Os conceitos e métodos provindos das teorias de integração morfológica e de genética quantitativa formam o arcabouço teórico para o estudo da evolução de estruturas complexas, compostas de múltiplos caracteres que interagem entre si. Nesse trabalho, utilizamos o crânio como modelo de estrutura complexa e estudamos sua diversificação nas espécies de sapo do grupo Rhinella granulosa. As perguntas do trabalho foram: (1) A organização da (co)variação é similar entre as espécies?; (2) A organização da (co)variação é modular nas espécies, conforme expectativas baseadas em desenvolvimento ou função?; (3) Fatores externos, como filogenia e clima, estruturam a similaridade no padrão de covariação entre as espécies?; (4) A diversificação da morfologia média do crânio se deu por deriva ou seleção natural?; (5) A divergência na morfologia média do crânio está associada à variação climática entre as espécies?; e finalmente (6) Restrições evolutivas atuaram na divergência entre as espécies? Os espécimes foram escaneados e validamos o uso de imagens 3D para a mensuração de 21 distâncias lineares. Os crânios das espécies foram representados como matrizes fenotípicas (P) de covariância e de correlação entre as distâncias. A similaridade entre as P das espécies é alta. As P de todas as espécies se conformam a um padrão modular compatível com interações funcionais entre ossos. As diferenças entre as P concentram-se no rostro e são associadas a diferenças no clima entre as espécies. Detectamos sinal de seleção natural nos nós mais basais da filogenia e variação local no crânio está associada à variação na sazonalidade da chuva entre as espécies. Restrições evolutivas atuaram na diversificação do crânio das espécies, defletindo as respostas evolutivas para tamanho. Concluímos que tanto seleção estabilizadora e direcional, conectadas à variação climática, quanto restrições evolutivas atuaram na diversificação do crânio das espécies / Concepts and methods within the theories of morphological integration and quantitative genetics characterize the foundation to study the evolution of complex structures, composed of several traits that interact with each other. In this work, we used the skull as a model of complex structure and we studied its diversification in toad species belonging to the Rhinella granulosa group. The questions addressed were: (1) Is the (co)variance structure similar across species?; (2) Is the (co)variance structure modular in the species, and compatible with developmental or functional interactions among traits?; (3) Do external factors, such as phylogeny and climate, structure the similarity in covariance pattern across species?; (4) Was the diversification of skull mean morphology driven by drift or natural selection?; (5) Is skull divergence associated to climatic variation across species?; and finally, (6) Is there a role for evolutionary constraints in species skull divergence? We scanned all specimens and we validated the use of 3D images to measure 21 linear distances. The skull was represented as covariance and correlation phenotypic matrices (P) among distances. P similarity is very high among species. All species\' P had a modular pattern compatible with functional interactions among bones. Differences in P were concentrated in the snout and associated to differences in climate across species. We detected a selection signal in the three most basal nodes of the phylogeny and local variation in the skull is explained by between-species variation in precipitation seasonality. Evolutionary constraints played a major role in species skull diversification, biasing evolutionary responses towards the direction of size. We conclude that stabilizing and directional selection, connected to climatic variation, as well as evolutionary constraints, acted in species skull diversification

Os conceitos e métodos provindos das teorias de integração morfológica e de genética quantitativa formam o arcabouço teórico para o estudo da evolução de estruturas complexas, compostas de múltiplos caracteres que interagem entre si. Nesse trabalho, utilizamos o crânio como modelo de estrutura complexa e estudamos sua diversificação nas espécies de sapo do grupo Rhinella granulosa. As perguntas do trabalho foram: (1) A organização da (co)variação é similar entre as espécies?; (2) A organização da (co)variação é modular nas espécies, conforme expectativas baseadas em desenvolvimento ou função?; (3) Fatores externos, como filogenia e clima, estruturam a similaridade no padrão de covariação entre as espécies?; (4) A diversificação da morfologia média do crânio se deu por deriva ou seleção natural?; (5) A divergência na morfologia média do crânio está associada à variação climática entre as espécies?; e finalmente (6) Restrições evolutivas atuaram na divergência entre as espécies? Os espécimes foram escaneados e validamos o uso de imagens 3D para a mensuração de 21 distâncias lineares. Os crânios das espécies foram representados como matrizes fenotípicas (P) de covariância e de correlação entre as distâncias. A similaridade entre as P das espécies é alta. As P de todas as espécies se conformam a um padrão modular compatível com interações funcionais entre ossos. As diferenças entre as P concentram-se no rostro e são associadas a diferenças no clima entre as espécies. Detectamos sinal de seleção natural nos nós mais basais da filogenia e variação local no crânio está associada à variação na sazonalidade da chuva entre as espécies. Restrições evolutivas atuaram na diversificação do crânio das espécies, defletindo as respostas evolutivas para tamanho. Concluímos que tanto seleção estabilizadora e direcional, conectadas à variação climática, quanto restrições evolutivas atuaram na diversificação do crânio das espécies / Concepts and methods within the theories of morphological integration and quantitative genetics characterize the foundation to study the evolution of complex structures, composed of several traits that interact with each other. In this work, we used the skull as a model of complex structure and we studied its diversification in toad species belonging to the Rhinella granulosa group. The questions addressed were: (1) Is the (co)variance structure similar across species?; (2) Is the (co)variance structure modular in the species, and compatible with developmental or functional interactions among traits?; (3) Do external factors, such as phylogeny and climate, structure the similarity in covariance pattern across species?; (4) Was the diversification of skull mean morphology driven by drift or natural selection?; (5) Is skull divergence associated to climatic variation across species?; and finally, (6) Is there a role for evolutionary constraints in species skull divergence? We scanned all specimens and we validated the use of 3D images to measure 21 linear distances. The skull was represented as covariance and correlation phenotypic matrices (P) among distances. P similarity is very high among species. All species\' P had a modular pattern compatible with functional interactions among bones. Differences in P were concentrated in the snout and associated to differences in climate across species. We detected a selection signal in the three most basal nodes of the phylogeny and local variation in the skull is explained by between-species variation in precipitation seasonality. Evolutionary constraints played a major role in species skull diversification, biasing evolutionary responses towards the direction of size. We conclude that stabilizing and directional selection, connected to climatic variation, as well as evolutionary constraints, acted in species skull diversification

The Loricariinae belong to the Neotropical mailed catfish family Loricariidae, the mostspeciose catfish family in the world, and are united by a long and flattened caudal peduncle and the absence of an adipose fin. Despite numerous works conducted on this group, no phylogeny is presently available. Prior to conduct evolutionary studies, an exhaustive and robust phylogeny was reconstructed using mitochondrial and nuclear data. Then, this phylogeny was used in multivariate and multi-table analyses to reveal the main evolutionary trends of the subfamily. The resulting phylogeny indicated that the Harttiini tribe, as classically defined, formed a paraphyletic assemblage and was restricted to three genera, and within the Loricariini tribe, two sister subtribes were distinguished, Farlowellina and Loricariina, both displaying complex evolutionary patterns. In addition several new taxa were highlighted and described. Subsequently using this phylogeny as exploratory tool, we demonstrated: (1) using co-inertia analysis that the diagnostic features provided to define the different genera were phylogenetically dependent; (2) using multiple co-inertia analysis that the underlying evolutionary forces shaping their diversification included intraphenotypic (morphology and genetics) and extraphenotypic (ecology and distribution) components; (3) using the RLQ analysis that co-dispersion events occurred between co-distributed species responsible for the current fish distribution; and (4) using the multi-scale pattern analysis that the co-evolution in traits related to the mouth characteristics was linked to reproductive functions responsible for a tertiary evolution of this organ.

Les Loricariinae appartiennent à la famille des poissons-chats néotropicaux cuirassés Loricariidae, la famille de poissons-chats la plus riche en espèce au monde, et se caractérisent par un pédoncule caudal long et aplati et par l’absence de nageoire adipeuse. Préalablement aux études évolutives réalisées, une phylogénie exhaustive et robuste a été établie sur la base de données mitochondriales et nucléaires. Cette phylogénie a ensuite été utilisée dans des analyses multivariées et multi-tableaux afin de révéler les principales tendances évolutives de la sous-famille. La phylogénie obtenue indique que la tribu Harttiini forme un groupe paraphylétique et est restreinte à trois genres, et que dans la tribu Loricariini, deux sous-tribus soeurs se distinguent, les Farlowellina et les Loricariina, chacune présentant des patterns évolutifs complexes. Plusieurs nouveaux taxa ont aussi été mis en évidence et décrits. En utilisant la phylogénie comme outil exploratoire, nous avons démontré : (1) avec l’analyse de co-inertie que les caractères diagnostiques fournis pour définir les différents genres étaient sous dépendance phylogénétique ; (2) avec l’analyse de co-inertie multiple que les forces évolutives sous-jacentes dirigeant leur diversification incluaient des composantes intraphénotypiques (morphologie et génétique) et extraphénotypique (écologie et distribution) ; (3) avec l’analyse RLQ que des évènements de co-dispersion entre espèces codistribuées avaient eu lieu et étaient responsables de la distribution actuelle des espèces ; et (4) avec l’analyse de patterns multi-échelles que la co-évolution des traits liés aux caractéristiques de la bouche était liée à des fonctions reproductrices responsables d’une évolution tertiaire de cet organe. / The Loricariinae belong to the Neotropical mailed catfish family Loricariidae, the mostspeciose catfish family in the world, and are united by a long and flattened caudal peduncle and the absence of an adipose fin. Despite numerous works conducted on this group, no phylogeny is presently available. Prior to conduct evolutionary studies, an exhaustive and robust phylogeny was reconstructed using mitochondrial and nuclear data. Then, this phylogeny was used in multivariate and multi-table analyses to reveal the main evolutionary trends of the subfamily. The resulting phylogeny indicated that the Harttiini tribe, as classically defined, formed a paraphyletic assemblage and was restricted to three genera, and within the Loricariini tribe, two sister subtribes were distinguished, Farlowellina and Loricariina, both displaying complex evolutionary patterns. In addition several new taxa were highlighted and described. Subsequently using this phylogeny as exploratory tool, we demonstrated: (1) using co-inertia analysis that the diagnostic features provided to define the different genera were phylogenetically dependent; (2) using multiple co-inertia analysis that the underlying evolutionary forces shaping their diversification included intraphenotypic (morphology and genetics) and extraphenotypic (ecology and distribution) components; (3) using the RLQ analysis that co-dispersion events occurred between co-distributed species responsible for the current fish distribution; and (4) using the multi-scale pattern analysis that the co-evolution in traits related to the mouth characteristics was linked to reproductive functions responsible for a tertiary evolution of this organ.

<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>

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.