Divergent natural selection for use of locally abundant resources can lead to diversification within and across species. However, the consequences of divergent selection for phenotypic evolution also depend on the development of variation. Because relationships among traits such as shared developmental timing or common involvement in an organismal function can channel variation generated during development, these relationships strongly influence the direction of evolution.During development of the mammalian mandible multiple tissues of distinct developmental origins interact with inputs from the functioning of attached muscles to produce a cohesive and well integrated trait. In soricid shrews, part of the mandible matures late in ontogeny, coinciding with the onset of foraging. In this case, foraging-linked muscle activity should influence the development of the late maturing mandibular region. Here, I show that variation in this late ossifying region reveals the local functional requirements of the jaw and results in an opportunity to decouple internal and external sources of variation (developmental and environmental respectively) in the mandible. Capitalizing on this feature of the Sorex system, I empirically examined the historical persistence of internal and external patterns of variation, the consequences of variation patterning for ecological and morphological diversification across taxa, and differences between early and late ossifying regions in their contribution to local adaptation in mandible morphology.I found that the functional requirements of diet directed mandible development and determined species similarity in both mandible morphology and function. Timing of bone maturation determined the morphological effects of foraging-linked muscle activity, resulting in differential expression of adaptive variation in the late maturing region. Further, I found higher levels of interspecific variation in the late maturing region of the mandible, and showed that interspecific divergence in foraging morphology occurs along the lines delineated by epigenetic inputs of muscle on bone formation during late ontogeny within species. These findings indicate that differences in functional requirements are critical for divergence among taxa in this system. Further, these results suggest that, when external inputs into trait development are indicative of local functional requirements, the same epigenetic mechanism of development can generate diversity both within and among taxa.

Genitálias de machos tendem a apresentar maior elaboração e taxas de evolução mais altas que as de fêmeas, fenômeno cujos mecanismos associados ainda são pouco compreendidos. Nesta tese buscamos explorar tais mecanismos através da compreensão sobre os papéis funcionais e do desenvolvimento por trás da diversificação genital em Pentatomidae. No primeiro capítulo, investigamos o funcionamento das peças genitais externas em machos e fêmeas de espécies de diferentes linhagens, identificando grupos de estruturas que funcionam de forma mais integradas entre si. Mostramos que existem marcas funcionais únicas a algumas linhagens, mas por outro lado algumas características são conservadas e se repetem em todas as espécies. No segundo capítulo, inferimos uma filogenia para Pentatomidae usando dados moleculares e morfológicos, com enfoque na subfamília mais diversa e confusa taxonomicamente, Pentatominae. Identificamos diversas linhagens propostas na literatura com grande suporte, corroboramos a monofilia de Pentatomidae e a polifilia de Pentatominae. No terceiro capítulo, testamos a hipótese de que diferentes componentes genitais em fêmeas possuem maior restrição evolutiva por serem mais integradas durante o desenvolvimento, em comparação aos machos. Corroboramos esta hipótese, e sugerimos que as genitálias dos machos são evolutivamente moldadas pela forma como funcionam na cópula, enquanto a diversificação nas fêmeas é restringida por sua origem ontogenética. / Male genitalia tend to show greater elaboration and evolve faster in relation to females, a phenomenon whose underlying mechanisms are still poorly understood. In this thesis, we explore these mechanisms seeking to understand the functional and developmental roles in the diversification of genitalia in Pentatomidae. In the first chapter, we investigated the function of the external genital parts in males and females from different lineages, with the goal of identifying structures that work in a more integrated fashion. We showed that each species have unique functional signals, but some characteristics are more conserved and invariable. In the second chapter, we inferred a phylogeny to the Pentatomidae using morphological and molecular data, focusing on the most diverse and taxonomically confusing subfamily, Pentatominae. We recognized several lineages proposed on the literature with strong support, corroborate the monophyly of Pentatomidae and the paraphyly of Pentatominae. In the third chapter, we test the hypothesis that different genital components in females are more evolutionarily constrained for being more integrated during the development, comparing to males. We corroborated this hypothesis and suggest that male genitalia are evolutionarily shaped by their functional roles in copula, while the diversification in female genitalia is more constrained due to its ontogenetic origin.

The overarching goal of my dissertation is to explore the potential and limits of a trait-based approach to plant ecology. Together, the different studies presented here address two explicit and implicit foundational assumptions underpinning the trait-based approach: (1) that the correlation patterns and biological significance of traits transfer across scales and (2) that the phenotypic complexity of plants can accurately be synthesized into a few meaningful traits to study their ecology. Moreover, the last chapter focuses on a third key assumption: (3) that traits are strong predictors of plant performance (Shipley et al. In Press). I examine these assumptions by exploring multivariate patterns of phenotypic variation and integration across different ecological scales (e.g., individuals, populations, species) while explicitly considering the phenotypic complexity of trees, both in terms of their multidimensional and integrated nature. Two themes thus permeate this body of work: scales and phenotypic complexity. Much of what we know about the relationships among key traits comes from species-scale studies. Trait variation at smaller scales are often interpreted in the context of these interspecific relationships, but it is not clear that interspecific patterns observed at global scales apply to smaller scales. Moreover, although plants are complex, integrated organisms with intricate relationships among their traits, single traits are often studied and interpreted without considering the rest of the phenotype. Yet, examining individual traits outside of their phenotypic context might provide limited insight or be misleading. To address these shortcomings, this body of work examines multidimensional patterns of trait variation and correlation across ecological scales. It uses (1) a set of six ecophysiological leaf traits from mature trees in a lowland tropical rainforest, and (2) a set of twenty leaf, root, stem, branch and whole-plant ecophysiological traits from deciduous saplings in a temperate forest. The combination of our findings point to three main conclusions: (i) local interspecific and intra-population trait integration structures differ from each other and from the global interspecific patterns reported in the literature, such that global-scale interspecific patterns cannot readily be transferred to more local scales; (ii) considering the complexity of the plant phenotype provides better insights into ecological patterns and processes than what we can learn from considering individual or a handful of traits; and (iii) traits strongly affect individual plant performance, although there is no relationship between a species' trait correlation structure and its environmental niche, which suggests that there are multiple alternative optimal phenotypes in a given environment.

The balance between the costs and benefits of conspicuous signals ensures that the expression of those signals is related to the quality of the bearer. Plastic signals could enable males to maximize conspicuous traits to impress mates and competitors, but reduce the expression of those traits to minimize signaling costs, potentially compromising the information conveyed by the signals. I investigated the effect of signal enhancement on the information coded by the biphasic electric signal pulse of the gymnotiform fish Brachyhypopomus gauderio. Increases in population density drive males to enhance the amplitude of their signals. I found that signal amplitude enhancement improves the information about the signaler’s size. Furthermore, I found that the elongation of the signal’s second phase conveys information about androgen levels in both sexes, gonad size in males and estrogen levels in females. Androgens link the duration of the signal’s second phase to other androgen-mediated traits making the signal an honest indicator of reproductive state and aggressive motivation. Signal amplitude enhancement facilitates the assessment of the signaler’s resource holding potential, important for male-male interactions, while signal duration provides information about aggressive motivation to same-sex competitors and reproductive state to the opposite sex. Moreover, I found that female signals also change in accordance to the social environment. Females also increase the amplitude of their signal when population density increases and elongate the duration of their signal’s second phase when the sex ratio becomes female-biased. Indicating that some degree of sexual selection operates in females. I studied whether male B. gauderio use signal plasticity to reduce the cost of reproductive signaling when energy is limited. Surprisingly, I found that food limitation promotes the investment in reproduction manifested as signal enhancement and elevated androgen levels. The short lifespan and single breeding season of B. gauderio diminishes the advantage of energy savings and gives priority to sustaining reproduction. I conclude that the electric signal of B. gauderio provides reliable information about the signaler, the quality of this information is reinforced rather than degraded with signal enhancement.

Le succès écologique des organismes dépend principalement de leur phénotype. Une composante important du phénotype est la morphologie fonctionnelle car elle influence la performance d’un organisme donné dans un milieu donné et donc reflète son écologie. Des disparités dans la morphologie fonctionnelle ou dans le développement entre espèces peuvent donc mener à des différences écologiques. Ce projet évalue le rôle des mécanismes de variation morphologique dans la production de différences écologiques entre espèces au sein des poissons hybrides du complexe Chrosomus eos-neogaeus. En utilisant la microtomodensitométrie à rayons X et la morphométrie géométrique 3D, la forme des éléments des mâchoires est décrite pour comparer la variation morphologique et les différences développementales entre les membres du complexe C. eos neogaeus. Les hybrides présentent autant de variation phénotypique que les espèces parentales et présentent des phénotypes nouveaux, dit transgressifs. Les hybrides présentent aussi des différences marquées avec les espèces parentales dans leur allométrie et dans leur intégration phénotypique. Finalement, ceux-ci semblent être plastiques et en mesure de modifier leur phénotype pour occuper plusieurs environnements. L’entièreté de ces résultats suggère que des changements dans le développement des hybrides entraînent une différenciation phénotypique et écologique avec les espèces parentales. / An organism’s success is highly dependent on its phenotype. A major component of the phenotype is functional morphology because of its role in determining performance in a given environment clues as to their ecology. Morphological disparity and developmental differences between species can thus lead to ecological differences. This project assesses the role that mechanisms capable of generating morphological variation have in producing ecological differences between species within the hybrid fishes of the Chrosomus eos-neogaeus complex. Using X-ray based computer-aided micro-tomography and 3D geometric morphometrics, the shape of bones composing the jaws was described to measure morphological variation and developmental differences in the C. eos-neogaeus complex. Chrosomus eos-neogaeus hybrids displayed as much variation as parental species and novel, deemed transgressive, phenotypes. Hybrids were also markedly different from parental species in their patterns of allometric scaling and phenotypic integration. Finally, hybrids appeared to be plastic and able to acclimate to different environments. The results of this study indicate that changes in the development of hybrids could lead to phenotypic and, in turn, ecological differences with parental species.

Le potentiel adaptatif d’une espèce peut être défini par sa capacité à faire face aux changements environnementaux. Le potentiel adaptatif augmente avec la variation du phénotype, du niveau intra-individuel au niveau inter-populations. Cependant, les facteurs qui contrôlent et expliquent cette variation phénotypique sont encore relativement peu compris. Nous avons étudié quatre espèces végétales des îles Kerguelen, en subantarctique, une des régions les plus sévèrement affectée par le changement climatique. Les espèces végétales des îles Kerguelen montrent un fort degré d’intégration phénotypique (i.e. forte corrélation entre les traits), suggéré comme étant une contrainte pour la variation des traits. Chez ces espèces, nous avons étudié les facteurs qui peuvent contraindre la variation phénotypique : les facteurs environnementaux extrinsèques, l’intégration phénotypique intrinsèque et les coûts associés de performance. Nous avons montré que la variation intra-individuelle, i.e. plasticité, peut être contrainte par la modification simultanée de multiple facteurs environnementaux, et par les coûts de performance qui en résultent. En revanche, la plasticité peut être favorisée par un fort degré d’intégration phénotypique (Chapitre 3). Nous avons montré que la variation interindividuelle, à l’intérieur des populations, n’était pas contrainte par des facteurs environnementaux, mais pouvait cependant être favorisée par un fort degré d’intégration phénotypique (Chapitre 1). Egalement, nous avons trouvé que la variation inter-populations à l’intérieur d’une région peut être contrainte par une variation environnementale limitée (Chapitre 1). Finalement, nous avons étudié des métabolites secondaires (amines et flavonols) qui font le lien entre variation environnementale et variation phénotypique. Nous avons trouvé que la composition et la fonction de ces métabolites varient entre régions, suggérant une différentiation évolutive entre régions (Chapitre 2). Les patrons de variation entre régions, suggèrent au niveau intra-spécifique une redondance et une versatilité fonctionnelle des métabolites, que nous somme, à notre connaissance, les premiers à mettre en évidence. Nous suggérons que le changement climatique des îles Kerguelen va avoir un impact négatif sur la performance des espèces végétales. La persistance d’habitats humides favorables à ces espèces sera alors un facteur déterminant de la capacité des espèces à faire face au changement climatique. De plus, ce projet a permis d’identifier des facteurs jusqu’alors peu reconnus qui pourtant favorisent le potentiel adaptatif des espèces. En particulier, le potentiel adaptatif peut être favorisé par (i) le degré d’intégration phénotypique (contrairement à ce qui est communément suggéré) et (ii) la redondance et la versatilité des métabolites (qui a peu été étudiée jusqu’alors). Qui plus est, nous avons mis en évidence pour la première fois, plusieurs coûts et limites de la plasticité, suggérant qu’une réponse plasticité de la plante n’est pas une garantie de succès dans le nouvel environnement. / The adaptive potential of a species can be defined as its capacity to cope with environmental change. Adaptive potential increases with phenotypic variation, from the intra-individual to the inter-population level, but factors controlling and explaining this variation still remain poorly understood.We studied four plant species from Iles Kerguelen in the sub-Antarctic region which is currently facing one of the strongest climate changes worldwide. Plant species from Iles Kerguelen are known to show high phenotypic integration (i.e. strong correlation among traits), a phenomenon that has been suggested to constrain trait variation. For these species we studied what constrains phenotypic variation, considering the external environment, the internal phenotypic integration and the associated performance costs. We found that intra-individual variation, i.e. plasticity, may be constrained by complex environmental change and the performance costs it triggers. In contrast, plasticity may be favored by high degree of phenotypic integration (Chapter 3). We found that inter-individual variation within populations may not be constrained by environmental factors, but may be favored by high phenotypic integration (Chapter 1). We found inter-population variation within regions may be constrained by restricted environmental variation (Chapter 1). Finally, we studied secondary metabolites (amines and flavonols) that connect environmental variation to phenotypic variation. We found that compositions and functions of these metabolites vary among regions, probably reflecting evolutionary differentiation among regions (Chapter 2). Patterns of variation betweenregions suggest that within species metabolites may be functionally redundant or versatile, for which to our knowledge our results are the first hint. Overall, we suggest that climate change in Kerguelen will impact plant species performance, and that the persistence of suitable wet habitats will be determinant in species capacities to cope with such changes. Furthermore, this project identified so far underestimated factors which may favor the adaptive potential of species. Particularly, we emphasize that the adaptive potential of species may increase due to (i) phenotypic integration, (contrary to common suggestion) and (ii) metabolite redundancy or versatility (only poorly studied so far). Moreover, we evidenced, partly for the first time, multiple costs and limits of plasticity and suggest that plasticity does not guarantee plant success in the new environment.

The aim of the present work was to investigate the impact of the varying environmental conditions on the taxonomic and phenotypic diversification of a geographically widespread and ecologically successful Old World primate genus, the macaques (Cercopithecidae: Macaca). To this end, the relationship between geography, ecology, phylogeny, and phenotypic variation among macaques was investigated. Constraints to phenotypic variation – and thus evolution – were also analysed in the form of observed amounts of phenotypic variation and patterns of phenotypic integration. A total of 72 standard linear measurements of teeth and associated cranial and mandibular structures were taken for a total sample of 744 specimens from 13 species of macaques. Climate and ecological data were collated from the literature. Univariate and multivariate statistics were employed for the analysis. Patterns of variation, covariation, and allometry were analysed in the dentition, both within and between species. The ecogeographical analysis was carried out by means of two-block partial least squares and a type of multivariate regression, both in a phylogenetic framework. Phylogenetic signal was tested for by means of Blomberg’s K. Macaque teeth differ in their variability. All teeth covary with each other, although correlations are strongest within tooth classes. Size was a strong contributing factor to dental integration, as evinced by lower correlations between teeth once allometric effects were removed. Integration patterns also showed modularity between the anterior and the posterior dentition. Between-species variation in overall craniodental size was associated with temperature, latitude, and body size. Species also varied, albeit to a lesser degree, along an antero-posterior contrast in relative tooth size. Larger anterior were found to be associated with frugivory and tropical ecology, whereas a larger posterior dentition was linked to a more folivorous diet and temperate environments. The latter pattern was largely a function of phylogenetic relatedness. Phylogenetic signal was generally strong in the dentition, although it was substantially greater in the anterior teeth (incisors and canines) than in the posterior teeth (premolars and molars). Macaques show adaptive differentiation in body size in response to temperature along a latitudinal cline, corroborating the presence of the Bergmann effect in macaques. There was no conclusive support for further adaptive differentiation, despite an association between relative tooth size and diet. Allometry appears to channel evolutionary divergence of macaques along a line of least evolutionary resistance, and developmental modularity allows for partly uncoupled evolution of the anterior and posterior dentition. Future research should be aimed at broadening the taxonomic scope to include craniodental variation of the African papionins and cercopithecins in order to put the observed macaque patterns in a broader evolutionary context.