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Le « masting » correspond à une dynamique de fructifications, commune chez de nombreuses plantes pérennes, et caractérisée par une production de fruits extrêmement fluctuante d’une année à l’autre et synchronisée à l’échelle populationnelle. Il a un impact important sur la démographie des populations de consommateurs de fruits, et par effet de cascade, sur l’ensemble de la dynamique des écosystèmes forestiers. Notre méconnaissance actuelle des causes proximales du « masting » empêche toute prédiction crédible sur la fréquence et l'intensité des fructifications, et sur leurs conséquences au niveau des écosystèmes forestiers, dans le contexte du changement climatique. Cette thèse vise à tester l’hypothèse selon laquelle le processus de pollinisation pourrait jouer un rôle clé dans le « masting » des chênes de région tempérée (Quercus petraea et Q. robur), ce qui pourrait le rendre extrêmement sensible au changement climatique. En combinant une approche empirique multi-sites à large échelle spatio-temporelle en France et une approche théorique basée sur l’utilisation d’un modèle mécaniste, j’ai montré que (i) la dynamique des fructifications est liée à la disponibilité en pollen pour la reproduction. Le pollen peut être limitant certaines années en raison d’une faible quantité de pollen produite ou de conditions météorologiques défavorables à l’émission et à la diffusion du pollen ; (ii) la phénologie pollinique est un caractère clé du « masting » : l’émission pollinique a lieu au début du printemps, dans des conditions météorologiques souvent défavorables à la pollinisation, ce qui conduit à de fréquents échecs de la fructification et explique le caractère rare et imprévisible des fructifications massives ; (iii) le « masting » deviendrait moins stochastique avec l’augmentation des températures printanières au cours des prochaines décennies, ce qui pourrait avoir des conséquences importantes sur la dynamique des consommateurs de fruits, et par effet de cascade, sur la capacité de régénération des chênaies / “Masting” designates a fruiting dynamics common in many perennial plants and characterized by seed production that is highly variable over the years and strongly synchronized among trees within populations. It is expected to strongly impact the population demography of seed consumers, and by cascade effect, the dynamics of forest ecosystems as a whole. Our lack of knowledge of the proximate causes of “masting” currently prevents any reliable prediction about the frequency and the intensity of fruiting, or about their consequences, under climate change. In this work I aimed to test the hypothesis that the pollination process is playing a key role in “masting” in temperate oak forests (Quercus petraea and Q. robur), which would make masting highly sensitive to climate change. By combining an empirical multi-site approach at large spatio-temporal scale in France and a theoretical approach based on a mechanistic model, I found that (i) fruiting dynamics depends on the annual amount of airborne pollen available for reproduction. This amount could be limited some years due to either low amounts of pollen produced at the population level, or harsh weather conditions affecting pollen release and aerial diffusion; (ii) pollen phenology is a key character of “masting”: pollen release takes place in early spring when weather conditions are typically unfavorable to pollination, which leads to frequent fruiting failure and explains thereby why mast years are rare and unpredictable; (iii) “masting” should become less stochastic in the upcoming decades because of the increase of spring temperatures, which should markedly influence the dynamics of seed consumers, and by cascading effect, oak forest regeneration

<p>In this thesis, quantitative genetic approaches, common-garden experiments, and field studies were combined to examine patterns of population differentiation and the genetic architecture of characters of putative adaptive significance in the widely distributed perennial herb Lythrum salicaria. In this work, I (1) documented patterns of population differentiation in phenology, life-history, and morphology along latitudinal gradients at different geographical scales, (2) investigated the genetic architecture of phenology, flower morphology, and inflorescence size, and (3) combined estimates of phenotypic selection in the field with information on the genetic variance-covariance matrix (G) to examine potential constraints to adaptive evolution. </p><p>A common-garden experiment demonstrated latitudinal variation in life-history, and phenology of growth and reproduction among L. salicaria populations sampled across Sweden. Flower morphology varied significantly among populations, but was, with the exception of calyx length, not related to latitude of origin. A second experiment, which included two Swedish, two Dutch, and two Spanish populations, indicated that the latitudinal gradient in reproductive and vegetative phenology might extend throughout Europe.</p><p>A quantitative-genetic study of two Swedish populations revealed significant additive genetic variation for all phenological and morphological traits investigated. The G matrices of the populations differed significantly according to common principal component analysis, and genetic correlations within the study populations did not strictly correspond to trait correlations observed among populations.</p><p>In a field study, I detected directional selection through female function for larger inflorescences in two consecutive years. Relative fitness increased disproportionately with inflorescence size in the year when supplemental hand-pollination indicated that pollen limitation was severe. Genetic correlations with inflorescence size considerably influenced predicted response to selection in other characters.</p><p>Taken together, the results suggest that among-population differences in phenology and life-history in L. salicaria have evolved in response to latitudinal variation in length of the growing season. They demonstrate that the evolutionary potential of local populations may be considerable. The genetic covariance structure substantially influences predicted short-term evolutionary trajectories. However, the weak correspondence between genetic correlations documented within populations and trait correlations among populations, suggest that the G matrix has not imposed strict constraints on patterns of among-population differentiation.</p>

<p>I combined comparative and experimental studies in the field and in the greenhouse to examine factors influencing reproductive success in two insect-pollinated herbs, the tristylous, selfincompatible perennial <i>Lythrum salicaria</i> and the self-compatible, biennial <i>Pedicularis palustris</i>. More specifically, I explored (i) the effects of plant population characteristics on the intensity and outcome of interactions with pollinators and seed predators, (ii) whether flower morphology affects duration of stigma receptivity, and (iii) whether damage-induced reduction in floral display reduces pollinator visitation and increases pollen limitation.</p><p>As predicted, the rate of pollinator visitation tended to increase and the number of flowers probed per plant tended to decrease with increasing population size in <i>L. salicaria</i>, but these relationships only approached statistical significance. By taking advantage of the pollen size polymorphism that is typical of many heterostylous plants, I could show that the number of compatible pollen grains received increased with population size, and that this was associated with a reduction in pollen limitation and increased seed output per flower. The deposition of high numbers of incompatible conspecific and heterospecific pollen grains did not appear to reduce seed set.</p><p>In <i>P. palustris</i>, fruit set and seed predation varied markedly among populations and years, but this variation could only partly be explained by variation in population size. Fruit set was positively related to population size, and seed predation was negatively related to population size, in one of three years. Similarly, the level of pollen limitation, which was quantified in two years, varied among populations, but was not related to population size, density or isolation.</p><p>In <i>L. salicaria</i>, both the duration of stigma receptivity and the effect of prior self-pollination on seed output varied among style morphs. These differences may contribute to morph-specific differences in pollen limitation and seed production documented in the field.</p><p>The results of a field experiment demonstrated that damage to the shoot apex may markedly reduce fruit production in <i>L. salicaria</i>, and suggested that this is mainly because damage reduces flower number. I found no evidence that a reduced floral display increased the severity of pollen limitation.</p><p>Taken together, the results show that interactions with both pollinators and herbivores may markedly affect reproductive output in the two plant species, and that the intensity of both mutualistic and antagonistic interactions vary considerably in space and time. Moreover, they are consistent with the hypothesis that pollination success should depend less on population size in selfcompatible than in self-incompatible plants.</p>

In this thesis, quantitative genetic approaches, common-garden experiments, and field studies were combined to examine patterns of population differentiation and the genetic architecture of characters of putative adaptive significance in the widely distributed perennial herb Lythrum salicaria. In this work, I (1) documented patterns of population differentiation in phenology, life-history, and morphology along latitudinal gradients at different geographical scales, (2) investigated the genetic architecture of phenology, flower morphology, and inflorescence size, and (3) combined estimates of phenotypic selection in the field with information on the genetic variance-covariance matrix (G) to examine potential constraints to adaptive evolution. A common-garden experiment demonstrated latitudinal variation in life-history, and phenology of growth and reproduction among L. salicaria populations sampled across Sweden. Flower morphology varied significantly among populations, but was, with the exception of calyx length, not related to latitude of origin. A second experiment, which included two Swedish, two Dutch, and two Spanish populations, indicated that the latitudinal gradient in reproductive and vegetative phenology might extend throughout Europe. A quantitative-genetic study of two Swedish populations revealed significant additive genetic variation for all phenological and morphological traits investigated. The G matrices of the populations differed significantly according to common principal component analysis, and genetic correlations within the study populations did not strictly correspond to trait correlations observed among populations. In a field study, I detected directional selection through female function for larger inflorescences in two consecutive years. Relative fitness increased disproportionately with inflorescence size in the year when supplemental hand-pollination indicated that pollen limitation was severe. Genetic correlations with inflorescence size considerably influenced predicted response to selection in other characters. Taken together, the results suggest that among-population differences in phenology and life-history in L. salicaria have evolved in response to latitudinal variation in length of the growing season. They demonstrate that the evolutionary potential of local populations may be considerable. The genetic covariance structure substantially influences predicted short-term evolutionary trajectories. However, the weak correspondence between genetic correlations documented within populations and trait correlations among populations, suggest that the G matrix has not imposed strict constraints on patterns of among-population differentiation.

I combined comparative and experimental studies in the field and in the greenhouse to examine factors influencing reproductive success in two insect-pollinated herbs, the tristylous, selfincompatible perennial Lythrum salicaria and the self-compatible, biennial Pedicularis palustris. More specifically, I explored (i) the effects of plant population characteristics on the intensity and outcome of interactions with pollinators and seed predators, (ii) whether flower morphology affects duration of stigma receptivity, and (iii) whether damage-induced reduction in floral display reduces pollinator visitation and increases pollen limitation. As predicted, the rate of pollinator visitation tended to increase and the number of flowers probed per plant tended to decrease with increasing population size in L. salicaria, but these relationships only approached statistical significance. By taking advantage of the pollen size polymorphism that is typical of many heterostylous plants, I could show that the number of compatible pollen grains received increased with population size, and that this was associated with a reduction in pollen limitation and increased seed output per flower. The deposition of high numbers of incompatible conspecific and heterospecific pollen grains did not appear to reduce seed set. In P. palustris, fruit set and seed predation varied markedly among populations and years, but this variation could only partly be explained by variation in population size. Fruit set was positively related to population size, and seed predation was negatively related to population size, in one of three years. Similarly, the level of pollen limitation, which was quantified in two years, varied among populations, but was not related to population size, density or isolation. In L. salicaria, both the duration of stigma receptivity and the effect of prior self-pollination on seed output varied among style morphs. These differences may contribute to morph-specific differences in pollen limitation and seed production documented in the field. The results of a field experiment demonstrated that damage to the shoot apex may markedly reduce fruit production in L. salicaria, and suggested that this is mainly because damage reduces flower number. I found no evidence that a reduced floral display increased the severity of pollen limitation. Taken together, the results show that interactions with both pollinators and herbivores may markedly affect reproductive output in the two plant species, and that the intensity of both mutualistic and antagonistic interactions vary considerably in space and time. Moreover, they are consistent with the hypothesis that pollination success should depend less on population size in selfcompatible than in self-incompatible plants.

Flowering plants display remarkable phenotypic diversity, especially in reproductive structures, much of which is thought to be associated with pollination by animals. Pollination syndromes are collections of floral traits (e.g. flower colour, shape, odour) that are associated with a plant attracting particular functional groups of animal pollinators. We explored the extent to which traits associated with the moth pollination syndrome translated into pollination by moths in the Pacific coast dune endemic Abronia umbellata and found mixed results: in one year of study, there was no difference in seed set by day- vs. night-pollinated inflorescences, but in another year of study, night-pollinated inflorescences set significantly more seed than those pollinated during the day. We integrate this work with tests of pollen and resource limitation of seed production and with seed set surveys of natural populations to address proximate and ultimate causes of low seed set, finding low rates of pollinator visitation, high pollen limitation of seed production in all populations studied, and no evidence of endogenous resource limitation of seed production. We propose that “excess” flowers may be functionally male, serving to increase outcross siring success. The transition from self-incompatibility and obligate outcrossing to self-compatibility and predominant selfing is the most common evolutionary transition among the flowering plants and traits associated with outcrossing may become reduced across such shifts, potentially through the action of natural selection, especially if pollinators are also herbivores, or if the signals that pollinators use to locate flowers are also used by herbivores. We examined the reduction of attractive visual and olfactory floral traits in A. umbellata across a shift from outcrossing to selfing and found a reduction of all floral traits considered. We found that floral volatile emissions were reduced more strongly than flower size or floral display (number of flowers per inflorescence), but there was no evidence of an ecological cost associated with conspicuousness: we did not find reduced leaf herbivory among selfers relative to outcrossers. / Thesis (Master, Biology) -- Queen's University, 2012-08-30 19:46:23.663

There is concern that climate change may cause mismatches between timing of flowering and activity of pollinators (phenology). However, concluding that mismatches will occur, and have serious consequences for pollination services, requires assumptions that have not yet been tested. I begin by discussing a set of these assumptions, bringing past research into the context of mismatch. Briefly, the assumptions are that 1) dates of first-flowering or emergence (DFFE) correctly describe phenology (and therefore mismatch); 2) differences in DFFE represent the magnitude of mismatch; 3) advancement of DFFE will be the primary phenological change; 4) shifts will be random and independent for each species; 5) populations of plants and pollinators are “bottom-up” regulated by their mutualistic interactions; 6) all interactions are of similar strength and importance; 7) dispersal, and the spatial context of phenological mismatches can be ignored; and ecological processes including 8) phenotypic plasticity and adaptive evolution of phenology, 9) competition and facilitation, and 10) emergence of novel interactions, will not affect mismatches. I then describe novel experiments, which could help to account for some of these assumptions, clarifying the existence and impacts of mismatches. Next, I present an original field experiment on factors affecting seed set in an alpine meadow in the Coast Range of British Columbia, Canada. I found evidence contradicting the assumption that seed set is primarily limited by pollination. My data highlight the roles of phenology, temperature (degree-days above 15°C, and frost hours), and interactions with pollinators (mutualists) and seed-predators (floral antagonists) in driving patterns of seed set. Seed set of early and late-flowering species responded differently to a 400m elevation gradient, which might be explained by phenology of bumble bees. My data suggest that the consequences of mismatch may be smallest for plants that are fly-pollinated and self-fertile. Non-selfing, bee-pollinated species might be more prone to reproductive limitation through mismatch (affected by snowmelt and cumulative degree-days). Plants that are limited by seed-predators might be negatively affected by warming temperatures with fewer frost hours, and extreme events such as late-season frosts and hail storms can prevent plants from setting seed entirely. Overall, my work emphasizes the importance of complementing theory, data-driven simulations, and meta-analyses with experiments carried out in the field. / Graduate

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