram avaliadas durante dois anos, no final da rotação. As concentrações de C e N no solo foram avaliadas no final da rotação e estimadas por meio do método NIRS (espectrometria de reflectância no infravermelho próximo) e calculados seus estoques no solo. A fixação simbiótica de N2 foi estimada pela diferença do estoque total de N entre os tratamentos. A produção de biomassa de lenho foi maior em 100E que em 50A:50E, devido ao menor crescimento da acácia, em comparação ao eucalipto, nas condições climáticas estudadas. A matéria seca de serapilheira depositada foi maior em 100E, mas o retorno médio de N ao solo, via deposição, em 100A foi 1,7 vez maior que em 100E. Os estoques de C e N no solo foram iguais nos diferentes tratamentos. Os estoques totais de C e N, na área com o maior crescimento de eucalipto, localizada em Santana do Paraíso, foram 30% e 50% maiores do que na área com menor crescimento, localizada em Itatinga. A mineralização de N no solo foi duas vezes maior em 100A que em 100E, e intermediária em 50A:50E, resultante de uma rápida ciclagem de N, da serapilheira acumulada, abaixo de árvores de acácia. O fluxo de N retornado ao solo com a deposição de serapilheira, no final da rotação, foi um bom indicador da disponibilidade de N no solo, em plantações tropicais de rápido crescimento. Nesse contexto, em termos de manejo florestal, os resultados sugerem que uma rotação com plantação homogênea com espécie arbórea fixadora de N (AFN) após várias rotações com plantios de eucaliptos, pode ajudar a manter a fertilidade de N do solo, com um manejo compatível com as práticas adotadas pelas empresas florestais. Outra opção de manejo em larga escala, seria a introdução de um sub-bosque com espécie AFN em plantações comerciais de eucalipto. No entanto o sucesso destas plantações consociadas, em grande parte, depende das espécies selecionadas.
Eucalyptus plantations cover 20 million hectares on highly weathered soils. Eucalyptus plantations intercropped with Acacia have the potential to improve the productivity of the stand, in relation to their mono-specific plantations, through the facilitatory effect of N fixation by acacia trees. The objective was to evaluate the effect of the consortium of genotypes of Eucalyptus with Acacia mangium on the production of biomass of each species and N cycling. Four experiments were established in areas managed with eucalypts for several rotations in a randomized block design with three treatments: mono-specific acacia plantation (100A), eucalyptus (100E) and mixed plantation at 1:1 (50A:50E). The experimental areas located in different ecological conditions in the southeastern region of Brazil, in the states of São Paulo and Minas Gerais. Tree growth was measured throughout the rotation of cultivation and allometric equations of biomass and mineral mass of N have been set for each area at the end of the rotation to the compartments of the trees. Litterfall and N mineralization in situ were assessed for two years, at the end of the rotation. Concentrations of C and N in the soil were evaluated at the end of the rotation and estimated by the NIRS method (near infrared reflectance spectrometry) and calculated stocks in the soil. The symbiotic N2 fixation was estimated by the difference of the total stock of N among treatments. The wood biomass production was higher in 100E than in 50A:50E, due to lower acacia growth compared to eucalyptus climatic conditions studied. The litterfall dry mass was higher in 100E, but the average return of N from litterfall to the soil was 1.7 times higher in 100A than in 100E. The stocks of C and N in the soil were similar in the different treatments. The total stocks of C and N, with more eucalypts growth, located in Santana do Paraíso, were 30% and 50% higher relative, by less growth, located in Itatinga. The mineralization of N in the soil in 100A was twice that at 100E, and intermediate in 50A:50E, resulting from faster turnover of N in the forest floor under acacia trees. Annual N flux from litterfall at the end of the rotation was a good indicator of soil N availability in fast-growing tropical plantations. In this context, in terms of forest management, results suggest that one rotation of mono-specific N-fixing tree species (NFT) after a certain number of of eucalypt rotations could help to maintain plantation soil fertility, with a management compatible with forest company practices. Other large scale management option for management is the introduction of NFT undergrowth by commercial eucalypts. However, the success of these mixed-species plantations is largely dependt on the selected species.

[Truncated abstract] In order to assess the sustainability of mixed plantings on saline land, this thesis examined the importance of leaf-litter trapping and microsites on recruitment in a salt affected alley-belted (tree/shrub) agricultural system in Western Australia. Located in the low rainfall region (MAR <330 mm) of the wheatbelt, the 60 ha site consists of concentric rows of Eucalyptus sargentii trees with mounded (6 - 11 cm high) 10 -15 m inter-rows of Atriplex spp. Sustainability of this system and fulfilment of productive and ameliorative functions is dependant on successful recruitment (perennials). Although the present study site was conducted on farmland in a Mediterranean-type climate, low annual rainfall and spatial arrangement of perennial shrubs and trees, allow useful comparisons to be made with naturally occurring banded semi-arid systems and vice-versa. Of key interest were leaf-litter redistribution and trapping by tree and shrub rows and whether litter-cover/microsites facilitated/interfered with seedling recruitment (establishment, growth and survival). Litter from the tree row, redistributed by prevailing winds and rain, accumulated adjacent to saltbush seeding mounds, creating a mosaic of bare and littered areas across the site (total litter 10 t/ha over 22 months). Accumulated litter was hypothesized to differentially influence seasonal soil abiotic parameters (depending on litter-cover density) including; salinity, water availability, infiltration rates, water repellency and temperature. These abiotic conditions were also hypothesized to vary between tree and shrub microsites. Biotically, recruitment at this site was also hypothesized to be determined by interactions (positive and negative) between perennial components and understorey annuals/perennial seedlings. Accumulation of litter and resultant heterogeneity was influenced by shrub morphology, microtopography, wind direction and distance from litter source, with increased litter on the leeward sides of hemispherical Atriplex undulata shrubs and shrubs closest to tree rows. ... The importance of tree/shrub microsites varied seasonally, with no influence in winter due to moderate temperatures and increased water availability. In warmer months saltbush mid-row microsites were most favourable for seedling recruitment due to moderate litter-cover; reducing salinity, temperatures and increasing infiltration; and reduced root-competition/shading by the tree row. Tree microsites also directly inhibited seedling recruitment through increased salinities and water repellency. However, trees also indirectly facilitated recruitment in adjacent areas through provision of leaf-litter. As litter-trapping and recruitment patterns at this site mirror those found in semi-arid natural and artificial systems, the results of this study provide useful insights into creating appropriate mimics of low rainfall natural banded woodland and chenopod shrublands. Saltbush seeding mounds, shrub morphology and litter were key components for litter trapping and recruitment heterogeneity at this site. In this tree/shrub alley planting, where litter quantities directly influence vegetation cover densities, future saline plantings need to consider appropriate tree/shrub row spacings and orientation for efficient resource (seeds, litter and water) capture.

La régénération forestière, naturelle ou non, est sujette à de nombreux échecs du fait d’une compétition importante entre les espèces de sous-bois et les jeunes plants d’arbre. L’objectif de ce travail de thèse a été de comprendre les réponses du chêne sessile (Quercus petraea) et d’une poacée, tous deux en compétition dans un contexte de modification de la disponibilité des ressources. Deux espèces de poacées ont été étudiées, la molinie bleue (Molinia caerulea) et la canche cespiteuse (Deschampsia cespitosa). L’accent a été mis sur le rôle des interactions entre différentes ressources (lumière, eau et azote inorganique) sur l’évolution de la compétition (intensité/importance) entre ces espèces et les mécanismes écophysiologiques sous-jacents. Grâce à des expérimentations en conditions semi-contrôlées et à une installation in situ, nous avons pu mettre en évidence une compétition très précoce, dès les premiers mois d’interaction entre le chêne et les poacées voisines. L’abondance de lumière conduit à une compétition plus forte des poacées sur le chêne, amplifiée par l’apport d’azote. Par ailleurs, la nature et l’ampleur de ces interactions varient selon le niveau de stress abiotique appliqué et selon les organes considérés. Nos résultats montrent aussi que le chêne réagit principalement en accumulant les ressources dans son système racinaire, ressources qui sont mobilisées l’année suivante et qui peuvent avoir un arrière effet positif sur le fonctionnement du chêne en absence de stress hydrique. Enfin de la facilitation a également été observée du chêne envers la canche cespiteuse suite à un apport d’azote. Ce dernier améliore la croissance du chêne en compétition, qui pourrait augmenter sa production d’exsudats et son turn-over racinaire au bénéfice de la canche qui valoriserait cette nouvelle source d’azote. Ce travail renforce l’idée d’intégrer l’effet des interactions des différents facteurs abiotiques dans les modèles de compétitions et dans les pratiques de régénération afin d’optimiser la coexistence des espèces forestières, et ce dès la mise en concurrence des jeunes arbres avec les poacées du sous-bois
Temperate forest ecosystems are prone to regeneration failures because of strong competition between understorey species and tree seedlings. This thesis aimed to improve our understanding of sessile oak seedlings (Quercus petraea) and poacea responses, both in competition among different levels of resource availabilities. Two poacea species were studied, Molinia caerulea and Deschampisa cespitosa. We focused on the role of the different resource combinations (light, water and inorganic nitrogen) on interaction variation (intensity and importance) between those species and underlying ecophysiological mechanisms. Our greenhouse and field experiments allowed us to infer a very early competition, from the first months of interaction between oak seedlings and poacea neighbours. High light level leads to stronger competition from poacea on oak seedlings, increased by nitrogen supply. Secondly nature and extent of these interactions depend on applied abiotic stress level and on considered organs. Thirdly Oak seedlings mainly respond by accumulating resources in coarse roots. These resources are remobilized next year and may have a positive carry-over effect on oak functioning if there is no water stress. Eventually we observed facilitation as well, from oak on D. cespitosa in fertilized environment. Nitrogen supply would foster oak seedling growth, increasing exudate production and root turn-over, to the benefit of D . cespitosa. The poacea would take up this extra nitrogen source for its own development. This study support the idea of including interaction effects of different abiotic factors in competition models. Silvicultural practices would also be enhanced by optimizing species coexistence in temperate forests as soon as tree seedlings and understorey species start to grow together

Mixed-species plantations of eucalypts and acacias have the potential to improve stand productivity over that of respective monocultures through the facilitative effect of nitrogen-fixation by acacias, and increased resource capture through above- and belowground stratification. However, growth in mixed-species plantations may not be improved compared to that of monocultures when competitive interactions outweigh the effects of improved nutrient availability and resource capture. Careful selection of sites and species is therefore critical to successfully improving stand productivity using mixed-species plantations. This study set out to examine some of the processes and interactions that occur in mixed-species plantations, and the effect nutrient and water availability can have on the growth of mixtures. In three out of four mixed-species field trials examined in this study, growth was not increased in mixtures compared to monocultures. However, in the fourth field trial, heights, diameters, stand volume and aboveground biomass were higher in mixtures of E. globulus and A. mearnsii from 3-4 years after planting. The range in outcomes from mixing species in these four trials shows that a fundamental understanding of the underlying processes is required to enable a greater predictive capacity for the circumstances under which mixtures will be successful. Therefore the growth dynamics, processes and interactions were examined in the mixtures of E. globulus and A. mearnsii. The difference in productivity between mixtures and monocultures in this trial increased with time up to age 11 years, when 1:1 mixtures contained twice the aboveground biomass of E. globulus monocultures. The positive growth response of trees in mixture compared to monocultures was the result of accelerated rates of nutrient cycling, a shift in C allocation and reductions in light competition through canopy stratification. Nitrogen contents of foliage and soil clearly showed that A. mearnsii influenced the N dynamics in this trial. If these changes in N contents were due to N fixation by A. mearnsii, then about 51 and 86 kg N ha-1 yr-1 was fixed in the 1:1 mixtures and A. mearnsii monocultures, respectively. Nitrogen fixation was also examined using the natural abundance method. The delta15N values of foliage collected at 10 years were grouped according to the mycorrhizal status of the host plant. Therefore the discrimination of 15N during transfer from mycorrhizae to the host plant appeared to vary with mycorrhizal status, and the natural abundance of 15N was not used to quantify N fixation. Rates of N and P cycling in litterfall were significantly higher in stands containing at least 25% A. mearnsii (more than 31 kg N ha-1 yr-1 and more than 0.68 kg P ha-1 yr-1) compared to E. globulus monocultures (24 kg N ha-1 yr-1 and 0.45 kg P ha-1 yr-1). Rates of litter decomposition and N and P release were about twice as high in 1:1 mixtures compared to E. globulus monocultures and were even higher in A. mearnsii monocultures. It is therefore important to select N-fixing species that are capable of cycling nutrients quickly between the plant and soil, and that have readily decomposable litter. The total belowground C allocation was not significantly different between mixtures and monocultures (14 to 16 Mg C ha-1 yr-1). However, since aboveground net primary production was greater in 1:1 mixtures, the changes in nutrient availability appears to have increased total productivity (both above- and belowground), and reduced the proportion of C allocated belowground in mixtures compared to E. globulus monocultures. In a pot trial containing mixtures of E. globulus and A. mearnsii both species grew larger in mixture than in monoculture at low N levels, and mixtures were more productive than monocultures. However, at high N levels, E. globulus suppressed A. mearnsii and mixtures were less productive than E. globulus monocultures. Similar effects were found for high and low levels of P. Therefore resource availability can have a strong influence on the interactions and growth of mixtures. The productivity of mixtures may only be increased on sites where the resource for which competition is reduced in mixture is a major limiting growth resource. For example, if N is not a limiting growth factor then an increase in N availability from N-fixation may not increase growth, and the N-fixing species may compete for other resources such as soil P, moisture or light. This study has shown that mixtures containing a N-fixing trees and a non-N-fixing trees can be more productive than monocultures, but that this increase in productivity will only occur on certain sites. Examination of the growth, interactions and processes that occurred in mixtures in this study provide useful information that can aid the selection of species combinations and sites.

Abstract Plant community composition and diversity are determined by the balance between rates of immigration and extinction. Processes of immigration to a local community, i.e. propagule availability and dispersal of propagules between and within habitats, set the upper limit for the pool of species potentially capable of coexisting in a community, while local biotic interactions, i.e., competition, facilitation, herbivory and interactions with below-ground ecosystem components, and environmental factors control colonisation and establishment, and determine the persistence and dynamics of already existing species. In this thesis, I studied (1) the interactions between propagule availability, biotic and environmental constraints on colonisation, and (2) the interdependence between biotic and environmental factors regulating community processes in already established resident vegetation. First, I found that both propagule availability and competition with adult plants limited the rates of colonisation and total community diversity in a relatively low-productive tundra ecosystem. Long-term exclusion of mammalian herbivores and alleviation of nutrient limitation by fertilization increased the intensity of competition with established vegetation, and diminished immigration rates. In addition, I also found that community openness to colonization depended on the initial community properties, i.e., the functional composition and the traits of dominant plants in resident vegetation, which mediate the effects of nutrient addition and biomass removal on immigration rates. Second, adult plants in the resident vegetation experienced an increased extent of neighbourhood competition and herbivory in nutrient enriched conditions and in naturally more fertile habitats. However, the effects were also species-specific. On a community level, release from heavy grazing favoured lichens over graminoids and increased species richness. Furthermore, I also showed that plant community composition was strongly linked with soil organic matter quality and microbial community composition, and that these vegetation-soil-microbe interactions varied along a gradient of soil pH. Overall, my results emphasise that propagule availability, biotic and environmental control over community processes are strongly interconnected in tundra ecosystems. Especially, my findings highlight the role of plant competition and herbivory and their dependence on soil nutrient availability in governing colonisation and resident community dynamics. My results also indicate that plant functional composition and traits of dominant plants are of great importance in channelling community responses to external alterations and dictating plant-soil interactions.

The global effort to rehabilitate and restore destroyed mangrove forests is unable to keep up with the high mangrove deforestation rates which exceed the average pace of global deforestation by three to five times. Our knowledge of the underlying processes of mangrove forest regeneration is too limited in order to find suitable techniques for the restoration of degraded mangrove areas. The general objective of my dissertation was to improve mangrove restoration by understanding regeneration processes and local plant-plant interaction in a regenerating Avicennia germinans forest. The study was conducted in a high-shore mangrove forest area on the Ajuruteua peninsula, State of Para, Northern Brazil. The dwarf forest consisting of shrub-like trees is recovering from a stand-replacing event caused by a road construction in 1974 which interrupted the tidal inundation of the study area. Consequently, infrequent inundation and high porewater salinity limit tree growth and canopy closure. All trees and seedlings were stem-mapped in six 20 m x 20 m plots which were located along a tree density gradient. Moreover, height, crown extent, basal stem diameter of trees were measured. The area of herbaceous ground vegetation and wood debris were mapped as well. The mapped spatial distribution of trees, seedlings and covariates was studied using point pattern analysis and point process models, such as Gibbs and Thomas point process, in order to infer underlying ecological processes, such as seed dispersal, seedling establishment, tree recruitment and tree interaction. In the first study (chapter 2), I analyzed the influence of abiotic and biotic factors on the seedling establishment and tree recruitment of A. germinans during the recolonization of severely degraded mangrove sites using point process modeling. Most seedlings established adjacent to adult trees especially under their crown cover. Moreover, seedling density was higher within patches of the herbaceous salt-marsh plants Blutaparon portulacoides and Sesuvium portulacastrum than in uncovered areas. The higher density of recruited A. germinans trees in herb patches indicated that ground vegetation did not negatively influence tree development of A. germinans. In addition, tree recruitment occurred in clusters. Coarse wood debris had no apparent effect on either life stage. These results confirm that salt-marsh vegetation acts as the starting point for mangrove recolonization and indicate that the positive interaction among trees accelerates forest regeneration. In the second study (chapter 3), I analyzed how intraspecific interaction among A. germinans trees determines their growth and size under harsh environmental conditions. Interaction among a higher number of neighboring trees was positively related to the development of a focal tree. However, tree height, internode length and basal stem diameter were only positively associated in low-density forest stands (1.2 trees m-2) and not in forest stands of higher tree density (2.7 trees m-2). These results indicated a shift from facilitation, i.e. a positive effect of tree interaction, towards a balance between facilitation and competition. In the third study (chapter 4), I used point process modeling and the individual-based model mesoFON to disentangle the impact of regeneration and interaction processes on the spatial distribution of seedlings and trees. In this infrequently inundated area, propagules of A. germinans are only dispersed at a maximum distance of 3 m from their parent tree. Furthermore, there is no evidence that the following seedling establishment is influenced by trees. I was able to differentiate positive and negative tree interactions simulated by the mangrove model mesoFON regardless of dispersal processes based on static tree size information using the mark-correlation function. The results of this dissertation suggest that mangrove forest regeneration in degraded areas is a result of facilitative and not competitive interactions among mangrove trees, seedling and herbaceous vegetation. This has important implications for the restoration of degraded mangrove forest. Degraded mangrove areas are usually restored by planting a high number of evenly spaced seedlings. However, high costs constrain this approach to small areas. Assisting natural regeneration could be a less costly alternative. Herbaceous vegetation plays a crucial role in forest recolonization by entrapping propagules and possibly ameliorating harsh environmental conditions. So far only competition among mangrove trees has been considered during restoration. However, facilitative tree interactions could be utilized by planting seedling clusters in order to assist natural regeneration instead of planting seedlings evenly-spaced over large areas. This dissertation also showed that point pattern analysis and point process modeling can enable forest ecologists to describe the spatial distribution of trees as well as to infer underlying ecological processes.

Les plantes en coussins sont connues pour leur capacité fondatrice de nouvel habitat pour les autres espèces. Par ailleurs, des études ont montré que la variation morphologique au sein d’espèces fondatrices peut induire une variation des effets compétiteurs sur les espèces subordonnées, ce qui peut affecter la composition des communautés. Cette thèse a pour objectif d’étudier les conséquences d’une variation intraspécifique de deux espèces fondatrices (Festuca gautieri et Onobrychis cornuta) occupant des micro-environnements contrastés dans deux systèmes montagneux [les Pyrénées (France) et le Mont-Liban (Liban),respectivement] sur les communautés végétales associées (subalpines etoroméditerranéennes, respectivement). Nous avons évalué par des approches descriptives et/ou expérimentales (jardins expérimentaux, transplantations réciproques) les contributions de la génétique et de la plasticité à la variation morphologique entre deux phénotypes de coussins (dense et lâche) pour chaque espèce fondatrice, et à leurs effets contrastés sur les espèces subordonnées. Nous avons également quantifié les effets rétroactifs de la communauté pour la reproduction des espèces fondatrices. Nos résultats montrent une contribution à la fois de la génétique et de la plasticité à la variation phénotypique. La base génétique des variations morphologiques entre les phénotypes a induit des différences héréditaires d’effets compétiteurs sur les espèces subordonnées, tout en contrecarrant l’augmentation de la compétition avec la diminution du stress – le résultat dominant dans la littérature sur les systèmes subalpins. Nous avons aussi trouvé des effets rétroactifs négatifs des espèces subordonnées pour l’espèce fondatrice, avec une diminution de la production de fleurs (par les coussins) due au nombre croissant d’espèces subordonnées. La diversité des espèces subordonnées était plus élevée dans les conditions environnementales favorables que dans les stressantes. Par conséquent, nous avons conclu que les effets génétiques surmontent les effets environnementaux, limitant la compétition dans les milieux favorables, maintenant ainsi une plus grande diversité dans ces milieux que dans les milieux stressants
Alpine cushion plants are foundation species known for their nursingability. Moreover, studies have shown that morphological variation in foundationspecies can trigger variation in competitive effects on subordinate species, likely toaffect community composition. We investigated the consequences of intraspecificvariation within two alpine cushion species (Festuca gautieri and Onobrychis cornuta)across heterogeneous environments in two mountain ranges [the Pyrenees (France)and Mount-Lebanon (Lebanon), respectively] for the associated plant communities(subalpine and oromediterranean, respectively). We assessed with observationaland/or experimental (common-gardens, reciprocal transplantation experiments)approaches the relative contribution of genetics and plasticity to the morphologicalvariation between two cushion phenotypes (tight and loose) of the foundationspecies, and to their differential effects on subordinate species. Communityfeedbacks were also quantified. Our results show that both genetics and plasticitycontributed to the phenotypic variation. The genetic basis of the morphologicaldifferences between phenotypes induced heritable differences in competitive effectson subordinate species, but however counteracted the general increase incompetition with decreasing stress dominantly found in the literature on subalpinesystems. We also found negative feedbacks of subordinates on foundation speciesfitness, with higher cover of subordinate species reducing the cushions flowerproduction. Subordinate species diversity was higher in benign than in stressedenvironmental conditions. Consequently, we concluded that genetic effects overcomethe environmental effects by limiting competition in benign physical conditions, thusmaintaining a higher diversity in benign than stressed conditions

Lors de ce travail de thèse nous avons adopté une approche fonctionnelle de la dynamique forestière méditerranéenne. L'objectif principal est de comprendre comment les réponses fonctionnelles à la disponibilité de la lumière et aux interactions biotiques vont conditionner la régénération des trois principales espèces impliquées dans cette dynamique : Pinus halepensis, Quercus ilex, Quercus pubescens. Une première expérimentation en pots a été menée en conditions contrôlées afin de déterminer progressivement : les différences de stratégies d'acclimations à l'ombre du pin d'Alep et du chêne pubescent, les compromis fonctionnels et les limites ontogéniques et nutritionnelles de ces stratégies, et l'impact de l'allélopathie et de la compétition sur cette réponse à l'ombre. Une expérimentation de terrain a ensuite permis de créer un gradient d'éclaircie dans une pinède de pin d'Alep, pour comprendre les principaux effets positifs et négatifs de la canopée sur les facteurs abiotiques et le développement du chêne pubescent et du chêne vert à différentes échelles de temps. Les résultats obtenus montrent que le pin d'Alep présente une stratégie d'évitement de l'ombre par une forte réponse plastique qui implique des compromis fonctionnels, des coûts nutritionnels élevés, et un temps d'expression limité. Cette stratégie présente un désavantage dans le cas d'une régénération sous canopée. A l'inverse, le chêne pubescent montre une stratégie plus conservatrice grâce à une réponse plastique à l'ombre limitée, mais une forte sensibilité aux interactions plantes-plantes. En conditions naturelles, le chêne pubescent et le chêne vert en régénération présentent des réponses différentes au gradient d'éclaircie de la canopée, le chêne pubescent montrant cette fois ci une moins forte tolérance à l'ombre. Cette thèse permet de mettre en évidence les liens entre limites fonctionnelles de la plasticité phénotypique, tolérance à l'ombre, et réussite d'installation sous couvert forestier
During this work we studied the Mediterranean forest dynamic through a functional approach. Our main goal consisted to know how functional responses to light availability and biotic interactions determine the regeneration of three main species involved in this dynamic : Pinus halepensis, Quercus ilex and Quercus pubescens.A nursery experiment was set up to determine specific shade acclimatation strategies of the Aleppo pine and the Downy oak, functional tradeoffs and nutritional and ontogenetic limits of these stratgies, and the competitive and allélopathic impact on the shade induced response.Then a field experiment was set up to make a light gradient in an Aleppo pins pinewood in order to know the main positive and negative canopy effects on abiotic factors and oak seedlings development at different scales of time. Results showed that Aleppo pine exhibits a shade avoiding strategy through a strong plastic response involving functional tradeoffs, high nutritional costs, and time-limited expression. This strategy is considered as maladaptive in case of regeneration under unovercome forest canopy. Reversly, the downy oak shows a more conservative strategy through a limited plastic response to shade but a strong vulnerability to plant-plant interactions. In field conditions, Quercus pubescens and Quercus ilex seedlings showed different responses to the canopy cover, with a fewer shade tolerance in Quercus pubescens. This work help to highlight relationships between functional limits of phenotypic plasticity, shade tolerance, and regenerative success in Aleppo pine understory

This thesis conducted a detailed examination of debilitative and facilitative dimensions of competitive state anxiety. Competitive anxiety was assessed using the Competitive State Anxiety Inventory-2 (CSAI-2) which was modified to measure not only the 'intensity' (level) but also 'direction' (facilitative/debilitative) of cognitive anxiety, somatic anxiety and self-confidence. A trait version of the questionnaire was also used. Three different research methodologies were employed in this thesis. The first two studies adopted a quantitative research methodology, Study 3 incorporated qualitative techniques and the final investigation addressed the research question via a single-subject design study. The first study investigated intensity and direction dimensions of state anxiety as a function of skill level in a sample of elite and non-elite swimmers. The results revealed that while no differences were evident between the groups on the intensity of cognitive and somatic anxiety, the elite group reported more facilitative interpretations of these symptoms than the non-elite group. Self-confidence was also higher in the elite group. The results suggested elite performers do not differ from nonelite performers on the level of anxiety they experience, but they do have a more positive interpretation of these symptoms in terms of consequence for performance. The second study examined the predictions of a control process model of anxiety proposed by Jones (1995a). Specifically, the study examined the directional perceptions of anxiety as a function of goal attainment expectations. Results showed that swimmers with favourable expectancies of their ability to achieve their goals reported no differences in intensity levels to the swimmers with negative or uncertain expectancies of goal achievement. However, more facilitative interpretations of pre-race symptoms were reported by the positive goal expectancy group. These findings highlighted important applied implications for achieving appropriate pre-performance states via setting appropriate goals that are within the control of the performer. Study three addressed how elite performers have acquired the ability to interpret their anxiety symptoms as being facilitative towards upcoming performance. Qualitative interview techniques and inductive content analysis revealed that the performers did experience negative cognitive and somatic anxiety symptoms when they first started competing. However, by appropriate education and natural learning experiences, the performers became aware that the nerves they experienced could be positive towards the upcoming race. Furthermore, the swimmers developed, and now follow, detailed pre-competition and pre-race routines to maintain this facilitative interpretation. The findings generated important practical implications for attempting to restructure negative interpretations of anxiety symptoms. The final study examined the effects of a multimodal intervention programme on performers debilitated by their anxiety symptoms, via a staggered multiple-baseline single-subject design over a series of ten competitive races. Following the intervention, the results showed that although intensity levels remained stable, the three performers receiving the intervention reported more facilitative interpretations of both cognitive and somatic anxiety. Self-confidence also increased following the intervention treatment. These findings demonstrated that information generated by elite performers can be transferred successfully in ecologically valid sport environments. The programme of research conducted in this thesis provides evidence that the conventional notion of anxiety as negative towards performance can be questioned and reconceptualised to include positive consequences for performance. The results highlight the importance of future research into other emotions that may have been formerly labelled as anxiety, but which denote a different emotional state altogether.

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Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES
The relationships between organisms and the mechanisms that enable their coexistence are important issues in ecology. Plant removal experiments are useful to elucidate these mechanisms because its cause and effect response. This thesis aims to analyze experiments removal plants. The first paper presents a scientometric review of the literature on plant functional groups removal 1991. We verify aspects such as the number of publications and citations in journals indexed in the Web of Science, which country was made in order to detect bias and geographic gaps, which ecological effects were tested, which functional groups were removed, which removal methods and which environments were made. We found an increase in the number of publications and citations on average during this period, however, was 12 times lower than publications on functional diversity, perhaps due to the difficulty of conducting experiments. The work was mainly carried out in temperate climates where taxonomic knowledge is greater, in addition, there is a prevalence of vegetation that is most commonly removed than trees. Testing interactions, especially competition, was the main reason for removal, perhaps as a matter of historical competition is considered the interaction that influences the co-occurrence of species. The cut at ground was the major method of removing, undisturbing the soil and avoiding undesirable effects of herbicides, and few studies have been made in a greenhouse and in the natural environment at the same time, which could elucidate responses to natural conditions and at the same time responses specific interactions. In the second paper we carried a case study on removal of capim-flecha ( Tristachya leiostachya Ness.), a dominant grass in Emas National Park (ENP), and what the consequences of such removal to the co-occurring grasses. We expected that the absence of capim-flecha allow further growth of grasses, however we found no difference in mean values between treatments indicating that there is no competition influencing the abundance of capim-flecha. The historical context burned in PNE may have influenced this high dominance of capim-flecha. We believe that experiments are useful in testing interactions, especially if they are made in the natural environment and greenhouse at the same time.
As relações entre os organismos e os mecanismos que permitem sua coexistência são importantes questões em ecologia. Experimentos de remoção de plantas são úteis em elucidar esses mecanismos, devido sua capacidade de resposta do tipo causa e efeito. Esta dissertação se propõe a analisar experimentos de remoção de plantas. O primeiro artigo apresenta uma revisão cienciométrica das publicações sobre remoção de grupos funcionais de plantas, desde 1991. Verificamos aspectos como o número de publicações e citações em periódicos indexados no Web of Science, em qual país foi feito, visando detectar vieses e lacunas geográficas, quais efeitos ecológicos testados, quais grupos funcionais removidos, quais os métodos de remoção e em quais ambientes foram feitos. Encontramos um crescimento no número de publicações bem como na média de citações nesse período, contudo foi 12 vezes menor do que publicações sobre diversidade funcional, talvez devido à dificuldade de realizar experimentos. Os trabalhos foram realizados principalmente em climas temperados onde o conhecimento taxonômico é maior, além disso, há um predomínio de vegetação rasteira que é mais comumente removida do que árvores. Testar interações, principalmente competição, foi o principal motivo de remoção, talvez por uma questão histórica de competição ser considerada a interação que mais influencia a co-ocorrência de espécies. O corte rente ao solo foi o principal método de remoção, evitando revolver o solo e efeitos indesejáveis de herbicidas, e poucos trabalhos foram feitos em casa de vegetação e ambiente natural ao mesmo tempo, o que poderia elucidar respostas à condições naturais e ao mesmo tempo respostas específicas de interações. No segundo capítulo fizemos um estudo de caso sobre remoção de capim-flecha (Tristachya leiostachya Ness.), uma gramínea dominante no Parque Nacional das Emas (PNE), e quais as consequências dessa remoção para as gramíneas co-ocorrentes. Esperávamos que a ausência do capim-flecha permitiria um maior crescimento das gramíneas, contudo não encontramos diferença nos valores das médias entre os tratamentos o que indica que não há competição influenciando a abundância do capim-flecha. O contexto histórico queimadas no PNE pode ter influenciado essa alta dominância do capim-flecha. Consideramos que experimentos são úteis em testar interações, sobretudo se forem feitos em ambiente natural e casa de vegetação ao mesmo tempo.

Les recherches récapitulées dans cette thèse de doctorat ont porté sur les causes de l’organisation spatiale des végétations périodiques. Ces structures paysagères aux motifs réguliers, tachetés, tigrés ou labyrinthiques, d’échelle décamétrique à hectométrique, couvrant des étendues considérables sur au moins trois continents, constituent un cas d’école dans l’étude des processus endogènes présidant à l’hétérogénéité du couvert végétal. Ces structures prennent place sur un substrat homogène, mis à part la rétroaction du couvert lui-même, et sont marquées par des écotones abrupts et la persistance d’une proportion considérable de sol nu. Plusieurs modèles ont mis en avant l’existence possible d’un phénomène d’auto-organisation du couvert, qui verrait une structure d’ensemble émerger des interactions locales entre individus. Ces modèles se basent sur le jeu simultané de la consommation de la ressource (compétition) et de l’amélioration de l’un ou l’autre des éléments du bilan de la même ressource par le couvert (facilitation). La condition à l’existence d’une structure d’ensemble spatialement périodique et stable réside dans une différence entre la portée de la compétition (plus grande) et celle de la facilitation. L’apparition de ces structures est modulée par le taux de croissance biologique, qui est le reflet des contraintes extérieures telles que l’aridité, le pâturage ou la coupe de bois. Le modus operandi des interactions spatiales supposées entre individus reste largement à préciser.

Nos recherches ont été menées au sud-ouest de la République du Niger, à l’intérieur et dans les environs du parc Régional du W. Trois axes ont été explorés :(i) Une étude de la dépendance spatiale entre la structure de la végétation (biovolumes cartographiés) et les paramètres du milieu abiotique (relief, sol), sur base d’analyses spectrales et cross-spectrales par transformée de Fourier (1D et 2D). (ii) Une étude diachronique (1956, 1975 et 1996) à large échelle (3000 km²) de l’influence de l’aridité et des pressions d’origine anthropique sur l’auto-organisation des végétations périodiques, basée sur la caractérisation de la structure spatiale des paysages sur photos aériennes via la transformée de Fourier en 2D. (iii) Trois études portant sur les interactions spatiales entre individus :En premier lieu, via l’excavation des systèmes racinaires (air pulsé) ;Ensuite, par un suivi spatio-temporel du bilan hydrique du sol (blocs de gypse) ;Enfin, via le marquage de la ressource par du deutérium.

Nous avons ainsi pu établir que les végétations périodiques constituent bien un mode d’auto-organisation pouvant survenir sur substrat homogène et modulé par les contraintes climatiques et anthropiques. Un ajustement rapide entre l’organisation des végétations périodiques et le climat a pu être montrée en zone protégée. La superficie et l’organisation des végétations périodiques y ont tour à tour progressé et régressé en fonction d’épisodes secs ou humides. Par contre, en dehors de l’aire protégée, la possibilité d’une restauration du couvert semble fortement liée au taux d’exploitation des ressources végétales. Ces résultats ont d’importantes implications quant à la compréhension des interactions entre climat et écosystèmes et à l’évaluation de leurs capacités de charge. La caractérisation de la structure spatiale des végétations arides, notamment par la transformée de Fourier d’images HR, devrait être généralisée comme outil de monitoring de l’état de ces écosystèmes. Nos études portant sur les modes d’interactions spatiales ont permis de confirmer l’existence d’une facilitation à courte portée du couvert végétal sur la ressource. Cependant, cette facilitation ne semble pas s’exercer sur le terme du bilan hydrique traditionnellement avancé, à savoir l’infiltration, mais plutôt sur le taux d’évaporation (deux fois moindre à l’ombre des canopées). Ce mécanisme exclut l’existence de transferts diffusifs souterrains entre sols nu et fourrés. Des transferts inverses semblent d’ailleurs montrés par le marquage isotopique. L’étude du bilan hydrique et la cartographie du micro-relief, ainsi que la profondeur fortement réduite de la zone d’exploitation racinaire, jettent de sérieux doutes quant au rôle communément admis des transferts d’eau par ruissellement/diffusion de surface en tant que processus clé dans la compétition à distance entre les plantes. L’alternative réside dans l’existence d’une compétition racinaire de portée supérieure aux canopées. Cette hypothèse trouve une confirmation tant par les rhizosphères excavées, superficielles et étendues, que dans le marquage isotopique, montrant des contaminations d’arbustes situés à plus de 15 m de la zone d’apport. De même, l’étude du bilan hydrique met en évidence les influences simultanées et contradictoires (facilitation/compétition) des ligneux sur l’évapotranspiration.

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This PhD thesis gathers results of a research dealing with the causes of the spatial organisation of periodic vegetations. These landscape structures, featuring regular spotted, labyrinthine or banded patterns of decametric to hectometric scale, and extending over considerable areas on at least three continents, constitute a perfect study case to approach endogenous processes leading to vegetation heterogeneities. These patterns occur over homogeneous substratum, except for vegetation’s own feedbacks, and are marked by sharp ecotones and the persistence of a considerable amount of bare soil. A number of models suggested a possible case of self-organized patterning, in which the general structure would emerge from local interactions between individuals. Those models rest on the interplay of competitive and facilitative effects, relating to soil water consumption and to soil water budget enhancement by vegetation. A general necessary condition for pattern formation to occur is that negative interactions (competition) have a larger range than positive interactions (facilitation). Moreover, all models agree with the idea that patterning occurs when vegetation growth decreases, for instance as a result of reduced water availability, domestic grazing or wood cutting, therefore viewing patterns as a self-organised response to environmental constraints. However the modus operandi of the spatial interactions between individual plants remains largely to be specified.

We carried out a field research in South-West Niger, within and around the W Regional Park. Three research lines were explored: (i) The study of the spatial dependency between the vegetation pattern (mapped biovolumes) and the factors of the abiotic environment (soil, relief), on the basis of spectral and cross-spectral analyses with Fourier transform (1D and 2D). (ii) A broad scale diachronic study (1956, 1975, 1996) of the influence of aridity and human induced pressures on the vegetation self-patterning, based on the characterisation of patterns on high resolution remote sensing data via 2D Fourier transform. (iii) Three different approaches of the spatial interactions between individuals: via root systems excavation with pulsed air; via the monitoring in space and time of the soil water budget (gypsum blocks method); and via water resource labelling with deuterated water.

We could establish that periodic vegetations are indeed the result of a self-organisation process, occurring in homogeneous substratum conditions and modulated by climate and human constraints. A rapid adjustment between vegetation patterning and climate could be observed in protected zones. The area and patterning of the periodic vegetations successively progressed and regressed, following drier or wetter climate conditions. On the other hand, outside protected areas, the restoration ability of vegetation appeared to depend on the degree of vegetation resource exploitation. These results have important implications regarding the study of vegetation-climate interactions and the evaluation of ecosystems’ carrying capacities. Spatial pattern characterisation in arid vegetations using Fourier transform of HR remote sensing data should be generalised for the monitoring of those ecosystems. Our studies dealing with spatial interaction mechanisms confirmed the existence of a short range facilitation of the cover on water resource. However, this facilitation does not seem to act through the commonly accepted infiltration component, but rather on the evaporative rate (twice less within thickets). This mechanism excludes underground diffusive transfers between bare ground and vegetation. Inverse transfers were even shown by deuterium labelling. Water budget study and micro-elevation mapping, along with consistent soil shallowness, together cast serious doubts on the traditional mechanism of run-off/diffusion of surface water as a key process of the long range competition between plants. An alternative explanation lies in long range root competition. This hypothesis find support as well in the excavated root systems, shallow and wide, as in isotopic labelling, showing contaminations of shrubs located up to 15 m of the irrigated area. Water budget study also evidenced simultaneous contradictory effects (facilitation/competition) of shrubs on evapotranspiration.

Doctorat en sciences agronomiques et ingénierie biologique
info:eu-repo/semantics/nonPublished

Les xarxes complexes serveixen als ec olegs per a estudiar les interaccions de pollinitzaci o a nivell comunitari. Actualment, es necessari construir xarxes que siguin representacions m es realistes del proc es de pol.linitzaci o per tal d'incrementar la seva utilitat pr actica. Per aix o, aquesta tesi es centra en incorporar a les xarxes la variaci o interindividual i informaci o sobre la magnitud i el signe (positiu, neutre o negatiu) dels efectes de les interaccions sobre la reproducci o de les plantes. Les esp ecies consten de poblacions d'individus amb fenotips, genotips i comportaments diferents, que per tant poden diferir en les interaccions. Aquesta variaci o intraespec ca es rellevant per molts processos ecol ogics i evolutius, per o no s'ha tengut en compte en la majoria d'estudis de xarxes planta-pol.linitzador. En els cap tols 1 i 2 s'estudien les c arregues pol.l niques d'exemplars d'insectes pol.linitzadors en dues comunitats de muntanya a Mallorca i es construeixen xarxes on els nodes dels pol.linitzadors representen individus en lloc d'esp ecies. Aquesta aproximaci o permet considerar la variaci o interindividual i aporta una nova perspectiva sobre l'estructura de les xarxes i els mecanismes que determinen les interaccions. Les xarxes de transport de pol.len a nivell d'individu tenen una densitat d'interaccions, connect ancia, grau d'anidament i diversitat d'interaccions m es baixa que les mateixes xarxes a nivell d'esp ecies, i per contra una modularitat major. Es troben aquestes difer encies perqu e les esp ecies de pol.linitzadors generalistes estan formades per individus especialistes i heterogenis en l' us de recursos. El grau d'especialitzaci o individual est a associat a la intensitat de compet encia inter- i intraespec ca. Per a que les xarxes planta-pol.linitzador representin millor les implicacions funcionals, han d'incloure mesures dels efectes de les interaccions sobre les esp ecies o de la magnitud real en qu e les esp ecies depenen d'aquestes interaccions. La freq u encia d'interacci o es considera una estimaci o v alida de la magnitud de l'efecte del pol.linitzador sobre l' exit reproductiu de la planta, per o no aporta cap informaci o del seu signe. Per exemple, en plantes amb mecanismes de producci o de llavors que no depenen d'insectes (e.g. autog amia, anemog amia), les interaccions poden tenir efectes neutres sobre la reproducci o. En el cap tol 3, per diferents plantes de dues comunitats d'estudi (costa i muntanya) es quanti ca el seu grau de depend encia dels pol.linitzadors, comparant experimentalment la producci o de llavors amb i sense insectes. L'objectiu es determinar si les esp ecies que s on m es depenents dels pol.linitzadors s on tamb e les que estan m es connectades en les xarxes de visites orals, es a dir si tenen major centralitat, nombre i diversitat d'interaccions. Nom es en una de les dues comunitats estudiades es troba aquesta relaci o, fet que suggereix que pot dependre del contexte comunitari. D'altra banda, hi ha interaccions que poden causar m es efectes negatius que positius sobre l' exit reproductiu de les plantes. Aix o pot passar quan els pol.linitzadors depositen pol.len conespec c en els estigmes, per o alhora tamb e pol.len heterospec c. La transfer encia interespec ca de pol.len es relativament comuna, perqu e sovint les esp ecies de oraci o simult ania comparteixen pol.linitzadors, i pot tenir efectes perjudicials per les plantes (e.g. p erdua de pol.len, obstrucci o dels estigmes). En el cap tol 4, s'estudien les transfer encies de pol.len interespec ques en tres comunitats andines al llarg d'un gradient altitudinal. Es construeixen xarxes dirigides representant la transfer encia de pol.len des de les esp ecies donants a les receptores i a cada interacci o de la xarxa s'hi associa un signe per representar l'efecte. Aquest signe s'obt e de la relaci o entre el pol.len conespec c i heteroespec c depositat sobre els estigmes. En totes les comunitats estudiades, les interaccions positives i neutres s on predominants, particularment en la comunitat de major altitud. Aquesta troballa suggereix que la facilitaci o entre plantes d'una comunitat pot augmentar quan les condicions pel servei de pol.linitzaci o es tornen menys favorables.
Las redes complejas sirven a los ec ologos para estudiar las interacciones de polinizaci on a nivel comunitario. Actualmente, para incrementar su utilidad pr actica, es necesario construir redes que sean representaciones lo m as realistas posibles del proceso de polinizaci on. Por este motivo, esta tesis se centra en incorporar a las redes la variaci on interindividual y informaci on sobre la magnitud y el signo (positivo, neutro o negativo) de los efectos de las interacciones sobre la reproducci on de las plantas. Las especies constan de poblaciones de individuos con fenotipos, genotipos y comportamientos distintos, que por tanto pueden diferir en sus interacciones. Esta variaci on intraespec ca es relevante para muchos procesos ecol ogicos y evolutivos, pero no se ha considerado en la mayor a de estudios de redes plantapolinizador. En los cap tulos 1 y 2 se estudian las cargas pol nicas de ejemplares de insectos polinizadores en dos comunidades de monta~na en Mallorca y se construyen redes donde los nodos de los polinizadores representan individuos en lugar de especies. Esto permite incorporar la variaci on interindividual y aportar una nueva perspectiva sobre la estructura de las redes y los mecanismos que determinan las interacciones. Las redes de transporte de polen a nivel de individuo tienen una densidad de interacciones, conectancia, anidamiento y diversidad de interacciones m as baja que las mismas redes a nivel de especies, pero una modularidad mayor. Estas diferencias ocurren porque las especies de polinizadores generalistas est an formadas por individuos especialistas y heterog eneos en la utilizaci on de recursos. El grado de especializaci on individual est a asociado a la intensidad de competencia inter- e intraespec ca. Para que las redes planta-polinizador sean m as representativas de las implicaciones funcionales deben incluir medidas de los efectos de las interacciones sobre las especies o de la magnitud real en que las especies dependen de estas interacciones. La frecuencia de interacci on se considera una estimaci on v alida del efecto del polinizador sobre el exito reproductivo de la planta, aunque no aporta informaci on sobre su signo. Por ejemplo, en plantas con mecanismos de producci on de semillas independientes de los insectos (e.g. autogamia, anemogamia), las interacciones pueden tener efectos neutros sobre la reproducci on. En el cap tulo 3, para diferentes plantas de dos comunidades de estudio (costa y monta~na) se cuanti ca el grado de dependencia de polinizadores comparando experimentalmente la producci on de semillas con y sin insectos. El objetivo es determinar si las especies m as dependientes de polinizadores son tambi en las m as conectadas en las redes, con mayor centralidad, n umero y diversidad de interacciones. S olo en una de las comunidades de estudio se encuentra esta relaci on, sugiriendo que es variable en funci on del contexto comunitario. Por otro lado, hay interacciones que pueden causar efectos m as negativos que positivos sobre el exito reproductivo de las plantas. Esto puede pasar cuando los polinizadores depositan polen conespec co y heterospec co en los estigmas. La transferencia interespec ca de polen es relativamente com un, porque a menudo las especies de oraci on simult anea comparten polinizadores, y puede tener efectos perjudiciales para las plantas (e.g. p erdida de polen, obstrucci on de estigmas). En el cap tulo 4, se estudian las transferencias de polen interespec cas en tres comunidades andinas a lo largo de un gradiente altitudinal. Se construyen redes dirigidas que representan la transferencia de polen desde las especies donantes a las receptoras y a cada interacci on se le asocia un signo mostrando el efecto. Este signo se obtiene de la relaci on entre el polen conespec co y heteroespec co depositado sobre los estigmas. En todas las comunidades estudiadas, predominan las interacciones positivas y neutras, particularmente en la comunidad de mayor altitud. Esto sugiere que la facilitaci on en una comunidad puede aumentar cuando las condiciones para la polinizaci on se vuelven menos favorables.
Ecologists use network analysis to study pollination interactions at a communitywide level. The construction of plant-pollinator networks which are realistic representations of the pollination process is fundamental to increase their usefulness and ecological meaning. For that reason, this thesis focuses on incorporating to such networks individual variation and information about the magnitude and sign (positive, neutral or negative) of interaction e ects on plant reproduction. Species consist of populations of phenotypically, genetically and behaviourally diverse individuals which thus di er in their interactions and foraging decisions. Despite its relevance for many ecological and evolutionary processes, intraspeci c variation has been overlooked in most plant-pollinator network studies. In chapters 1 and 2, pollen loads of insect pollinator individuals are studied in two mountain communities of Mallorca, and networks where pollinator nodes depict individuals instead of species are built. Such approach (network downscaling) permits to account for the variation within species and provides new insights on network interaction patterns and their causal mechanisms. Pollen-transport networks at the individual level had lower linkage density, connectance, nestedness and interaction diversity, than the same networks at the species level, but higher modularity. These di erences occur because generalist pollinator species are composed of heterogeneous specialist individuals in the use of resources. The degree of individual specialization is associated with inter- and intraspeci c competition. Plant-pollinator networks which represent more accurately functional implications should include measures of interaction e ects on species or estimates of the actual degree to which species depend on such interactions. In general, interaction frequency is considered a good surrogate of the magnitude of interaction e ects, although it does not give any information about the e ect sign. For instance, in plant species with mechanisms for producing seeds independently of pollinators (e.g. autogamy, anemogamy), interactions may have a neutral reproductive e ect. In chapter 3, the degree of reproductive dependence on pollinators is quanti ed for several plant species in two study communities (coast and mountain), comparing seed set with and without insects. The objective is to determine whether plant species which depend more on pollinators are also more connected in ower-visitation networks, i.e. have high centrality, number and diversity of interactions. Such relationship is only found in one of the two study communities, which suggests that it is community-context dependent. On the other hand, there are interactions which may cause more negative than positive e ects on plant reproductive success. This may happen when owervisitors deposit both conspeci c and heterospeci c pollen on stigmas. Such interspeci c pollen transfer is common because co- owering plants often share pollinators, and can have detrimental e ects on plant tness (e.g. pollen loss and stigma clogging). In chapter 4, interspeci c pollen transfers are studied in three high-Andean communities along a mountain altitudinal gradient. Directed networks depicting pollen transfers from donor to receptor species are constructed, and a sign is associated to each network link to represent the e ect. This sign is obtained from the study of the relationship between conspeci c and heterospeci c pollen deposited on stigmas. In all study communities, facilitative and neutral pollinator-mediated interactions among plants prevail over competition, particularly in the highest elevation community. This nding suggests that pollination facilitation in communities can increase under less favourable conditions for the pollination service, supporting previous predictions of higher incidence of facilitative interactions in stressful environments.

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This study aims to examine some of the main respects the principle of facilitation of consumer protection in court, provided for by article 6, section VIII, the code of consumer protection, within the individual deal. Will address issues pertaining to competition, shifting the burden of proof, the authority of states obligations to do and not do, and the special civil courts, and how these institutions and instruments to give effect to act principle in question. The analysis takes place from the beginning governing the protective system to the consumer in the CDC, taking account of the imbalance inherent in the relationship of consumption, which stems from the situation of vulnerability of the consumer, and the difficulties for him found with regard to access to justice and their situation in judge
O presente trabalho tem por finalidade analisar alguns dos principais aspectos do princípio da facilitação da defesa do consumidor em juízo, previsto pelo art. 6º, inciso VIII, do Código de Defesa do Consumidor, no âmbito das lides individuais. Serão abordadas questões atinentes à competência, à inversão do ônus da prova, à tutela específica das obrigações de fazer e de não fazer, e aos Juizados Especiais Cíveis, e como estes institutos e instrumentos atuam para conferir efetividade ao princípio em questão. A análise desenvolve-se a partir dos princípios que regem o sistema protetivo ao consumidor, contido no CDC, levando-se em conta o desequilíbrio inerente às relações de consumo, que decorre da situação de vulnerabilidade do consumidor, e as dificuldades por ele encontradas no que respeita ao acesso à Justiça e à sua atuação em juízo

Les écosystèmes herbacés méditerranéens, caractérisés par une importante biodiversité, ont subi de considérables changements d‟usage. Désormais, leur conservation est devenue insuffisante au regard de l‟augmentation des dommages causés par les activités humaines. La restauration écologique apparaît donc comme un outil approprié pour participer à leur préservation. Des recherches préalables sur les différents compartiments de l‟écosystème ainsi que sur les facteurs responsables de l‟assemblage de leurs communautés avant et après dégradations sont donc nécessaires afin de mettre en place des méthodes de restauration adaptées à ces écosystèmes. Ainsi, en utilisant comme modèle biologique les Coussouls de Crau et leurs friches post-culturales (Bouches-du-Rhône, France), représentatif de nombreux écosystèmes steppiques méditerranéens, nous avons choisi de déterminer et d‟étudier les principaux filtres potentiellement responsables de l‟organisation de cette communauté végétale. Le pâturage ovin, les changements trophiques du sol, les phénomènes de compétition et la capacité de reproduction et/ou de dispersion des espèces, identifiés lors d‟une première étude sur la régénération de la steppe, ont ainsi été étudiés dans trois expérimentations différentes. Deux protocoles de restauration écologique in situ (transfert de foins et semis d‟espèces structurantes) et un protocole d‟expérimentation ex-situ testant les interactions d‟une sélection d‟espèces de la steppe ont permis de réaliser ces recherches. Nos principaux résultats montrent que les filtres de reproduction/dispersion, le niveau trophique du sol, la compétition et le régime de perturbations récurrentes lié au pâturage ovin ont un rôle déterminant dans les changements de la composition et de la structure de la communauté et ils expliquent ainsi en partie le ralentissement de sa dynamique successionnelle vers la steppe de référence. Nous avons également pu démontrer que ces filtres interagissaient entre eux. Au niveau de la recherche appliquée, les protocoles testés favorisent le retour de la végétation steppique, néanmoins la composition des parcelles restaurées reste toujours très différente de l‟écosystème de référence à très court terme (deux à trois années après la restauration). Cela démontre que même si nous comprenons mieux aujourd‟hui les facteurs expliquant l‟organisation de cette communauté, il est cependant toujours difficile de la restaurer dans son intégralité car il n‟est pas aisé de manipuler et de trouver un équilibre entre l‟action de ces différents facteurs. Il est donc désormais plus que nécessaire de poursuivre ces recherches fondamentales et appliquées notamment au niveau des processus de maturation et de structuration des vieilles communautés végétales herbacées
Mediterranean herbaceous ecosystems which are characterized by a high biodiversity have been submitted to a lot of changes in use. Their conservation is not sufficient enough and ecological restoration seems to be an adapted solution to improve conservation efforts as it can mitigate damages induced by human activities. To develop restoration methods adapted to these ecosystems, preliminary researches on the different communities composing an ecosystem and the filters which are responsible for community assemblage before and after disturbance are necessary. Representative of numerous Mediterranean steppe ecosystems, the “Coussouls de Crau” and their ex-arable-fields (Bouches-du-Rhône, France) are the biological models used to identify and study the principal filters potentially responsible for organizing the plant communities. Sheep grazing, trophic changes, competition and reproduction and /or species dispersal, were identified in a first study on steppe regeneration; these filters were then the core of the following three research experiments. We carried out two in-situ restoration protocols (hay transfer and sowing foundation species) and one ex-situ experiment testing interactions between a selection of steppe species. The main results show that reproduction, seed dispersal, soil trophic level, competition, extensive sheep grazing are filters which have a determinant role in structuring the communities, in determining plant composition and partly in explaining the slow dynamics towards the reference steppe. Also, we show that filters interact between them. For applied restoration, the tested methods promote steppe vegetation come back, but in the short term (two or three years after restoration), the composition of restored areas remains different to the reference ecosystem. Even if we understand better the different factors which explain community organization, these results demonstrate that it is difficult to restore it integrally because it is not easy to manipulate and find an equilibrium between the different filters actions. Now it is necessary to continue these fundamental and applied researches, in particular on the maturation and structure of the old herbaceous plant communities

En aquesta tesi presento quatre capítols, en els quals es discuteix com diferents espècies subarbustives interactuen amb plantes coexistents, sota règims variables d’estrès. Aquesta recerca ha estat centrada en ecosistemes de gran valor ambiental, ecològic i de conservació, i alhora sensibles als canvis ambientals, en quatre regions distintes situades en zones de latitud o altitud elevades. Per ordre latitudinal, els ecosistemes estudiats han estat: 1) el límit supraforestal dels Pirineus Centrals, situats en una zona temperada; 2) el gradient de successió primària d’un ecosistema situat en una zona boreal, a Finlàndia,; 3) el límit supraforestal situat a la zona subàrtica de Lapònia; i 4) la tundra situada en una zona de l’alt àrtic, al nord‐est de Grenlàndia . Els dos primers capítols es basen en una aproximació experimental i se centren en investigar com els subarbustos més comuns trobats prop del límit supraforestal interactuen amb plançons d’arbres de les espècies que formen el límit del bosc en dos ambients contrastats, i.e. el límit supraforestal subàrtic, i el límit supraforestal dels Pirineus Centrals. A més, donat que els arbres que viuen prop del límit de distribució són molt sensibles als canvis ambientals , especialment a l’augment de temperatures durant l’estació de creixement (Körner 2003), també s’ha investigat com podrien afectar el creixement dels plançons d’arbre i la seva supervivència al llarg de l’ecotò, diferents escenaris de canvis ambientals. El primer capítol es titula ‘Les interaccions arbre‐arbust i els canvis ambientals dirigeixen la dinàmica supraforestal a la zona subàrtica’, i s’hi exposa l’experiment de tres anys de durada que es va executar al ‘Parc Nacional d’Abisko’, en els Scandes subàrtics, al nord de Suècia; les plantes estudiades varen ser plançons de Betula pubescens i el subarbust Vaccinium myrtillus. En aquest estudi es va certificar la importància de les interaccions arbust‐arbre, tant facilitatives com competitives, com a elements clau en la dinàmica supraforestal subàrtica. A més, vàrem demostrar que la gran sensibilitat dels plançons a l’escalfament tenia fortes implicacions per la dinàmica supraforestal, tenint en compte l’escenari previst d’augment de temperatures en latituds elevades. També es va observar que les interaccions complexes entre arbusts i herbívors són claus per preveure canvis futurs. El segon capítol es titula ‘Els plançons d’arbres situats als límits supraforestals Pirinencs i subàrtics mostren respostes semblants a la presència d'arbustos i a les simulacions de canvis ambientals’. Aquí es presenta una comparació entre els resultats de l’experiment presentat en el primer capítol i els obtinguts en un experiment paral∙lel dut a terme durant un període de temps semblant prop del límit supraforestal en el ‘Parc Natural de l’Alt Pirineu’, als Pirineus Centrals catalans. Aquest experiment es va basar en el mateix disseny factorial que l’anterior estudi, però amb diferents espècies (i.e. plançons de l’arbre Pinus uncinata i l’arbust Rhododendron ferrugineum). Segons ens consta, és el primer estudi que avalua experimentalment les respostes de les plantes vers diferents escenaris ambientals en un ecosistema d’alta muntanya als Pirineus. En aquest capítol es presenten alguns mecanismes que ajudaran a comprendre la variabilitat recentment observada de les respostes locals de límits supraforestals de zones temperades i subàrtiques com a resultat del clima canviant, i també identifiquem alguns paral∙lelismes que poden utilitzar‐se per generalitzar les respostes a gran escala dels límits supraforestals al canvi climàtic. El tercer capítol se centra en els efectes d’un subarbust dominant (i.e. Empetrum nigrum) en plançons de Pinus sylvestris al llarg del gradient de successió primària en un ecosistema boreal en una illa emergent de la badia de Bòtnia, a Finlàndia. Aquest capítol s’anomena ‘Un arbust ericoide exerceix la doble funció de reclutar pins i els seus simbionts fúngics al llarg d’un gradient de successió primària’. Aquí hem mostrat que els efectes facilitadors i competidors dels subarbustos determinen fortament l’establiment de plançons i la seva colonització fúngica al llarg d’aquest gradient de successió. Segons ens consta, aquests són els primers resultats que demostren que un arbust ericoide micorrízic pot millorar tant el desenvolupament de l’arbre hoste ectomicorrízic com el dels simbionts fúngics de l’arbre. L’estudi presentat al quart capítol es va realitzar al llarg d’un gradient de nivositat en un ecosistema extrem de tundra àrtica al ‘Parc Nacional del nord‐est de Grenlàndia’, el Parc Nacional més gran del món. El capítol es titula ‘Interaccions vegetals i composició de la vegetació àrtica al llarg d’un gradient de nivositat al nord‐est de Grenlàndia’. Aquest ecosistema és probablement el més sensible i fràgil de tots els ecosistemes estudiats en aquesta tesi, donat que s’espera que a la costa est de Grenlàndia s’hi esdevinguin canvis substancials en el clima com a resultat de canvis destacables en els règims de precipitació de neu i de les temperatures (Brown i Mote 2009). Aquí es va avaluar la riquesa d’espècies de plantes, així com els patrons d’establiment i composició de diverses formes de creixement existents en comunitats vegetals àrtiques associades a una variació de la cobertura de neu durant els mesos d’hivern. Aquest estudi ajudarà a preveure la diversitat potencial i els canvis en la vegetació a la zona de l’alt Àrtic, si els règims de precipitació de neu canvien en el futur com es preveu.
In this thesis I present four chapters, and in all of them I discuss how dwarf shrubs interact with co‐occurring plants under varying regimes of stress. This research involved ecosystems of great environmental, nature conservation and ecological value, yet highly sensitive to environmental changes, in four contrasting cold regions at high altitude or high latitude. Following a latitudinal order, the selected ecosystems were: 1) a temperate alpine treeline in the Central Pyrenees; 2) a primary succession gradient in a boreal ecosystem in Finland; 3) a subarctic alpine treeline in Lapland; and 4) a high‐arctic tundra in north‐eastern Greenland. The first two chapters are based on an experimental approach and focus on how shrubs commonly found near the treeline interact with tree seedlings of treelineforming species in two contrasting environments, i.e. in a subarctic forest‐tundra ecotone in Lapland, northern Sweden, and in a more southern, temperate forestalpine pasture ecotone in the Central Pyrenees. In addition, since trees living near their limit of distribution are very sensitive to environmental changes, especially to increased temperature during the growing season (Körner 2003), we also assessed how distinct environmental change scenarios may affect tree seedling growth and survival across the ecotone. The first chapter is entitled ‘Shrub‐tree interactions and environmental changes drive treeline dynamics in the Subarctic’, where we explain the three‐year‐long experiment performed in the Abisko National Park, in the subarctic Scandes, Northern Sweden; the species studied were Betula pubescens tree seedlings and the shrub Vaccinium myrtillus. In this study we showed the importance of facilitative and competitive shrub‐tree interactions as drivers of subarctic treeline dynamics. Furthermore, we demonstrated that the great sensitivity of tree seedlings to warming had strong implications for treeline dynamics under the predicted warmer scenario at high latitudes, and we identified that complex interactions between shrubs and herbivores are critical to predicting future changes. The second chapter is entitled ‘Similar tree seedling responses to shrubs and to simulated environmental changes at Pyrenean and subarctic treelines’. Here we presented a comparison between the results obtained in the experiment presented in the first chapter and those obtained in a parallel experiment performed during a similar period near the treeline in the Alt Pirineu Natural Park, in the Central Pyrenees, Catalonia. This experiment was based on the same factorial design but with different species (i.e. Pinus uncinata tree seedlings and the shrub Rhododendron ferrugineum). To our knowledge, it is the first study which experimentally tests the responses of plants to distinct environmental scenarios in a high mountain ecosystem in the Pyrenees. In this chapter we presented some mechanisms for understanding the recently observed variability of local responses of both subarctic and alpine treelines to currently changing climate while identifying some commonalities that can be used to generalise large scale response of treelines to climate warming. The third chapter focuses on the effects of a dominant dwarf shrub (i.e. Empetrum nigrum) on Pinus sylvestris tree seedlings along a primary succession within a boreal ecosystem on an uplifting island in Bothnian Bay, Finland. This chapter is called ‘An ericoid shrub plays a dual role in recruiting both pines and their fungal symbionts along primary succession gradients’. Here we showed that facilitative and competitive effects of shrubs markedly determined tree seedling establishment and their fungal colonisation along this succession gradient, but in this chapter we did not relate these findings to any environmental changes. As far as we know, we presented the first finding that an ericoid mycorrhizal shrub may enhance both the performance of the ectomycorrhizal host tree and the tree’s fungal symbionts. The study presented in the fourth chapter was performed along a snow‐depth gradient in an extreme arctic tundra ecosystem in the Northeast Greenland National Park, the largest national park in the world. The chapter is entitled ‘Plant interactions and higharctic vegetation composition along a snow‐depth gradient in NE Greenland’. This ecosystem is probably the most sensitive and fragile among the ecosystems studied in this thesis as the eastern coast of Greenland is expected to experience substantial changes in climate due to marked changes in snow precipitation and temperature regimes (Brown and Mote 2009). Here we assessed plant species richness, establishment and composition patterns in distinct growth forms occurring in common arctic plant communities associated with varying snow‐depth during the winter season. This study will help to predict potential diversity and vegetation changes in the high Arctic if snow precipitation regime changes in the future as anticipated.

A redução da cobertura florestal e o uso agrícola intensivo do solo resultam na necessidade de adoção de intervenções de restauração bastante onerosas, o que restringe a expansão dessa atividade e a mitigação dos impactos das atividades humanas na biodiversidade e nos serviços ecossistêmicos. Nesse sentido, a exploração de madeira em florestas em processo de restauração pode ser uma alternativa para a viabilização econômica dessa atividade. Em particular, merece destaque as oportunidades oferecidas pela exploração de espécies arbóreas exóticas de ciclo curto, que antecipam o retorno econômico da restauração e podem auxiliar na amortização dos custos de implantação dos reflorestamentos. No entanto, embora o cultivo intercalado de árvores exóticas seja autorizado pela legislação ambiental como um método de recomposição ambiental, pouco se sabe sobre os efeitos negativos do uso dessas espécies nos modelos de restauração. Nesse contexto, o objetivo deste trabalho foi avaliar o impacto ecológico e silvicultural do uso e colheita de eucalipto consorciado com espécies arbóreas nativas para a restauração da Mata Atlântica. Avaliamos três experimentos implantados nos estados da Bahia e Espírito Santo, os quais adotaram o eucalipto como espécie pioneira comercial, utilizada para gerar renda no primeiro ciclo de corte (em torno de 4 a 5 anos) e ser então removida da floresta em restauração. Em geral, o uso do eucalipto nesses experimentos foi menos prejudicial do que o esperado. Não houve influência negativa do consórcio de eucalipto com espécies nativas, em comparação com plantios exclusivos de espécies nativas, na sobrevivência e crescimento de espécies nativas madeireiras, bem como na regeneração natural de espécies nativas no sub-bosque dos plantios. Nós avaliamos também os impactos da exploração da madeira do eucalipto em dois desses experimentos, onde verificamos que a operação de colheita do eucalipto gera danos nas árvores nativas plantadas e uma abertura significativa do dossel florestal (considerando que o eucalipto representa 50% dos indivíduos plantados e 90% da área basal nos modelos avaliados), porém esses danos já foram parcialmente minimizados em apenas quatorze meses após a colheita. Assim, concluímos que para as condições ambientais avaliadas, o uso do eucalipto em modelos de restauração florestal não prejudicou o desenvolvimento das árvores de espécies nativas plantadas em consórcio e não prejudicou a expressão da regeneração natural, o que torna seu uso válido para essas condições. Mais estudos são necessários, no entanto, para que se acompanhe as trajetórias sucessionais de modelos de restauração florestal a médio e longo prazo, a fim de validar a recomendação do seu uso a partir de bases científicas mais consolidadas.
The reduction of forest cover and the intensive agricultural use of the soil result in the necessity of adopting expensive restoration interventions, which restrict the expansion of this activity and the mitigation of the impacts of human activities on biodiversity and ecosystem services. Thus, the exploitation of wood in restoration forests may be an alternative for the economic viability of this activity. Particularly noteworthy are the opportunities offered by the exploitation of short rotation exotic tree species, which anticipate the profit obtained from restoration and may contribute to the amortization of implantation costs. However, even though the intercropping with exotic trees is authorized by the current environmental legislation as a method for forest restoration, little is known concerning the possible negative effects of the use of these species in restoration models. This research aimed to evaluate the ecological and silvicultural impacts of the use and harvest of Eucalyptus temporarily mixed with native tree species in the restoration of the Atlantic Forest. We evaluated three experiments in the states of Bahia and Espírito Santo, which used Eucalyptus as a commercial pioneer species to generate income after the first rotation (around 4-5 years) and then removed from the system. Eucalyptus had no negative influence over native species\' survival and growth when compared to pure native species plantations, as well as over natural regeneration of native species in the understory. We also evaluated the impacts of harvesting Eucalyptus in two of these three experiments and found that the Eucalyptus harvesting operation causes significant damage to planted native trees and results in opening of the forest canopy (considering that Eucalyptus represents 50% of the individuals planted and 90% of the basal area), but these damages were partially minimized within only 14 months after harvesting. Thus, we conclude that for the environmental conditions evaluated, the mixture with Eucalyptus in this alternative forest restoration approach did not affect the development of native tree species neither the expression of natural regeneration, which makes its use valuable. Further studies are necessary, however, to follow the succession trajectories of forest restoration models in the medium and long runs, to validate the recommendation of their use with more consolidated scientific basis.

Includes bibliographical references.
The central question and motivation behind this paper is to determine whether trade preference agreements facilitate permanent economic development, or if they merely temporarily increase the volume of exports over the period of preferential market access. The paper will evaluate this, by using the case study of the African Growth and Opportunity Act (AGOA) enacted by the United States (US) in 2000. The literature and empirical data show that exports do increase in certain cases under trade preference agreements, however what is missing to a large extent is the impact that these increased exports have on facilitating competitive industrial development through learning-by-doing spill over effects. The objective of this paper is to identity evidence which supports the notion that trade preferences have the potential to advance competitive industrial development, by specifically looking at the impact that AGOA has had on eligible Sub-Saharan African (SSA) countries exports to third countries since its enactment in 2000 as the indicator of trade induced permanent economic development. This is one indicator of many, such as labour productivity, manufacturing output, foreign direct investment, and GDP per capita which could also be used as indicators of development. The remainder of this paper is organised as follows. In the second section, a review of the theoretical and empirical literature with respect to trade preferences and specifically AGOA preferences is discussed. Section three presents an empirical analysis, firstly in terms of a graphical analysis which is then followed by an econometric analysis. The aim of the empirical analysis is to firstly understand the aggregate story of apparel exports under AGOA preferences, and secondly to test the relationship that trade preferences facilitate economic development through enhanced trade. The fourth section concludes the paper by emphasising the key findings, issues and policy recommendations of the paper.

The general aim of my dissertation was to investigate the role of plant interactions in driving population dynamics. Both theoretical and empirical approaches were employed. All my studies were conducted on the basis of metabolic scaling theory (MST), because the complex, spatially and temporally varying structures and dynamics of ecological systems are considered to be largely consequences of biological metabolism. However, MST did not consider the important role of plant interactions and was found to be invalid in some environmental conditions. Integrating the effects of plant interactions and environmental conditions into MST may be essential for reconciling MST with observed variations in nature. Such integration will improve the development of theory, and will help us to understand the relationship between individual level process and system level dynamics. As a first step, I derived a general ontogenetic growth model for plants which is based on energy conservation and physiological processes of individual plant. Taking the mechanistic growth model as basis, I developed three individual-based models (IBMs) to investigate different topics related to plant population dynamics: 1. I investigated the role of different modes of competition in altering the prediction of MST on plant self-thinning trajectories. A spatially-explicit individual-based zone-of-influence (ZOI) model was developed to investigate the hypothesis that MST may be compatible with the observed variation in plant self-thinning trajectories if different modes of competition and different resource availabilities are considered. The simulation results supported my hypothesis that (i) symmetric competition (e.g. belowground competition) will lead to significantly shallower self-thinning trajectories than asymmetric competition as predicted by MST; and (ii) individual-level metabolic processes can predict population-level patterns when surviving plants are barely affected by local competition, which is more likely to be in the case of asymmetric competition. 2. Recent studies implied that not only plant interactions but also the plastic biomass allocation to roots or shoots of plants may affect mass-density relationship. To investigate the relative roles of competition and plastic biomass allocation in altering the mass-density relationship of plant population, a two-layer ZOI model was used which considers allometric biomass allocation to shoots or roots and represents both above- and belowground competition simultaneously via independent ZOIs. In addition, I also performed greenhouse experiment to evaluate the model predictions. Both theoretical model and experiment demonstrated that: plants are able to adjust their biomass allocation in response to environmental factors, and such adaptive behaviours of individual plants, however, can alter the relative importance of above- or belowground competition, thereby affecting plant mass-density relationships at the population level. Invalid predictions of MST are likely to occur where competition occurs belowground (symmetric) rather than aboveground (asymmetric). 3. I introduced the new concept of modes of facilitation, i.e. symmetric versus asymmetric facilitation, and developed an individual-based model to explore how the interplay between different modes of competition and facilitation changes spatial pattern formation in plant populations. The study shows that facilitation by itself can play an important role in promoting plant aggregation independent of other ecological factors (e.g. seed dispersal, recruitment, and environmental heterogeneity). In the last part of my study, I went from population level to community level and explored the possibility of combining MST and unified neutral theory of biodiversity (UNT). The analysis of extensive data confirms that most plant populations examined are nearly neutral in the sense of demographic trade-offs, which can mostly be explained by a simple allometric scaling rule based on MST. This demographic equivalence regarding birth-death trade-offs between different species and functional groups is consistent with the assumptions of neutral theory but allows functional differences between species. My initial study reconciles the debate about whether niche or neutral mechanisms structure natural communities: the real question should be when and why one of these factors dominates. A synthesis of existing theories will strengthen future ecology in theory and application. All the studies presented in my dissertation showed that the approaches of individual-based and pattern-oriented modelling are promising to achieve the synthesis.

L'équilibre arbre-herbe dans la savane tropicale est reconnu comme l'une des principales

énigmes de l'écologie des plantes. Les origines du difficile équilibre entre ces formes de vie

sur des dizaines de millions de kilomètres carrés à l'échelle mondiale sont en partie attribuées

aux perturbations fréquentes induites par les feux de végétation dont les effets varient dans

l'espace et dans le temps selon les conditions environnementales. Les résultats de recherche

dépendent de l’échelle et les conclusions tirées d’études locales sont rarement transposables à

d’autres échelles. La question du transfert d'échelle s’avère donc cruciale dans l'étude des

effets du feu, et nécessite une approche transdisciplinaire.

En raison de la variété des échelles couvertes, cette étude constitue une première dans la

confrontation de données sur l’historique des feux dérivé de l'imagerie satellitaire à des

données de terrain incluant des mesures détaillées sur la structure et la composition de

végétation, ainsi que des propriétés édaphiques et topographiques. Elle s’est focalisée sur la

composante ligneuse de par son caractère pérenne et son influence sur les processus

écologiques majeurs. Sur une zone de plus de 2000 km², le Parc National du W du Niger

(PNWN), où le feu est utilisé comme outil pour la gestion et la conservation des écosystèmes

semi-arides, une carte de l'historique des feux a été élaborée à partir d'images MODIS de 250

m de résolution spatiale et de résolution temporelle journalière couvrant une période de sept

années (2002-2009). Pour comprendre la variabilité, à la fois dans l'espace et le temps, de la

propagation du feu dans la végétation, nous avons étudié les caractéristiques de distribution

des feux en termes de régime du feu (i.e. période d'occurrence et fréquence) et de structure

spatiale (métriques paysagères). Les relations causales plausibles entre les régimes du feu, les

conditions édaphiques et topographiques à l'échelle régionale comme locale, et les

caractéristiques de la végétation ligneuse (composition et structure) ont été examinées à

travers des analyses multivariées et des modèles d'équations structurales. Nous avons aussi

examiné plus en détails les stratégies adaptatives mises en oeuvre par les ligneux, et les

interactions biologiques qui sous-tendent l'organisation spatiale des ligneux à travers une

approche des processus ponctuels.

Les résultats montrent que l'activité du feu dans le PNWN se caractérise par une hétérogénéité

spatio-temporelle induite principalement par les conditions édapho-topographiques via la

structure de la végétation ligneuse. Les feux précoces de gestion créent des pare-feux

efficaces, limitant une large extension des feux de saison tardifs. Cependant, ces feux tardifs

pourraient ne pas être aussi destructifs comme qu’on le suppose généralement. En effet,

l'adaptation des espèces aux différents régimes defeu correspond à des stratégies de croissance

contrastées. Dans le cas des feux tardifs, les surfaces terrières et hauteurs moyennes les plus

fortes sont rencontrées, permettant aux arbres de résister au feu. Quant aux zones non

affectées par les feux l'analyse "patron-processus" désigne clairement la facilitation entre

ligneux comme un processus fondamental de l'organisation spatiale périodique du couvert, une

structure émergente qui empêche le passage du feu. Bien qu’ils ne se substituent pas aux

études expérimentales, ces résultats basés sur une expérimentation naturelle à large échelle

apportent des informations nouvelles précieuses tant au niveau fondamental que pour la mise

en place d'une gestion raisonnée du PNWN.

The tree-grass equilibrium in tropical savanna is recognized as one of plant ecology's main

conundrums. The origins of the difficult balance between these life forms over tens of millions

of square kilometers worldwide are in part attributed to the frequent disturbances caused by

vegetation fires effects of which vary in space and time depending on local environmental

factors. Research results are scale-dependent and findings from local studies are rarely

transposable to higher levels of ecosystem organization. The question of scaling (scale

transfer) is therefore crucial in the study of fire effects, and requires a multidisciplinary

approach.

Because of the variety of scales covered, this study is a premiere in the confrontation of

satellite-imagery derived fire history data with detailed field data including measurements of

vegetation parameters (structure and composition), as well as soil and topographic properties.

The study focuses on the woody component, because of its perennial character and its

influence on major ecological processes. On an area of more than 2000 km², the W National

Park of Niger (WNPN) where fire is used as a tool for the management and conservation of

semi-arid ecosystems, a fire history map was elaborated from MODIS images with a 250 m

spatial resolution and a daily temporal resolution over a period of seven years (2002-2009). To

understand the variability, both in space and time, of fire propagation in vegetation, we studied

the fire distribution characteristics in terms of fire regime (i.e. timing and frequency) and

spatial structure (landscape metrics). Plausible causal relationships at regional and local scales

between fire regimes, edaphic and topographic conditions, and the woody vegetation

(composition and structure) characteristics were examined through multivariate analyses and

structural equations models. We also examined in detail the woody species adaptive strategies

as well biological interactions, which underlie their spatial organization, using point statistics.

Results show that the WNPN fire's activity is characterized by spatial and temporal

heterogeneity induced mainly by edaphic and topographic conditions via the structure of the

ligneous component. Prescribed early season fires create effective firewalls, limiting wide late

season fires. However, these late fires might not be as destructive as is commonly assumed.

Indeed, species adaptation to different fire regimes corresponds to contrasting growth

strategies. In the case of late fires, increased basal areas and mean tree heights were

encountered, enabling trees to resist fire and escape flames. As for the unburned areas, the

"pattern-process" analysis clearly indicates that facilitation between shrubs is a fundamental

process determining the woody cover periodic spatial organization, an emergent structure that

prevents fire spread.

Although they do not replace experimental studies, these results based on a large-scale natural

experiment provide valuable new information both on a fundamental level and for setting up

the rational management of the WNPN.
Doctorat en Sciences
info:eu-repo/semantics/nonPublished

The role of competition in tree communities is increasingly well understood, while little is known about the patterns and mechanisms of the interplay between above- and belowground competition in tree communities. This knowledge, however, is crucial for a better understanding of community dynamics and developing adaptive near-natural management strategies. We assessed neighbourhood interactions in an unmanaged old-growth European beech (Fagus sylvatica) forest by quantifying variation in the intensity of above- (shading) and belowground competition (crowding) among dominant and co-dominant canopy beech trees during tree maturation. Shading had on average a much larger impact on radial growth than crowding and the sensitivity to changes in competitive conditions was lowest for crowding effects. We found that each mode of competition reduced the effect of the other. Increasing crowding reduced the negative effect of shading, and at high levels of shading, crowding actually had a facilitative effect and increased growth. Our study demonstrates that complementarity in above- and belowground processes enable F. sylvatica to alter resource acquisition strategies, thus optimising tree radial growth. As a result, competition seemed to become less important in stands with a high growing stock and tree communities with a long continuity of anthropogenic undisturbed population dynamics. We suggest that growth rates do not exclusively depend on the density of potential competitors at the intraspecific level, but on the conspecific aggregation of large-diameter trees and their functional role for regulating biotic filtering processes. This finding highlights the potential importance of the rarely examined relationship between the spatial aggregation pattern of large-diameter trees and the outcome of neighbourhood interactions, which may be central to community dynamics and the related forest ecosystem services.

Anxiety is one of the most studied research topics in sport psychology literature (Guillen & Sanchez, 2009); however, even though the Olympics are considered to be one of the most pressure-filled sporting events (Birrer, Wetzel, Schmidt, & Morgan, 2012), to date there has been no research aimed specifically at investigating Olympic athletes’ competitive state anxiety and its impact on subsequent performance. Furthermore, according to Nesti (2011), in order to support athletes in dealing with their experience of anxiety, researchers must turn towards the phenomenological, real-lived experience of the athlete to uncover what might best support positive anxiety management and interpretation in competition. The purpose of this study, therefore, was to respond to the gap in Olympic athlete anxiety research by examining the phenomenological experience of competitive state anxiety for female beach volleyball players at the 2012 London Olympics. Six in-depth, phenomenological interviews were conducted with these Olympic female beach volleyball players. Results indicated that, while all athletes in this study experienced anxiety at the 2012 Olympics, it was not the reduced intensity of anxiety that positively impacted their performance but rather the athletes’ ability to recognize, manage, and positively interpret their anxiety. In addition, it was shown that self-confidence further buffered the potentially negative impacts of anxiety. It is recommended that future research focus on extending phenomenological anxiety research to other sports and genders, and to specifically examine the impact of trait anxiety, team dynamics, and the experience of flow on athletes’ anxiety interpretation.