At broad scales, community ecologists study how biogeographic factors like environmental dissimilarity and geographic distance influence community assembly and composition. At small scales, community ecologists study how one or several species interact to determine habitat partitioning and coexistence. In this dissertation, I present studies at both scales. Chapter One investigates community assembly across the Caribbean, Central, and South American radiations of Anolis lizards and Eleutherodactylid frogs to test whether oceanic islands are unique in their assembly processes. Such uniqueness is suggested by high levels of endemism on islands; however, comparable levels of endemism can be found in mainland communities. I modeled the rate of species turnover between mainland communities, with respect to geographic distance and environmental dissimilarity, and then used the mainland model to predict turnover among islands. Turnover among island communities was significantly higher than predicted from the mainland model, confirming the long-held but untested assumption that island assemblages accumulate biodiversity differently than their mainland counterparts. Chapter Two reviews the evidence for ecological character displacement (ECD), an evolutionary process whereby two resource competitors diverge from one another in phenotype and resource use, facilitating coexistence in a community. I find that, despite current scientific opinion, the evidence for ECD is equivocal; most cases of ECD pattern fail to rule out processes alternative to resource competition that could create the same pattern. I conclude that better evidence may come from real time tests of ECD. Chapters Three and Four describe just such a test in small island populations of Anolis carolinensis. In Chapter Three, I find that small island populations of A. carolinensis that have come into sympatry with a novel competitor, the invasive A. sagrei, shift their habitat use to become more arboreal, compared to allopatric populations. Consistent with prediction, individuals from sympatric populations have larger toepads with additional adhesive scales - a common adaptation to arboreality in Anolis. In Chapter Four, I describe a common garden experiment that finds that the observed toepad divergence is an evolved response, suggesting rates of divergence for toepad area and scale number on par with well known examples of contemporary evolution.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第22286号 / 理博第4600号 / 新制||理||1660(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 工藤 洋, 教授 田村 実, 准教授 高山 浩司 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Plants adjust to challenging environments by genetically fixed changes and phenotypically plastic response. Alpine environments pose multiple challenges to plant life including cold, high irradiance and short vegetative period. To survive such specific conditions, plants often significantly alter their morphology. In my thesis I studied to which extent specific traits of alpine ecotypes repeatedly appear among independently formed alpine populations and to which extent these changes represent fixed genotypic differentiation vs phenotypic plasticity. To address these questions I performed an experiment in which Arabidopsis arenosa plants from sixteen populations belonging to two ecotypes (alpine and foothill) were grown in conditions resembling alpine vs foothill conditions. Specifically, I modified levels of irradiance and temperature and complemented alpine-like and foothill-like treatment by additional two extreme treatments to reach full-factorial design. I used discriminant and classificatory analysis to examine the overall morphological differentiation characterised by set of twenty measured traits. Then I examined variation in each trait by statistical Bayesian model that I designed for this purpose. I found out that although ecotypes are predominantly differentiated by fixed morphological...

Phenotypic plasticity plays an important role in organisms’ adaptability to environmental change such as global warming caused by greenhouse-gas emissions. One plastic response to increased temperatures is for organisms to shift their phenology. It is of great concern that the phenologies of interacting species, such as plants and pollinators, may be shifting at different rates, causing temporal mismatches, which for plants can lead to unsuccessful reproduction. The “reproductive assurance hypothesis” states that plants capable of self-pollination should be under high selection to employ this as their main reproductive strategy in the event of pollinator scarcity to ensure reproduction, and consequently invest less in attracting pollinators. This study examines how soil warming in the Hengill geothermal area in Iceland affects the flowering phenology, reproductive strategy and investment in attractiveness to pollinators in the self-compatible herb Cerastium fontanum (Caryophyllaceae), when grown in a common garden in Stockholm, Sweden. Previous research showed that C. fontanum from warmed soils flowered earlier in situ than plants from colder soils, and later when grown in a common environment. In this study, C. fontanum plants collected along a temperature gradient followed the same counter-gradient pattern, where plants from warmer soils flowered later than plants from colder soils. Soil temperature at site of origin positively affected flower number but had no effect on flower size, seed production from autogamous self-pollination or visitation rate. Based on my findings it does not appear that C. fontanum, despite having an earlier flowering phenology in situ, is under any selection to alter its reproductive strategy or investment in attractiveness to pollinators when grown in a common temperature, and therefore it seems unlikely that plants are experiencing a temporal mismatch with insect pollinators. However, it would be worthwhile to conduct a similar experiment in Iceland to better understand how an earlier flowering affects pollination systems.

Cette thèse visait à étudier la variabilité génétique et plasticité phénotypique de la vulnérabilité à la cavitation chez le hêtre (Fagus sylvatica L.). A cette fin, nous avons tout d’abord testé les variations dues à la méthodologie de mesure de la vulnérabilité à la cavitation. Notre étude a montré que les variations de vulnérabilité à la cavitation au cours du temps sont faibles. Néanmoins, il est préférable d’éviter de récolter les échantillons durant le début de la période de formation du bois. Par ailleurs il est possible de conserver au frais les échantillons sans dégrader les mesures durant au moins un mois. Nos résultats montrent également que la vulnérabilité à la cavitation peut être variable aussi bien au sein d’un même arbre qu’entre populations in situ. Ensuite, nous avons voulu déterminer la part de plasticité phénotypique de la part de variabilité génétique dans la variabilité de la vulnérabilité à la cavitation. Pour cela nous avons étudié 17 populations de hêtre venant de toute l’Europe et poussant dans une plantation comparative. Parmi les 17 populations 6 d’entre elles ont été étudiées dans trois plantations comparatives différentes (France, Espagne et Slovaquie). Nos résultats indiquent que chez le hêtre la vulnérabilité à la cavitation varie considérablement au sein de chaque population. Ils indiquent également que l’on observe peu de différenciation génétique entre populations. Les résultats montrent également que la plasticité phénotypique de la vulnérabilité à la cavitation est importante entre les plantations, et que le degré de cette plasticité peut varier d’une population à l’autre. Pour finir, nous avons regardé s’il existait des relations entre la vulnérabilité à la cavitation et des paramètres d’efficience hydraulique ainsi qu’avec l’efficience d’utilisation de l’eau. Nos résultats ne permettent pas de mettre en évidence des preuves d’un trade-off entre l’efficience et la sureté du xylème. / This work aimed at studying the genetic variability and the phenotypic plasticity of vulnerability to cavitation in beech (Fagus sylvatica L.). We tested the variations due to the methodology of measuring vulnerability to cavitation. Our study showed that the variations in vulnerability to cavitation over time are low. However, it’s better to avoid collecting samples during the period of initial wood formation. Furthermore it’s possible to store samples without degrading measures for during one month. Our results also show that vulnerability to cavitation can be variable both within the same tree and between populations in situ. We determined the proportion of phenotypic plasticity and the proportion of genetic variability in the variability of vulnerability to cavitation. For this we studied 17 beech populations from all Europe and growing in a common garden. Among the 17 populations 6 of them have been studied in three different common gardens (France, Spain and Slovakia). Our results indicate that for beech, vulnerability to cavitation varies considerably within populations. They also indicate that there are few genetic differentiations between populations. The results show that the phenotypic plasticity of vulnerability to cavitation is important among common garden, and the degree of this plasticity can vary between populations. To complete our study, we looked at the relationships between vulnerability to cavitation and hydraulic efficiency parameters and with the water use efficiency. Our results do not reveal evidence of relationships vulnerability to cavitation and other physiological traits.

La compréhension des mécanismes déterminant le succès des espèces invasives est une étape essentielle dans la gestion des invasions biologiques actuelles et la prédiction des futurs risques d’invasion. En adoptant un cadre d’étude conceptuel intégrant les processus écologiques et évolutifs, l’objectif de ce travail était d’analyser les déterminants de la colonisation de l’ambroisie à feuilles d’armoise en France. Tout d’abord, l’étude des interactions biotiques et abiotiques a permis de montrer la capacité de tolérance de l’ambroisie à l’herbivorie et au stress hydrique. L’ambroisie est capable de tolérer la défoliation grâce à une croissance compensatoire efficace, sans que sa reproduction en soit affectée. Cette forte tolérance à l’herbivorie est maintenue chez les populations introduites, malgré la faible pression des ennemis naturels dans la zone d’introduction. En condition de stress hydrique, l’ambroisie produit une biomasse racinaire supérieure aux espèces présentes dans les communautés qu’elle envahit. De plus, les différences dans les valeurs moyennes pour les traits mesurés suggèrent une occupation différente par l’ambroisie des niches écologiques disponibles. La comparaison en jardins communs de populations de l’aire d’origine avec des populations de l’aire d’introduction isolées et issues du foyer central d’invasion a montré que l’adaptation de l’ambroisie à son environnement reposait principalement sur la plasticité phénotypique plutôt que sur la différenciation des traits. Les études de génétiques quantitatives ont mis en évidence un potentiel évolutif élevé pour les traits liés à la germination. Les traits liés à la morphologie, à la phénologie et à la physiologie de la plante montrent une variance additive et une héritabilité plus faibles et donc un potentiel évolutif moindre. En revanche, la variation dans les normes de réaction indique un potentiel évolutif important de la plasticité phénotypique. La tolérance au stress hydrique et à l’herbivorie sont des facteurs qui potentiellement augmentent la capacité de l’ambroisie à coloniser une large gamme d’habitat. De plus, la plasticité phénotypique et le potentiel évolutif important peuvent favoriser une augmentation ou un déplacement de la niche écologique de l’espèce et ainsi favoriser l’expansion de son aire de répartition / Understanding of the mechanisms behind the success of the invasive species is essential to manage current biological invasions and to prevent the risks of the futures ones. Using a conceptual framework integrating ecological and evolutionary processes, this work aimed to analyse the factors of the common ragweed colonization in France. First of all, the study of biotic and abiotic interactions has shown the ability of common ragweed to tolerate herbivory and water stress. Common ragweed is able to buffer defoliation through an efficient compensatory growth with no consequence on the reproduction. Herbivory tolerance has been maintained in introduced populations even if herbivory pressure is low in the introduction area. Under water stress conditions, common ragweed displayed higher root biomass than the other species present in the communities it invades. In addition, differences in mean trait values suggest different niche occupation by common ragweed compared with its companion species. Common garden experiments comparing native populations vs. invasive isolated and invasive core populations have shown that adaptation ability is mainly due to phenotypic plasticity rather than trait differentiations. Quantitative genetic studies suggest a high evolutionary potential for germination traits. Morphological, phenological and physiological traits exhibit lower standard genetic variation and lower heritability and thus a lower evolutionary potential. However, variation in reaction norms suggests a high evolutionary potential for phenotypic plasticity. Herbivory and water stress tolerance are factors that potentially increase the ability to colonize à large range of habitats. Furthermore, phenotypic plasticity and evolutionary potential may also favour an increase or a shift in species ecological niche and hence may favour the distribution range expansion

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