Historically, the Kentucky Inner Bluegrass blue ash-oak savanna-woodland was the primary ecosystem of the Inner Bluegrass Region (IBR) of Kentucky. After European settlement, the majority (>99%) of Bluegrass savanna was converted to agricultural and urban land uses. Currently remnant savanna tree species are failing to recruit. Therefore, a long-term restoration ecology project researching competition and disturbance on seedling establishment, survival, and growth has been established at Griffith Woods (the largest remaining savanna in Kentucky) in Harrison Co., KY. Fourteen native hardwood tree species (a total of 6,168 seedlings) have been experimentally planted. Light, soil, surrounding vegetation, and herbivory, factors thought to influence seedling survival, have been initially assessed. Results show that soils differed spatially in P, Ca, Mg, Zn, pH, N percent and soil organic matter percent. Light was significantly reduced by diffusive filtering through vegetation. Vegetation biomass was influenced by pH and Mg. Initial seedling survival was high, but significantly differed by species type, location, and soil pH, Mg, and Zn. This research demonstrates that under a similar range of conditions, native hardwood tree seedling establishment is possible. Therefore, the potential exists to restore Bluegrass savanna-woodland in order to return proper ecological functioning into a degraded landscape.

Crop rotation may be used to prevent the continuous selection of particular weed species adapted to one crop type. This might be useful for weed management, economy in herbicide applications and promoting biodiversity. Common simple crop sequences might be diversified by introducing perennial forage crops. Impacts of such perennial crops on weeds were studied with four approaches : 1) Large-scale weed surveys in 632 fields in western France showed that weed species composition differed most strongly between perennial alfalfa crops and annual crops. Comparisons of fields before, during and after perennial alfalfa suggested that community composition varies in a cyclic way during such crop rotations. Several weed species problematic in annual crops were suppressed during and after perennial crops, but the appearance of other species led to equal or even higher plant diversities. 2) A 3-year field experiment with contrasting crop management options allowed an investigation of the underlying mechanisms for this: The absence of soil tillage reduced weed emergence but increased the survival of established plants. The permanent vegetation cover and frequent hay cuttings reduced weed growth, plant survival and seed production. 3) Greenhouse experiments testing the regrowth ability of individual plants after cutting showed strong differences between species and functional groups. An two-factorial experiment suggested that the negative impacts of cutting and competition on weed growth were mainly additive. 4) Special measurements of weed seed predation in the field experiment showed positive correlations with vegetation cover, indicating that this ecosystem service may be particularly fostered by perennial crops. Consistent preferences of seed predators for certain weed species indicates that seed predation may be another cause of the observed weed community shifts.

Un des principaux challenges pour prédire la composition, la structure et la dynamique des communautés végétales est de déterminer comment l’environnement biotique et abiotique va modifier la direction et l’amplitude des interactions entre plantes. L’objectif de ma thèse est de mettre en évidence le rôle prépondérant des interactions complexes (impliquant plus de deux compartiments biotiques) dans la dynamique forestière et plus particulièrement dans la dynamique de régénération des ligneux d’intérêt sylvicole tel que le chêne sessile. Les résultats soulignent l’importance de prendre en compte : (i) l’effet des cervidés sur les patrons de réponses des interactions entre plantes le long des gradients de ressources, (ii) les interactions indirectes et notamment la facilitation indirecte, (iii) des paramètres démographiques différents en relation avec la phénologie des espèces en présence, (iv) la séparation des mécanismes sous-jacents à une interaction écologique multiple via un suivi des conditions environnementales. Ainsi, ma thèse appuie l’idée que les gestions se focalisant sur une unique pression et ignorant les autres pressions ne sont pas capables de maintenir des populations d’espèces cibles car elles ne tiennent pas compte des interactions multiples. Ceci souligne l’importance d’utiliser des stratégies complémentaires pour permettre la pérennité des écosystèmes forestiers et notamment une régénération suffisante dans le cadre des changements globaux que sont le changement climatique et la surabondance de cervidés. / One of the main challenges to predict vegetation dynamics and plant community composition is to identify how biotic and abiotic factors modify the nature and magnitude of plant-plant interactions. The objective of my thesis is to highlight the leading role of multiple interactions (involving more than two biotic compartments) in forest understory dynamics and more specifically regeneration dynamics of target species such as sessile oak. The results emphasize the importance of: (i) effects of deer on response patterns of plant-plant interactions along resource gradients, (ii) indirect interactions, in particular indirect facilitation, (iii) demographic parameters in relation to species phenology, (iv) distinguishing among underlying mechanisms of multiple interactions thanks to environmental monitoring. Thus, my thesis supports the idea that management practices focusing on a single pressure, while ignoring others, are unable to conserve populations of target species because they don’t consider multiple interactions. This highlights the importance to use complementary management strategies to achieve sustainability in the context of global changes (climatic change and deer overabundance).

Les communautés végétales constituent des systèmes complexes au sein desquels de nombreuses espèces, pouvant présenter une large variété de traits fonctionnels, interagissent entre elles et avec leur environnement. En raison de la quantité et de la diversité de ces interactions les mécanismes qui gouvernent les dynamiques des ces communautés sont encore mal connus. Les approches basées sur la modélisation permettent de relier de manière mécaniste les processus gouvernant les dynamiques des individus ou des populations aux dynamiques des communautés qu'ils forment. L'objectif de cette thèse était de développer de telles approches et de les mettre en oeuvre pour étudier les mécanismes sous-jacents aux dynamiques des communautés. Nous avons ainsi développés deux approches de modélisation. La première s'appuie sur un cadre de modélisation stochastique permettant de relier les dynamiques de populations aux dynamiques des communautés en tenant compte des interactions intra- et interspécifiques et de l'impact des variations environnementale et démographique. Cette approche peut-être aisément appliquée à des systèmes réels et permet de caractériser les populations végétales à l'aide d'un petit nombre de paramètres démographiques. Cependant nos travaux suggèrent qu'il n'existe pas de relation simple entre ces paramètres et les traits fonctionnels des espèces, qui gouvernent pourtant leur réponse aux facteurs externes. La seconde approche a été développée pour dépasser cette limite et s'appuie sur le modèle individu-centré Nemossos qui représente de manière explicite le lien entre le fonctionnement des individus et les dynamiques de la communauté qu'ils forment. Afin d'assurer un grand potentiel d'application à Nemossos, nous avons apportés une grande attention au compromis entre réalisme et coût de paramétrisation. Nemossos a ainsi pu être entièrement paramétré à partir de valeur de traits issues de la littérature , son réalisme a été démontré, et il a été utilisé pour mener des expériences de simulations numériques sur l'importance de la variabilité temporelle des conditions environnementales pour la coexistence d'espèces fonctionnellement différentes. La complémentarité des deux approches nous a permis de proposer des éléments de réponse à divers questions fondamentales de l'écologie des communautés incluant le rôle de la compétition dans les dynamiques des communautés, l'effet du filtrage environnementale sur leur composition fonctionnel ou encore les mécanismes favorisant la coexistence des espèces végétales. Ici ces approches ont été utilisées séparément mais leur couplage peut offrir des perspectives intéressantes telles que l'étude du lien entre le fonctionnement des plantes et les dynamiques des populations. Par ailleurs chacune des approches peut être utilisée dans une grande variété d'expériences de simulation susceptible d'améliorer notre compréhension des mécanismes gouvernant les communautés végétales. / Plant communities are complex systems in which multiple species differing by their functional attributes interact with their environment and with each other. Because of the number and the diversity of these interactions the mechanisms that drive the dynamics of theses communities are still poorly understood. Modelling approaches enable to link in a mechanistic fashion the process driving individual plant or population dynamics to the resulting community dynamics. This PhD thesis aims at developing such approaches and to use them to investigate the mechanisms underlying community dynamics. We therefore developed two modelling approaches. The first one is based on a stochastic modelling framework allowing to link the population dynamics to the community dynamics whilst taking account of intra- and interspecific interactions as well as environmental and demographic variations. This approach is easily applicable to real systems and enables to describe the properties of plant population through a small number of demographic parameters. However our work suggests that there is no simple relationship between these parameters and plant functional traits, while they are known to drive their response to extrinsic factors. The second approach has been developed to overcome this limitation and rely on the individual-based model Nemossos that explicitly describes the link between plant functioning and community dynamics. In order to ensure that Nemossos has a large application potential, a strong emphasis has been placed on the tradeoff between realism and parametrization cost. Nemossos has then been successfully parameterized from trait values found in the literature, its realism has been demonstrated and it has been used to investigate the importance of temporal environmental variability for the coexistence of functionally differing species. The complementarity of the two approaches allows us to explore various fundamental questions of community ecology including the impact of competitive interactions on community dynamics, the effect of environmental filtering on their functional composition, or the mechanisms favoring the coexistence of plant species. In this work, the two approaches have been used separately but their coupling might offer interesting perspectives such as the investigation of the relationships between plant functioning and population dynamics. Moreover each of the approaches might be used to run various simulation experiments likely to improve our understanding of mechanisms underlying community dynamics.

La rotation de cultures peut être utilisée pour empêcher la sélection continue d’espèces adventices adaptées à un type de culture. Elle pourrait favoriser la gestion des adventices, l’économie d’herbicides et la biodiversité. Les successions de cultures simples d’aujourd’hui pourraient être diversifiées par des cultures fourragères pérennes. Les impacts des ces cultures sur les adventices ont été étudié utilisant quatre approches : 1) Des relevés d’adventices sur 632 champs dans l’ouest de la France ont montré que la composition spécifique varie le plus entre des cultures fourragères pérennes et des cultures annuelles. Une comparaison des champs avant, pendant, et après des cultures fourragères pérennes a suggéré que la composition des communautés varie d’une manière cyclique pendant ces rotations. Plusieurs espèces problématiques dans des cultures annuelles ont été supprimées pendant et après les cultures pérennes, mais l’apparition d’autres espèces a produit une diversité de plantes comparable, voire supérieure. 2) Une expérimentation au champ de trois ans avec des modes de gestion contrastés a permis d’étudier les mécanismes sous-jacents: L’absence de travail du sol a réduit la levée des adventices, mais a augmenté la survie des plantes adultes. Le couvert végétal permanent et les fauches fréquentes ont réduit la croissance, la survie des plantes et la production de graines. 3) Des expérimentations sous serre analysant la croissance poste fauche de plantes individuelles ont montré des différences importantes entre espèces et groupes fonctionnels. Une expérimentation à deux facteurs a suggéré que les impacts négatifs de la fauche et de la compétition sur la croissance des adventices ont été additifs. 4) Des mesures spéciales de prédation de graines d’adventices sur l’expérimentation au champ ont montré des corrélations positives avec le couvert végétal et la prédation, indiquant une importance particulière de ce service écosystémique dans des cultures pérennes. La préférence des graines de certaines espèces montre que la prédation de graines peut être une autre cause des changements de communautés d’adventices. / Crop rotation may be used to prevent the continuous selection of particular weed species adapted to one crop type. This might be useful for weed management, economy in herbicide applications and promoting biodiversity. Common simple crop sequences might be diversified by introducing perennial forage crops. Impacts of such perennial crops on weeds were studied with four approaches : 1) Large-scale weed surveys in 632 fields in western France showed that weed species composition differed most strongly between perennial alfalfa crops and annual crops. Comparisons of fields before, during and after perennial alfalfa suggested that community composition varies in a cyclic way during such crop rotations. Several weed species problematic in annual crops were suppressed during and after perennial crops, but the appearance of other species led to equal or even higher plant diversities. 2) A 3-year field experiment with contrasting crop management options allowed an investigation of the underlying mechanisms for this: The absence of soil tillage reduced weed emergence but increased the survival of established plants. The permanent vegetation cover and frequent hay cuttings reduced weed growth, plant survival and seed production. 3) Greenhouse experiments testing the regrowth ability of individual plants after cutting showed strong differences between species and functional groups. An two-factorial experiment suggested that the negative impacts of cutting and competition on weed growth were mainly additive. 4) Special measurements of weed seed predation in the field experiment showed positive correlations with vegetation cover, indicating that this ecosystem service may be particularly fostered by perennial crops. Consistent preferences of seed predators for certain weed species indicates that seed predation may be another cause of the observed weed community shifts. / Fruchtfolgen können dazu dienen, die kontinuierliche Selektion von Unkrautarten zu verhindern, die an eine bestimmte Kultur angepasst sind. Dies könnte dem Unkrautmanagement, der Einsparung von Herbiziden, und der Biodiversität dienen. Heutige, sehr einfache Furchtfolgen könnten durch mehrjährige Futterkulturen diversifiziert werden. Die Einflüsse solcher mehrjähriger Kulturen auf Unkräuter wurden in vier Ansätzen untersucht : 1) Vegetationsaufnahmen auf 632 Feldern in Westfrankreich zeigten, dass die Unkrautzusammensetzung zwischen mehrjährigen Futterkulturen und einjährigen Kulturen stark variiert. Der Vergleich von Feldern vor, während und nach mehrjährigen Futterkulturen legte nahe, dass die Pflanzengemeinschaft während solcher Fruchtfolgen zyklisch variiert. Mehrere problematische Unkrautarten wurden während und nach den mehrjährigen Kulturen zurückgedrängt. Das Auftauchen anderer Arten führte jedoch zu einer gleichbleibenden oder leicht erhöhten Pflanzenvielfalt. 2) Ein dreijähriger Feldversuch mit verschiedenen Bearbeitungsoptionen ermöglichte es, die zugrunde liegenden Mechanismen zu untersuchen: Die fehlende Bodenbearbeitung hat das Auflaufen der Unkräuter reduziert und das Überleben der adulten Pflanzen erhöht. Die permanente Vegetationsbedeckung und die häufigen Heuschnitte haben das Wachstum, das Überleben und die Samenproduktion vermindert. 3) Gewächshausexperimente zum Nachwachsen von Unkrautpflanzen nach Heuschnitten zeigten große Unterschiede zwischen verschiedenen Arten und funktionellen Gruppen. Ein Experiment mit zwei Faktoren lässt vermuten, dass die negativen Effekte der Schnitte und der Konkurrenz auf das Unkrautwachstum sich addieren. 4) Spezielle Messungen der Prädation von Unkrautsamen auf den untersuchten Feldern zeigten positive Korrelationen mit der Vegetationsbedeckung, was auf eine besondere Wichtigkeit dieser Ökosystemdienstleistung in ausdauernden Kulturen hindeutet. Die Präferenz von bestimmten Samenarten deutet darauf hin, dass Samenprädation ein weiterer Grund für die beobachteten Änderungen der Unkrautgemeinschaften sein kann.

L’urbanisation transforme et fragmente le paysage en plus de changer son climat et d’induire une homogénéisation biotique mondiale, c’est-à-dire une perte de biodiversité inter-habitats. Qu’elles soient historiques ou contemporaines, naturelles ou humaines, les nombreuses perturbations au sein d’une ville compliquent l’étude des relations entre les plantes et leur environnement, mais l’originalité de cette thèse est de les décortiquer soigneusement à diverses échelles de temps et d’espace. Mes recherches visent à comprendre l’effet de l’urbanisation sur la diversité floristique et fonctionnelle des boisés. L’étude se déroule à Montréal et sur trois îles environnantes, au cœur de la région la plus peuplée du Québec. J’examine d’abord les changements temporels des communautés végétales d’un boisé, et identifie les processus en cause. La comparaison d’inventaires effectués au Bois-de-Saraguay (96 ha), avec ceux d’une étude similaire menée 30 ans plus tôt, a permis d’atteindre cet objectif. Je regarde ensuite s’il est possible de recourir aux ptéridophytes pour estimer l’intégrité forestière urbaine. À cette fin, la diversité ptéridologique a été répertoriée au sein de 82 boisés (paysage) et de 225 quadrats (microhabitats). Des variables révélatrices des perturbations ont servi à définir les niveaux d’intégrité. En troisième lieu, j’identifie les rôles des contraintes (filtres) associées à l’urbanisation et des processus sous-jacents dans l’assemblage des communautés de sous-bois. Pour ce faire, la diversité des plantes vasculaires de sous-bois a été recensée dans 50 boisés et 431 quadrats. Avec ces données, 18 aspects de la diversité taxinomique et fonctionnelle ont été examinés. Des filtres de grande (paysage) et fine (habitat) échelles ont été définis à partir de variables environnementales et spatiales. Mon dernier objectif consiste à dresser le portrait de la flore de l’aire d’étude. Mon analyse des communautés du Bois-de-Saraguay révèle qu’une homogénéisation fonctionnelle du sous-bois s’y est produite. Celle-ci résulte d’une réorganisation spatiale des populations dans laquelle des espèces similaires — comprenant les plantes ligneuses à fruits charnus et clonales — sont devenues dominantes. Selon mes résultats, ces changements ont commencé grâce à des événements de dissémination des espèces, puis en suivant les perturbations anthropiques en place. Mon analyse de la diversité ptéridologique montre qu’elle diminue dans les boisés perturbés par l’effet d’îlots de chaleur et par l’effet de lisières. Des niveaux d’intégrité du paysage forestier et des microhabitats peuvent donc être indiqués par les ptéridophytes. Mon analyse de la flore de sous-bois montre que les caractéristiques contemporaines des forêts, de la matrice urbaine et des habitats locaux représentent les trois catégories de filtres qui façonnent le plus d’aspects de la diversité végétale. L’histoire forestière influence peu d’aspects, hormis la présence de plantes exotiques par exemple. L’assemblage des communautés est principalement contrôlé par l’environnement et très secondairement par des processus spatiaux tels que la dissémination végétale. L’analyse des filtres révèle que les grands boisés sont riches en espèces, alors que les petits ou ceux de formes allongées favorisent les plantes clonales ou de grande taille. Les plantes se disséminant sans assistance se rencontrent davantage dans les boisés des zones faiblement urbanisées, les arbustes exotiques dans les zones résidentielles et les espèces à fruits adhésifs dans les zones fortement bâties. Les îlots de chaleur font chuter la diversité de traits fonctionnels. À petite échelle, la couverture arborée et arbustive influence fortement le type de diversité des sous-bois. De plus, mes inventaires dévoilent que les boisés étudiés sont très riches en plantes indigènes, qu’ils recèlent encore de nouvelles espèces pour le Québec, et ne comptent pas moins de 20 % de la flore provinciale. En somme, ma thèse milite en faveur de la conservation de l’ensemble des boisés urbains, même ceux de petite taille. / Urbanization transforms and fragments the landscape. It also changes its climate and induces a global biotic homogenization, namely a loss of between-habitat biodiversity. Whether historical or contemporary, natural or human, the numerous disturbances within a city make it difficult to study the relationships between plants and their environment. The originality of this thesis is to dissect them at various scales of time and space carefully. In this context, my main aim is to understand the effect of urbanization on the floristic and functional diversity of woodlands. My study takes place in Montreal and on three surrounding islands of the most populated region of Quebec. I first examine the temporal changes of communities in a forest park and identify the processes involved. The comparison of inventories carried out in the Bois-de-Saraguay (96 ha), with similar surveys conducted 30 years ago, made it possible to achieve this objective. Second, I consider the possibility of using pteridophytes to estimate the urban forest integrity. To this end, I recorded pteridological diversity among 82 woodlands (landscape) and 225 quadrats (microhabitats). I used metrics of disturbances to define the integrity levels. Third, I identify the roles of constraints (filters) associated with urbanization and underlying processes in the assembly of understorey communities. To this aim, I identified the diversity of understorey vascular plants in 50 woodlands and 431 quadrats. From these data, I examined 18 taxonomic and functional diversity aspects. I defined large (landscape) and fine (habitat) scale filters based on environmental and spatial variables. My final objective is to describe the flora of the study area. My analyses reveal that a functional homogenization occurred in the understorey communities of the Bois-de-Saraguay. This homogenization results from a spatial reorganization of communities in which species with similar traits — including woody plants with fleshy fruits and clonality — became dominant. According to my results, these changes began through species dispersal events, then following anthropogenic disturbances in place. Pteridological diversity decreases in woodlands disturbed by the heat island effect and by the edge effect, suggesting that this group of species is a good surrogate for urban forest integrity. My analyses of the understorey flora show that the contemporary features of forests, urban matrix and local habitats represent the three categories of filters that shape the most aspects of plant diversity. Forest history influences few aspects, apart from the presence of exotic plants for example. The assembly of communities is mainly controlled by the environment and secondarily by spatial processes such as plant dispersal. Analysis of the filters reveals that large woodlands are species rich, while small or elongated ones favor tall or clonal plants. Plants with unassisted dispersal are more common in forests located in weakly urbanized areas, exotic shrubs in residential areas, and adhesive-fruit species in highly built-up areas. Urban heat islands significantly reduced the diversity of functional traits. At the fine scale, tree and shrub cover strongly influences the type of understorey diversity. Finally, my inventories reveal that the studied forests are very rich in native plants. They still harbor unrecorded species for Québec and account for no less than 20% of the provincial flora. Overall, my thesis argues for the preservation of all the urban forests, even small ones.

Le brachypode rameux (Brachypodium retusum) est une espèce herbacée pérenne qui domine les pelouses sèches de Méditerranée occidentale. Dans notre zone d’étude du Sud de la France, la recolonisation spontanée est très faible après perturbation du sol. Cette observation ne correspond pas aux résultats d'études menées dans d'autres régions montrant une capacité de colonisation élevée. L'objectif principal de cette thèse était donc de tester les différentes hypothèses pouvant expliquer les différents patrons de colonisation.Dans un premier chapitre, nous avons testé si ces différents patrons de colonisation résultent d'une différenciation génétique entre les populations. Des marqueurs AFLP ont été utilisés pour analyser la structure génétique de 17 populations de Méditerranée occidentale dont la différenciation neutre entre populations. Dans un sous-échantillon de 13 populations françaises, la différenciation de traits phénotypiques sous sélection a été testée dans un jardin commun et comparée à la différenciation neutre. Dans le deuxième chapitre, nous présentons une étude sur la différenciation adaptative des traits phénotypiques et avons testé une potentielle réponse différentielle à la manipulation de facteurs environnementaux clés. Les deux derniers chapitres de la thèse ont analysé les facteurs environnementaux qui limitent la recolonisation in situ, indépendamment du génotype ou de la population. Dans le troisième chapitre, nous avons testé l’effet du pâturage et du feu sur le recouvrement végétatif ainsi que sur la reproduction sexuée de B. retusum installé. Enfin, dans le quatrième chapitre, nous avons mesuré l’effet de l’arrosage initial et du pâturage sur l’installation de plantules transplantées initialement germées en serre et de plantules issues de semis in situ. Nos résultats ont montré que les populations de B. retusum sont génétiquement différenciées pour les marqueurs neutres mais également pour les traits phénotypiques. Cette différenciation est supérieure à la dérive seule et suggère une adaptation aux conditions environnementales, en particulier aux températures estivales et à la fréquence du gel en hiver. Une réponse différentielle à la manipulation expérimentale de facteurs environnementaux (sol, pâturage, humidité du sol) a confirmé le caractère adaptatif de la différenciation génétique. Les expériences in situ ont montré que le feu a un effet positif sur la reproduction de B. retusum et sur la communauté végétale associée, alors que son recouvrement végétatif n’était pas supérieur à la moyenne de la communauté. L'exclusion du pâturage pendant deux saisons n'a eu d'incidence sur aucun des paramètres mesurés chez les populations adultes. L'arrosage initial a affecté l’installation des plantules au cours de la première saison. Au cours de la deuxième saison, l'effet seul de l’arrosage n'était pas significatif, cependant son effet est resté positif sur la survie uniquement dans les parcelles pâturées. Le pâturage, au début du cycle de vie de B. retusum, a eu un effet négatif sur le recrutement et la croissance des plantules. En conclusion, la différenciation adaptative entre les populations peut avoir contribué aux différences régionales en termes de capacité de colonisation et doit être prise en compte lors du ciblage des populations sources pour l'introduction de plantes en restauration écologique. La réponse positive de B. retusum a indiqué que le feu était une force sélective importante dans le passé qui pourrait être utilisée pour favoriser l'espèce et la communauté végétale associée dans des opérations de conservation et de restauration. L'exclusion du pâturage à court terme est tolérée par les populations adultes de B. retusum, mais l'abandon à long terme entraîne une diminution de son recouvrement par rapport aux graminées pérennes à croissance rapide. / Ramose false brome (Brachypodium retusum) is a perennial herbaceous species that dominates dry grasslands of the Western Mediterranean. In our southern French study area, spontaneous re-colonisation is very low after soil disturbance. This observation does not correspond to the results of studies from other regions showing a high colonisation capacity. The major objective of this PhD thesis was to test different hypotheses potentially explaining the different colonisation patterns.In the first chapter, we tested whether these different colonisation patterns are the result of genetic differentiation among populations. AFLP markers were used to analyse genetic structure including neutral population differentiation in 17 Western Mediterranean populations. In a sub-sample of 13 French populations, differentiation in phenotypic traits under selection was tested in a common garden and compared to neutral differentiation. In the second chapter, we present a study on adaptive differentiation in phenotypic traits testing a potentially differential response to the manipulation of key environmental factors. The last two chapters of the PhD analysed environmental factors that limit re-colonisation in the field independent of genotype or population. In the third chapter, we tested the effect of grazing and fire on vegetative recovery as well as on sexual reproduction of established B. retusum and in the fourth chapter we measured the effect of initial watering and grazing on the establishment of transplanted seedlings pre-grown in a greenhouse and of field-sown seedlings.Our results showed that populations of B. retusum are genetically differentiated in neutral markers but also in phenotypic traits. This differentiation is superior to drift alone and suggests adaptation to environmental conditions, particularly to summer temperature and winter frost frequency. A differential response to experimental manipulation of environmental factors (soil, pasture, soil moisture) confirmed the adaptive character of genetic differentiation. The field experiments showed that fire has a positive effect on B. retusum reproduction and on the associated community whereas vegetative recovery was not higher than community average. Two seasons of grazing exclosure did not affect any of the measured parameters in adult populations. Initial watering affected seedling establishment in the first season. In the second season, the watering main effect was not significant but interestingly the effect remained positive on survival in grazed plots whereas no such effect was observed in exclosures. Grazing in early life cycle stages of B. retusum had a negative effect on seedling recruitment and growth.In conclusion, adaptive differentiation between populations may have contributed to regional differences in colonisation capacity and needs to be taken into account in targeting source populations for plant introduction in ecological restoration. The positive response of B. retusum indicated that fire was an important selective force in the past which may be used to favour the species and its associated plant community in current conservation and restoration management. Short-term grazing exclosure is tolerated by mature B. retusum populations but long-term abandonment results in a decrease of cover relative to high-growing perennial grasses. In early stages of seedling establishment grazing should be avoided to guarantee introduction success - or grazing stress needs to be compensated by watering.

Identifying mechanisms that determine who lives and dies is the first step in developing successful restoration techniques for rare species and endangered habitats. We studied interactions that affect establishment of native plant forbs of conservation concern at the seedling stage to support the theoretical basis for restoration activities in Pacific Northwest prairies. Specifically, we tested the hypothesis that seedling establishment is controlled by 1) competition with or 2) facilitation by existing vegetation and that the interaction is mediated in part by environmental stress. We direct-seeded or planted vegetative plugs of Lupinus oreganus, Castilleja levisecta, Erigeron decumbens, Iris tenax and Sidalcea malviflora ssp. virgata into 20 plots with a range of community compositions in high-stress upland prairies at each of three sites. We counted seedlings and estimated cover of plant functional groups as well as litter, bare soil and disturbance then used linear regression to test for effects of these factors on seedling establishment. We found evidence of indirect facilitation of grass on seedling establishment in the first year: higher accumulations of leaf litter increased seedling numbers at two sites. In the second year, there was evidence of facilitation by live vegetation and litter on seedlings at one site, but no net effect of either competition or facilitation at the other two sites. Overall, we found more evidence for positive interactions than we did for competition. In particular, litter appeared to have a positive effect on seedling establishment of L. oreganus and S. malviflora ssp. virgata. This is contrary to the common perception that litter inhibits plant establishment but supports the theory that facilitation is more common in high stress sites; practitioners should consider seeding into leaf litter at some sites. To support a robust approach to conservation and reintroduction of species with dormant seed, we characterized dormancy types and developed germination protocols for S. malviflora ssp. virgata and I. tenax. S. malviflora ssp. virgata has physical dormancy and may have physiological dormancy. Scarification followed by four weeks of cold moist stratification was effective in initiating germination. I. tenax has morphophysiological dormancy which is overcome by four weeks of warm moist stratification followed by 6-12 weeks of cold stratification. We also conducted a meta-analysis of experiments that tested pre-sowing seed scarification of L. oreganus and conclude that breaking physical dormancy prior to direct seeding does not support higher establishment relative to unscarified seeds in this species. / Graduation date: 2012

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

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