Perissodus microlepis is a species of cichlid fish endemic to Lake Tanganyika (Africa). Adult P. microlepis are lepidophages, feeding on the scales of other living fish. As an adaptation for this feeding behavior P. microlepis exhibit lateral asymmetry with respect to jaw morphology: the mouth either opens to the right or left side of the body. Field data illustrate a temporal phenotypic oscillation in the mouth-handedness, and this oscillation is maintained by frequency-dependent selection. To better understand the oscillation, Takahashi and Hori model frequency-dependent selection in P. microlepis using a population genetic model. Their results are intriguing, and the purpose of this dissertation is to improve and extend their model, which fails to account for important biological aspects.We model P. microlepis with a novel approach that fuses the disparate modeling traditions of population genetics and population dynamics; we account for both processes since, in the case of P. microlepis, they occur on the same time scale (a case of microevolution). We construct our models using systems of difference equations. We prove the existence and uniqueness of a positive equilibrium, which corresponds to a 1 : 1 phenotypic ratio. Using a local stability and bifurcation analysis, we show that the equilibrium becomes unstable when frequency-dependent selection is sufficiently strong. We determine necessary and sufficient conditions for onset of oscillation. Local bifurcation analysis indicates key features of the oscillation that may suggest critical experiments.We determine the role of stage structure and the role of strong and weak intraspecific competition. We show that stage-structure is not necessary for, but enhances, oscillatory behavior. Finally we demonstrate the complicated interplay between population dynamic and population genetic processes. Our findings indicate that classical population genetic models can fail to elucidate complex dynamics.

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Les signaux d'avertissement que portent les proies toxiques (ou autrement défendues) offrent une opportunité unique de développer une vision intégrative de la diversification biologique. Ces signaux sont soumis à une forte sélection naturelle et sexuelle. D'une part, la stratégie d'échantillonnage utilisée par les prédateurs, caractérisée par l'apprentissage des signaux associés à la toxicité, protège les signaux en forte fréquence dans la communauté de proies. Cette sélection fréquence-dépendante positive favorise l'uniformité des phénotypes et la convergence de signaux entre espèces toxiques (mimétisme mutualiste dit "Müllerien") dans de nombreux taxons. D'autre part, les signaux d'avertissement sont utilisés comme critères de choix entre partenaires sexuels et sont donc soumis à de la sélection sexuelle avec des conséquences importantes pour l'émergence de l'isolement reproducteur et la spéciation. Paradoxalement, malgré une forte sélection naturelle favorisant la convergence, les signaux d'avertissement sont incroyablement diversifiés, à la fois au sein de la même espèce et entre espèces. Cette diversification morphologique est souvent associée à une importante diversification des espèces à l'échelle des clades. Dans cette thèse, j'apporte quelques explications à ce paradoxe et j'affine ainsi notre compréhension de l'effet de la sélection fréquence-dépendante positive et du mutualisme sur la diversification à l'échelle micro- et macro-évolutive. Premièrement, je montre que la stratégie d'échantillonnage des prédateurs peut favoriser la diversification des signaux d'avertissement malgré une sélection fréquence-dépendante positive. Deuxièmement, je dissèque les conditions permettant l'évolution d'un isolement reproducteur stable, nécessaire à la spéciation, dans certaines situations écologiques où les signaux mimétiques sont sous sélection naturelle et sexuelle. Troisièmement, je décris des effets indirect de la sélection fréquence-dépendante sur la diversification à l'échelle macro-évolutive par le biais de contraintes spatiales et de convergences écologiques secondaires. / The display of warning signals by unpalatable (or otherwise defended) prey provides a wonderful opportunity for establishing an integrative view of biological diversification. Warning signals are known to be under strong natural and sexual selection. On the one hand, the sampling strategy of predators, characterized by a learned avoidance of signals associated with unpalatability, generates natural selection in favour of warning signals in high frequency in the prey community. Such positive frequency-dependent selection favours phenotypic uniformity and causes unpalatable species to converge on common warning signals (mutualistic "Müllerian" mimicry), as seen in a large panel of taxa. On the other hand, warning signals are used as a phenotypic cue for mate choice, generating sexual selection with important consequences for reproductive isolation and speciation. Paradoxically, despite powerful selection favouring phenotypic convergence, warning signals are fantastically diverse, both within and between species, and this morphological diversification is often associated with extensive species diversification at the clade level. In this thesis, I tackle this apparent paradox from the ground up and I thereby refine our understanding of the role of positive frequency-dependent selection and mutualistic interactions for evolutionary diversification at micro- and macroevolutionary scales. First, I show that the predator sampling strategy can favour the emergence of diversity of warning signals despite positive frequency-dependent selection. Second, I dissect the conditions allowing the evolution of strong and stable reproductive isolation, necessary for speciation to occur, in a number of ecological situations where warning signals are under natural and sexual selection. Third, I highlight important indirect effects of frequency-dependent selection on diversification at macro-evolutionary scale via spatial constraints and by-product ecological convergence.

The question of the tempo of evolution is amongst the oldest conundrums in evolutionary biology and has not been satisfactorily answered yet. One of the attempts to do so is the frozen plasticity theory, which postulates that a sexually reproducing species is only capable of evolution within short periods of time after its genetic polymorphism decreases e.g. as a consequence of peripatric speciation. In the longer periods of evolutionary stasis, its evolution is limited by frequency- dependent selection and pleiotropy. In this work, I have produced an open source software simulating the respon- ses of populations of sexually reproducing individuals to varying environmental conditions. Using this software, I simulated evolution of populations with different probabilities of arising of alelles affecting more phenotypic traits and frequency- dependent selected alleles that have opposing phenotypic effects when present at the locus in one copy, respectively two copies. I observed trends predicted by the theory of frozen plasticity: slower adap- tation to instant environmental changes, lower achieved fitness and more frequent extinctions of populations with higher portions of investigated types of alleles, but only with low effect sizes and without statistical significance. For future research, it would be desirable...

Fisher’s adaptive sex ratio theory predicts that organisms should invest equally in sons and daughters and the sex ratio at conception should be 1:1. Hamilton’s theory predicts that organisms should adjust sex ratios based on the relative strength of competition within a mating group. Testing sex ratio and sex allocation theories requires variation in sex ratio. Different sex allocation and sex allocation adjustment mechanisms can produce skewed sex ratios. I used Tigriopus californicus, a harpacticoid copepod with extrabinomial variation in sex ratios, to test sex ratio evolution and socially-mediated sex determination. Using artificially selected sex-biased populations, the trajectory of population sex ratios were as expected under Fisher’s theory and sex ratios approached/reached 0.5 proportion males. Populations with overlapping generations had a slower rate of change towards 0.5 than populations with non-overlapping generations. I show that these data are supported by multiple different models: a mechanistic and simulation model. I tested socially-mediated sex determination using seawater conditioned with different local sex ratios of copepods. There were detectable effects found in both wild populations and isofemale lines. However, these effects may be trivial as differences were small between treatments. Sex determination in T. californicus is a complex mechanism, with multiple genetic and environmental components. The complex nature of sex determination in T. californicus and the dynamic nature of their habitat in highly ephemeral splash pools provide a possible explanation for the non-Fisherian sex ratios we see. / Graduate

Minority phenotype preference, or the negative frequency-dependent selection is a type of selection, where a feature is more preferred, the lower it's frequency is in the population. Even a weak effect in other-preference based sexual selection can result in a sustainable polymorphism. This study reviews the phenomenon in the context of human visual facial features. Common trends in attractiveness shape the morphology of the human face towards uniformity. Individual recognition however, as a condition for most social relations, uses the wast variability of different features. This variability could be formed and maintained by minority phenotype preference. In our study we tested the effect of minority phenotype preference in the selection for rare hair and eye colors. In 120 unique sets, each containing six photographs, we experimentally manipulated the frequency of each color type. These sets were then shown to 226 human raters. We tested whether the relative frequency of each color type affected the rating of individual stimuli. In hair color, significant effect of minority phenotype preference was detected when females rated the photographs of men. When males rated the photographs of females, the effect was significant in eye color only. Key words: face perception, sexual selection,...

The human eye is a conspicuous and unique component of facial appearance. From other mammalian eyes it is set apart mainly by its unusual shape, visible white sclera, and a wide range of iris colours, which is something unique within a single species. Whether alone or in the context of the face as a whole, eye colour is an underexplored area within research into the perception of various personality traits. In this thesis, the author first reviews the physiological factors connected with eye colour and reported correlations between eye colour and different psychological and behavioural conditions. This is followed by an investigation of a possible relationship between eye colour and perceived dominance and attractiveness. The first and second study examines whether and to what extent eye colour is associated with facial morphology responsible for perceived dominance. The results are ambiguous. A cross-cultural comparison in the third study revealed that faces with blue eyes are judged as more attractive only in populations where individuals with darker eyes predominate. It is thus discussed whether this population-specific pattern is the consequence of a negative frequency-dependent selection that may have contributed to the present-day eye colour diversity. In short, the aim of this thesis was to...

There is repeated evidence that hybridization is a major contributor to the production of adaptive diversity; however, the evolutionary fate of hybrids in natural populations remains poorly understood. In Heliconius butterflies, hybridization is common and responsible for generating a variety of warning color patterns across the genus. Predator avoidance of warning colorations appears to largely be learned, which drives strong positive frequency-dependent selection. This creates a paradox for hybrid lineages: how do novel hybrid forms manage to establish and persist under such strong selection? In this dissertation, I present a series of studies centered on the selection dynamics of Heliconius hybrid zones, to elucidate how novel adaptive traits establish in nature. Clines across hybrid zones have often been analyzed to estimate selection on ecologically important loci. Here, warning color clines were characterized and compared across multiple transects along a Heliconius hybrid zone in the Guiana Shield. Furthermore, a mark-resight experiment and communal roost observations were completed near the center of this hybrid zone to determine the survival and likelihood of establishment of native and foreign forms. These studies reveal similar survivorship of hybrid and pure color patterns, and specifically demonstrate that a rare putative hybrid form can survive and establish within a hybrid zone. Both hybrids and pure color patterns showed comparable life expectancies in the mark-resight experiment and similar patterns of presence at nocturnal roosts. These results suggest that selection on warning color pattern is relatively weak within the hybrid zone. Analyses of color pattern clines uncovered strong selection bounding the hybrid zone in bi-race areas, while weaker selection was estimated for a tri-race area. In fact, the tri-race area was three times wider than the bi-race areas. Collectively, these studies suggest that the selection dynamics across hybrid zones may play an integral role in the establishment of new adaptive traits, and offers a route by which a reputed hybrid race may have arisen. The investigations within this dissertation also provide a new view of hybrid zone dynamics, and improve our understanding of how hybridization and selection shapes the evolution of biodiversity.

In this thesis, I combined comparative and experimental approaches to examine selection on reproductive traits and population differentiation in the insect-pollinated, self-incompatible, perennial herb Primula farinosa. More specifically, I (1) determined whether the effects of floral display and interactions with pollinators and seed predators, and plant reproductive success were frequency-dependent and affected by surrounding vegetation context, (2) examined the consequences of intermittent drought years on population dynamics using numerical simulations based on demographic data collected over seven years, (3) analyzed among-population differentiation in flowering phenology and reproductive allocation, and its relationship to soil-depth at the site of origin. A field experiment suggested that conspicuous plants facilitate inconspicuous plants in terms of pollinator attraction, and that the facilitation effect is contingent on the height of the surrounding vegetation. Further experiments revealed that both mutualistic and antagonistic interactions can result in frequency-dependent selection on floral display. Among inconspicuous plants, both fruit initiation, and damage from seed predators increased with the proportion of the conspicuous morph. The relative strength of these effects, and therefore their net outcome on the relationship between morph ratio and seed production varied among years. I combined information on vital rates and their relation to environmental conditions in simulations to predict future population viability in changing environments. Simulated stochastic population growth rate decreased with increasing frequency of drought years. Reproductive allocation varied significantly among populations both in the field and in a common-garden experiment, but was correlated with soil depth at the site of origin only in the field. The results suggest that among-population variation in reproductive effort in the field mainly reflects plastic responses to environmental conditions, and that this plasticity may be adaptive. The common-garden experiment suggested that the study populations have diverged genetically in flowering time.

La sélection fréquence-dépendante est un mécanisme d’évolution selon lequel l’aptitude d'un type varie en fonction de sa fréquence dans la population. Ce mécanisme joue un rôle important dans de nombreuses interactions autant interspécifiques (parasitisme, prédation, compétition), qu'intra-spécifiques entre les différents phénotypes d'une même espèce. La sélection fréquence-dépendante peut être positive ou négative et favoriser alors les phénotypes communs ou rares, respectivement. Elle a été mise en évidence dans le contexte du choix de partenaire chez plusieurs espèces, notamment chez certaines espèces d'insectes (ex.: demoiselles, drosophiles, cantharide de Pennsylvanie) et de poissons (ex.: guppys, xiphos), mais elle a été aussi récemment découverte chez l’humain. L'importance de la sélection fréquence-dépendante dans le choix de partenaire chez les espèces monogames reste tout de même peu explorée et cette étude vise à combler cette lacune en utilisant le diamant mandarin, un passereau monogame, comme modèle biologique. Nous avons étudié l'importance de ce mécanisme lorsqu'un trait est neutre et lorsque celui-ci constitue un indicateur de qualité. De plus, nous avons tenté de déterminer si la présence de rivales peut modifier la préférence initiale des femelles pour les phénotypes rares ou communs. / Frequency-dependant selection is an evolution mechanism in which the fitness of a type depend of its frequency in the population. This mechanism play an important role in several interspecific (parasitism, predation, competition) and intraspecific interactions within different phenotypes of a same species. Frequency-dependant selection can be positive or negative and favor, respectively, either common or rare phenotype. This selection has been found in context of mated choice of several species, especially in insects (damselfly, drosophila, soldier beetle) and fishes (guppy, swordtail) and it has even been recently discover in humans. The significance of frequency-dependent selection in mated choice of monogamous species is still less explored and this study used the zebra finches (a monogamous passerine) in order to explored this field. We studied the significance of this mechanism when a character is neutral and when it is rather a quality cue. Also, we attempt to establish if rivals' presence affect or modify the initial preference of females for common or rare phenotypes.