Master of Science / Biology / Michael Tobler / Environmental variation can shape phenotypic variation in organisms. Most evidence for trait differentiation along environmental gradients comes from analyses of dichotomous habitat types that differ in only one or few environmental factors. In reality, however, environmental variation is often more subtle, gradual, and multifarious. I investigated geographic variation in body shape, trophic resource use, and individual diet specialization in two species of darters (Etheostoma spectabile and E. flabellare; Percidae) that occur along river gradients. I explicitly tested how abiotic and biotic environmental factors shape trait variation within and between species. Results indicated significant among population variation in the body shape of both species. Population differences in body shape were correlated with variation in substrate composition. Although body shape analyses revealed a small but significant signal of convergent evolution of body shape when both species occur in sympatry, E. spectabile and E. flabellare mostly exhibited unique responses to shared sources of selection. The analyses of darter trophic resource use uncovered significant resource partitioning between the two species and geographic variation in diets that is likely driven by differences in resource availability. Furthermore, the majority of populations exhibited significant individual specialization. Variation in individual specialization in populations of E. flabellare was related to invertebrate density and competitor richness, and in E. spectabile to the combined effects of invertebrate density and invertebrate diversity. My results indicate substantial variation in trophic resource use among individuals, populations, and species of small-bodied fishes that are typically assumed to be generalist insectivores. Variation in diet specialization may be more widespread than previously considered, and ecological opportunity is an important factor in shaping trophic resource use of individuals and populations. Overall, the results indicate that even subtle and gradual environmental variation can induce substantial variation in phenotypes on a relatively small spatial scale.

Les estuaires sont constitués d'une vaste mosaïque d'habitats naturels et semi-naturels. L'objectif de cette thèse est d'étudier les végétations et les paysages végétaux des systèmes estuariens atlantique en France, en essayant de comprendre le fonctionnement et les schémas de distribution des différentes unités de végétation. La présente étude tente de mettre au point un modèle théorique commun de fonctionnement et de gradients écologiques, afin de compléter la classification et la connaissance écologique des estuaires, et une aide au suivi et à l’évaluation de l’utilisation des sols, des impacts humains, en développant un modèle spatio-temporel prédictif fondé sur la végétation réelle et potentielle, et en utilisant l'approche dynamico-caténale. Dans 8 estuaires sélectionnés, nous avons effectué des travaux sur le terrain pour une superficie totale de 98 318 ha, mettant en évidence 2 séries de végétation et 4 géopermaséries, correspondant à 131 associations végétales, 60 alliances, 43 ordres et 28 classes. Nous avons cartographié la végétation de trois estuaires représentatifs, ce qui représente une superficie totale de 74 433 ha. Un schéma synthétique des paysages végétaux estuariens est proposé, en intégrant les gradients géographiques et écologiques et les formes géomorphologiques. / Estuaries generally include a wide mosaic of natural and semi-natural habitats. The objective of this thesis is to study the vegetation and Plant Landscape of French Atlantic estuarine Systems, trying to understand the functioning and the plant distribution patterns. The present study tries to carry out a theoretical common model of functioning and ecological gradients, in order to make a basis to improve their classification and ecological studies, and to Help the monitoring and assessment of land uses, land forms transformation and human impacts : developping a spatio-temporal predictive model based on actual and potential vegetation, using the dynamico-catenal approach.The study area corresponds to the Atlantic French estuaries. In 8 selected estuaries, we undertook fieldworks for a total of 98315 ha, highlighting 2 vegetation series and 4 geopermaseries, corresponding to 131 plant associations, 60 alliances, 43 ordos and 28 classes. We mapped the vegetation of three representative estuaries for a total of 74433 ha. A synthetic scheme of estuary vegetation landscape is proposed, integrating geographical and ecological gradients as well as geomorphologic forms.

Abstract Gradient, which is largely considered to be related to water level in mires, is referred to as a microtopographic mud bottom to carpet to lawn to hummock level gradient or the hummock level to intermediate level (lawn) to flark level gradient. The relationship of this vegetation gradient to various physical water level characteristics was studied. The general classification used in the present summary paper divides the aro vegetation of the inland of Northern Ostrobothnia into two main groups: (a) treeless fen aro vegetation (Juncus supinus, Carex lasiocarpa, Rhynchospora fusca, Molinia caerulea) and (b) heath aro vegetation (Polytrichum commune). The first group (a) was divided into fen aro wetlands with an approximately10 cm peaty layer at most and into aro fens with a peat layer thicker than 10 cm. The treatment of the water level gradient was divided into three main groups. (1) The mean water level correlated with mire surface levels (microtopographic gradient) within mires with slight water level fluctuations and partly within mires with considerable water level fluctuations. (2) Three habitat groups could be distinguished on the basis of the range of water level fluctuation i.e. mires with slight water level fluctuations, mires with considerable water level fluctuations and the aro vegetation with extreme water level fluctuations. (3) The timing of water level fluctuations indicated that there are different types of patterns within aro wetlands, the seasonal pattern being mainly a response to yearly snow melt and the several-year-fluctuation pattern being related to the regional groundwater table fluctuation in mineral soils (heath forests). A link was suggested between the stability of the water regime and peat production in local aapa mire – aro wetland environments. From the point of view of peatland plants the direction of variation from a stable to an unstable water regime in aapa mire – aro wetland environments represents a transition towards more and more harsh ecological conditions, partly forming a gradient through natural disturbance. A qualitative functional model was provided for the mire – aro wetland systems of Northern Ostrobothnia. The model supposes differences in the characteristics of peat between two functional complexes within a mire system. Finally, the model for local mire – aro wetland systems was converted to a general from: diplotelmic (acrotelm) mires were divided into two subtypes (diplotelmic water stabilization mires, diplotelmic water fluctuation mires) and the relationship of those subtypes to percolation mires and seasonal wetlands was considered.

En contexte tempéré, les études sur l'instabilité spatio-temporelle de la sensibilité des essences forestières au climat sont rares et souvent conduites à des échelles locales et régionales ; de telles approches ne permettant pas d'obtenir une vision globale de la réponse à l'environnement et à ses variations. La fusion de jeux de données dendrochronologiques (plus de 4500 arbres carottés) a permis d'analyser le comportement de croissance de sept essences européennes tempérées majeures (Quercus petraea, Fagus sylvatica, Abies alba, Picea abies, Pinus sylvestris, Pinus nigra, Pinus uncinata) dans des contextes climatiques variés (océanique à subalpin) et sur l'ensemble du 20ème siècle. Ce travail a également permis de préciser dans quelles mesures les conditions écologiques locales modulaient cette sensibilité au climat. Les relations cerne-climat ont été évaluées par le calcul de fonctions de corrélation. Quelque soit l'essence et le contexte écologique, la sécheresse estivale est le principal facteur limitant la croissance radiale (mais non l'unique), suivie par la sécheresse de l'automne précédent et enfin le froid hivernal. La variabilité spatiale de la réponse dépend plus fortement de la pluviométrie que des températures, une pluviométrie faible conduisant à une sensibilité plus forte au froid hivernal et aux sécheresses estivale et automnale. Ce comportement général est modulé par les conditions écologiques locales, avec une sensibilité à la sécheresse moindre sur sol profond. Les différences interspécifiques s'expriment principalement hors saison de végétation (novembre à mars), même si les corrélations sont rarement significatives. La croissance des résineux est généralement stimulée par des fins d'hiver chauds (février à avril), alors que celle des feuillus est corrélée négativement aux températures et positivement aux précipitations en décembre et janvier. Ces différences entre essences s'avèrent plutôt stables le long des gradients climatiques. Enfin, l'analyse temporelle révèle de fortes instabilités des relations cerne-climat au cours du siècle dernier. Le sens et l'ampleur de ces variations sont homogènes le long des gradients écologiques, mais en revanche peu synchrones avec les instabilités climatiques (automne, hiver, printemps) ou écophysiologiquement peu logiques (été). Cette faible cohérence entre tendances climatiques et instabilité de la sensibilité au climat pourrait s'expliquer par l'absence d'une contrainte climatique de croissance unique en contexte tempéré, où la largeur de cerne est sous le double contrôle du froid hivernal et du stress hydrique estival (et automnal). Elle pourrait également provenir de phénomènes non climatiques, tels que l'effet biologique lié au vieillissement ou l'évolution progressive des pratiques de gestion forestière. Les analyses des variations spatio-temporelles de sensibilité au climat questionnent également sur la précision des relations cerne-climat, estimée le plus souvent au travers du calcul des fonctions de corrélation. En effet, les comparaisons inter-région, inter-site et inter-période des réponses révèlent souvent des variations de corrélations dont les grandeurs pourraient être de l'ordre de la précision liée à l'échantillon considéré. Nous proposons ici de quantifier l'effet de la taille (nombre d'arbres carottés) et des caractéristiques de l'échantillon (nombre de placettes, nombre d'arbres par placette, statuts sociaux couverts) sur la qualité de l'estimation du signal environnemental contenu dans la chronologie moyenne et des fonctions de corrélation. Cette analyse a permis également de préciser dans quelles mesures les différences (1) de traits fonctionnels entre espèces et (2) de contextes climatiques (plus ou moins limitants) modulent cet effet « échantillon ». [...] Suite et fin du résumé dans la thèse. / In temperate conditions, studies dealing with spatio-temporal instabilities in climate sensitivity of forest tree species are scarce and often led at local and regional scales, which prevents from drawing global responses to the environment and its variations. The dendrochronological dataset merging (more than 4500 cored trees) allowed analyzing the growth pattern of seven major European species (Quercus petraea, Fagus sylvatica, Abies alba, Picea abies, Pinus sylvestris, Pinus nigra, Pinus uncinata) in various climatic contexts (oceanic to subalpine) and over the 20th century. This thesis also investigated the climate sensitivity modulation by local ecological conditions. Climate-growth relationships were studied through the calculation of correlation functions. Regardless of the species and the ecological context, summer drought is the main growth limiting factor (but not the unique one), followed by previous autumn drought and winter frost. Spatial variability in response to climate depends more heavily on pluviometry than temperature, decreasing amount of precipitation leading to increasing sensitivities to summer and previous autumn droughts and also winter frost. This general pattern is modulated the local ecological conditions, with especially a lower sensitivity to drought on deep soils. Species-specific responses to climate are mainly evidenced out of the growing season (November to March), even if correlations are rarely significant. The growth of conifers is generally enhanced by warm late winters (February to April), while that of broadleaves is negatively correlated to temperatures and positively to precipitation in December and January. These between-species differences turn out to be stable along the climatic gradients. Lastly, the temporal analysis evidences strong climate-growth relationships instabilities over the last century. The way and the magnitude of these variations are rather homogenous along the ecological gradients, but display low synchronicity with climatic instabilities (autumn, winter and spring) and are ecophysiologically difficult to explain (summer). Such incoherencies between climatic trends and climate sensitivity trends could be related to absence of a single growth limiting factor under temperate context, since tree-ring is under the control of both winter frost and summer (and autumn) drought. They could also result from non-climatic phenomenon, such as the biological the age-related biological effect or progressive changes in forest management. The analyses of spatio-temporal variations in sensitivity climate question on the precision of the climate-growth relationships, most of the time estimated with correlation functions. Indeed, inter-plot, inter-region and inter-period comparisons of responses often highlight differences in correlations which could be of the same magnitude than that of the precision related to the investigated sample. We thus propose to quantify the effect of the sample size (number of cored trees) and characteristics (number of plots, number of trees per plot, sampled social statuses) on the accuracy of the estimation of both the environmental signal estimation contained in the growth chronology and the correlation functions. This analysis also investigates the modulation of such effects by the species-specific functional traits and the strength of the environmental growth limitation. [...] Last and final summary in the thesis.

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / Biological diversity is a broad, deep and multifaceted concept that describes all variability of living forms on different spatial and temporal scales and levels of organization. Even among species, diversity can assume distinct characters, including a taxonomic, a phylogenetic and a functional one. Semiarid ecosystems such as the Caatinga were poorly addressed from an integrated perspective of its biodiversity. The region possess high biotic variability and undergoes intense pressures from abiotic and anthropogenic factors. With around 500 species known in the biome, birds compose an expressive biological group in the Caatinga, with distinct evolutionary lineages and ecological features. This study aims to understand the evolutionary and ecological aspects of the diversity of birds of the Caatinga and their environmental drivers. Specifically, we aim to (i) summarize the existing knowledge on birds assemblages in the biome, their phylogenetic relationships and ecological (i.e., functional) differences; (ii) investigate the interrelationships among taxonomic, phylogenetic and functional diversity, including their distinct measures; (iii) evaluate the effects of different environmental (climatic and structural) gradients on these different dimensions of bird diversity; and (iv) evaluate the effects of these factors on the functional diversity based on individual traits. We gathered information of 123 birds assemblages from the Caatinga based on the available literature, and used a species-level phylogenetic tree and a set of speciesspecific functional traits to estimate the phylogenetic and functional diversities, respectively. Also, we explored different diversity metrics, aiming to understand their interrelationships and select those less biased by species richness and sampling effort. 523 species composed the regional diversity of the Caatinga. The phylogenetic and functional diversity patterns were coincident regarding spatial distribution and environmental hypothesis, with current climate as the leading factor. Nonetheless, the diversity of individual traits diverged qualitatively and quantitatively from the total functional diversity. These results reinforce the need to intensify the sampling efforts in the Caatinga, allow us to comprehend the effects of environmental conditions on birds assembling in ecological and evolutionary terms and, finally, warns to the problem of selecting ecological traits to describe functional diversity. / Diversidade biológica é um conceito amplo, profundo e multifacetado que descreve toda variabilidade de formas de vida em diferentes escalas espaciais, temporais e níveis de organização. Mesmo entre espécies, a diversidade pode assumir distintos caráteres, incluindo o taxonômico, o evolutivo e funcional. Ecossistemas semiáridos como a Caatinga foram pouco estudados sob uma perspectiva integrada da biodiversidade. A região possui alta variabilidade biótica e sofre intensa pressão de fatores abióticos e antrópicos. Com cerca de 500 espécies conhecidas no bioma, as aves compõem um grupo biológico expressivo da Caatinga, com distintas linhagens evolutivas e características ecológicas. Este trabalho objetiva compreender os aspectos evolutivos e ecológicos da diversidade de aves da Caatinga e seus determinantes ambientais. Especificamente, buscamos (i) sumarizar o conhecimento existente sobre as assembleias de aves do bioma, suas relações filogenéticas e diferenças ecológicas (funcionais); (ii) investigar as inter-relações entre a diversidade taxonômica, filogenética e funcional, incluindo distintas medidas destas; (iii) avaliar os efeitos de diferentes gradientes ambientais (climáticos e estruturais) sobre essas diferentes dimensões da diversidade; e (iv) avaliar os efeitos desses fatores sobre a diversidade funcional de traços individuais. Reunimos informações de 123 assembleias de aves da Caatinga a partir da literatura disponível e utilizamos uma árvore filogenética das espécies e um conjunto de traços funcionais espécie-específicos para estimar a diversidade filogenética e funcional, respectivamente. Ainda, exploramos diferentes métricas de diversidade, a fim de entender suas inter-relações e selecionar aquelas menos enviesadas pela riqueza e esforço amostral. 523 espécies compuseram a diversidade regional da Caatinga. Os padrões de diversidade filogenética e funcional foram coincidentes quanto à distribuição espacial e a hipótese ambiental, tendo o clima atual como fator principal. Entretanto, a diversidade de traços funcionais individuais divergiu qualitativa e quantitativamente da diversidade funcional total. Esses resultados reforçam a necessidade de intensificar os esforços amostrais na Caatinga, permitem compreender o efeito das condições ambientais na formação das assembleias de aves em termos ecológicos e evolutivos, e por fim, alerta para o problema de selecionar traços ecológicos para descrever a diversidade funcional.