With close to 2100 species, the flowering plant genus Carex (Cyperaceae; sedges) is an example of an evolutionary radiation. Despite its potential for use as a model taxon in evolutionary studies, the diversification of sedges remains largely unexplored. This thesis realizes the potential of Carex as an evolutionary model group by using it to ask questions about species richness patterns. More specifically, it seeks to determine the relationship, if any, between rates of trait evolution and species richness. This tests the hypothesis that organisms with increased abilities to evolve new traits, speciate more rapidly. Morphological and ecological (habitat and climatic niche) traits are modelled on a nearly complete regional (North America north of Mexico) phylogeny and rates of trait evolution are compared among non-nested sister groups. However, before trait evolution is modelled, this work evaluates the sensitivity of climatic niche estimates to underlying distribution datasets. It tests the agreement of niche estimates derived from the commonly used online repository GBIF (the Global Biodiversity Information Facility) and county-level distributions via BONAP (the Biota of North America Program). Results showed that in the context of phylogenetic comparative analyses, it is not vital to obtain highly accurate climatic niche estimates. The second study found significant positive correlations between the rates of climatic niche, habitat and reproductive morphological evolution and species richness. This result supports the role of high trait lability in generating species richness and more generally, the idea that high trait disparity through evolutionary time leads to species success.

The aim of this dissertation is to study the systematics and evolution of Euphorbiaceae vines (tribe Plukenetieae), a diverse pantropical lineage (~365 species and 18 genera) composed of three morphologically distinct subtribes, Dalechampiinae, Plukenetiinae, and Tragiinae. Through the course of my research I largely resolved the evolutionary history of Plukenetieae and made broader contributions to the study of pollen and seed evolution, pantropical biogeography, and plant diversification. In chapter two I developed the first well-sampled molecular phylogeny for Plukenetieae (154 terminals, ~93 species, 2,207 character dataset composed of ITS and psbA-trnH with indel gap-scored data), and determined baseline species group relationships of the tribe. Molecular phylogeny largely agreed with pollen morphology hypotheses and confirmed that the large genus Tragia was para- and/or polyphyletic and should be split into smaller genera. Analysis of pollen morphology revealed a trend towards aperture reduction and loss in Tragiinae, with four origins of weakly defined apertures and up to three origins of inaperturate pollen. In chapter three, I studied the seed size evolution of Plukenetia, a pantropical genus with large edible oil-rich seeds, by developing a near-exhaustive phylogeny (83 terminals, 20 of ~24 species, 5,069 bp dataset of ETS, ITS, KEA1 introns 11 and 17, TEB exon 17, matK, ndhF) and conducting ancestral state estimation and phylogenetic regression. Seed size evolution in Plukenetia was dynamic and associated with competing selective pressures of plant size, fruit type (and inferred dispersal syndrome), and seedling ecology. In chapter four I presented a revised sectional classification of Plukenetia based on phylogeny and morphological evidence, including three new taxa from South America. Chapters three and five included biogeographical investigations on Plukenetia and Plukenetieae. Analyses revealed that pantropical disjunct distributions arose one to three times in each subtribe via periodic long-distance dispersals from the Oligocene to the Pliocene, most often from South America to Africa and then Southeast Asia. Lastly, in chapter five, I developed an improved phylogeny for Plukenetieae (289 terminals, ~109 species, 5,160 bp dataset of ETS, ITS, KEA1 intron 11, TEB exon 17, matK, ndhF) to study the influence of innovative traits (twining growth form, stinging hair defences, and pseudanthial inflorescences) on diversification in the tribe. However, increased diversification was not associated with innovative traits. Instead, diversification was associated with clades that shifted into drier open habitats, aided by habitat expansion following the Late Miocene cooling period.

The morphologies and behaviors of animals evolve and diversify, filling ecological niches in their environments. In this study I examine how a morphological trait, body mass, and three ecological traits, namely diel activity patterns, diving/non-diving locomotion, and diet, evolve in the Anseriformes (waterfowl). Through ancestral state reconstructions using a maximum likelihood approach the evolution of these traits was compared to see if any patterns of trait coevolution emerged. Body mass was compared to each ecological trait using a phylogenetic ANOVA to test if there were body size differences between ecological groups. The pattern of male and female body mass evolution across the clade was found to be remarkably similar, indicating that selection effected body mass in similar ways between the sexes. Diving locomotion appears to be the ancestral state for Anseriformes with non-diving independently evolving probably five times. The ancestral state of diet appears to be either herbivory or omnivory, with carnivory secondarily evolving twice independently. For diel activity patterns, the ancestral state reconstruction showed little resolution at the internal nodes, indicating the high degree of plasticity in this trait among the species studied. Body mass in both males and females was not significantly correlated with any particular diet, diving locomotion, or diel activity pattern.

Studies assessing the evolution of plant traits frequently focus on pollinators as the primary drivers of floral trait evolution. However, herbivores can also play a role, and, under some circumstances, may even impose stronger selection on floral display than pollinators. This is especially true when the traits under selection are linked to anti-herbivore defense. Here I describe a study in which I quantified the relative role of herbivores and pollinators in selection for floral traits in the North American invasive plant, purple loosestrife (Lythrum salicaria). Because L. salicaria responds to leaf-chewing herbivores by producing compensatory tissue growth, and this in turn alters the architecture of the floral display, I further tested whether herbivores can indirectly modify pollinator-mediated selection through this pathway. Using a split-plot design, I measured pollen limitation and reproductive output in experimentally manipulated plants in the presence and absence of simulated herbivore damage in order to quantify the effects of damage and pollination on natural selection for floral display. My results showed that damage significantly increased direct selection (beta-i) for earlier flowering time and decreased selection on the number of inflorescences, even more than pollinators did. Because damage did not modify pollinator-mediated selection for floral display traits, the selection imposed by herbivores is likely only having direct effects. My findings demonstrate the importance of considering multiple agents of selection and their potential interactions when quantifying natural selection in a study system. In particular, it is important to consider that the agent most frequently studied may not be imposing the brunt of selection.

Compreender os fatores que afetam a distribuição das espécies tem sido um dos principais objetivos dos ecólogos. Atualmente, sabe-se que os processos ecológicos e evolutivos moldam a dinâmica de especiação e extinção de espécies, e determinam a distribuição e abundância das mesmas. Ao longo dos últimos anos, tem havido um aumento no número de estudos que utilizam informação filogenética para explicar as dinâmicas populacionais e as distribuições de espécies, e que buscam identificar os mecanismos responsáveis pela montagem das comunidades. Interações das espécies, sejam elas intraespecífica, interespecífica ou com o ambiente, ocorrem baseadas nas diferenças e semelhanças fenotípicas. Essas variações fenotípicas tem origem na evolução das espécies, e com isso espera-se que as espécies proximamente relacionadas tendam a ser ecologicamente mais semelhantes entre si do que as espécies distantemente relacionadas. Esta concepção tem dado origem a um conceito importante, com implicações para estudos tanto ecológicos quanto evolutivos: o conceito de conservação filogenética de nicho, isto é, quando as espécies relacionadas mantêm seus nichos ancestrais ao longo do tempo evolutivo. Esse padrão tem importância para diversas áreas de ecologia, permitindo a ligação das espécies aos processos ecológicos e auxiliando na maior compreensão da ecologia evolutiva das diferentes linhagens. Devido à sua importância, é fundamental o desenvolvimento de métodos estatísticos adequados para quantificar esses padrões e inferir os processos que o subjazem. Atualmente, os métodos utilizados para inferir conservação filogenética de nicho são, em sua maioria, incompatíveis com determinados conceitos ecológicos e não abrangem todos os tipos de dados e esse fato explica uma visão incompleta dos processos presentes nas comunidades e conflitante com o objetivo de muitos estudos ecológicos e conservacionistas que buscam vincular as espécies aos processos ecológicos e evolutivos. Desta forma, o principal objetivo desta tese é propor novos métodos para quantificar o sinal filogenético que integrem diferentes aspectos do conceito de nicho ecológico. Apresentamos aqui os novos métodos em detalhes e avaliamos suas propriedades estatísticas (erro tipo I e poder estatístico) por meio de dados simulados. No capítulo 1, nós propomos um método para medir sinal filogenético utilizando o teste de Mantel, incorporando modelos evolutivos para testar hipóteses específicas da evolução dos atributos. No capítulo 2, descrevemos um conjunto de funções e um novo pacote estatístico para explorar os padrões filogenéticos no nível de metacomunidade. Este pacote permite explorar a distribuição de linhagens filogenéticas através de gradientes ecológicos, a análise de sinal filogenético no nível da metacomunidade e explorar a associação entre clados e gradientes ecológicos. No capítulo 3, investigamos a relação entre sinal filogenético dos atributos com os padrões de coocorrência das espécies nos níveis da comunidade. Esta abordagem permite testar se espécies filogeneticamente relacionadas que coocorrem expressam as suas dimensões de nicho com maior semelhança do que seria esperado por modelos neutros de evolução. Por fim, testamos as propriedades estatísticas destes métodos em relação dois modelos nulos, que incorporam diferentes aspectos da estrutura da comunidade e evolução dos atributos das espécies. Os três capítulos representam diferentes trabalhos que se interconectam no sentido de elucidar o conceito de sinal filogenético e conservação filogenética de nicho. / Understanding the factors that can affect species distributions has been a main goal of ecologists. Currently, it is known that evolutionary and ecological processes shape the speciation dynamics, species extinction and determine the distribution and abundance of species. Over the last years, there has been an increase in the number of studies using phylogenetic information to explain the dynamics of population, species distribution and identifying the mechanisms of community assembly. Species interactions – intraspecific, interspecific or with the environment – occur based on their phenotypic differences and similarities. As phenotypic variation has a basis in evolutionary history, it is expected that closely related species tend to be more ecologically similar to each other than distantly related ones. This notion has given rise to an important concept, with implications for both evolutionary and ecological studies: the concept of phylogenetic niche conservatism, that is, when related species maintain their ancestral niches over evolutionary time. This pattern is important for several areas of ecology, and allows to link species to ecological processes and to understand the evolutionary ecology of different lineages. Despite its importance, it is crucial the development of appropriate statistical method to measure this pattern and to infer the processes behind it. The methods currently available to infer phylogenetic niche conservatism are sometimes incompatible with some ecological concepts and do not cover all kind of data, this fact leads to an incomplete view of the process acting in the currents communities and conflict with the goal of many ecological and conservation studies that need to link species to ecological and evolutionary processes. The main goal of this dissertation is to propose novel methods to measure phylogenetic signal incorporating different aspects of ecological niche. We introduce novel methods in detail and evaluate its statistical properties (type I error and statistical power) by means of simulated data with known structure. In chapter 1 we propose a method to measure phylogenetic signal using the Mantel test, incorporating evolutionary model to test specific hypothesis of trait evolution. In chapter 2, we describe a set of function and a new statistical package for exploring the phylogenetic patterns at the metacommunity level. This package allows the exploration of distribution of phylogenetic lineages across ecological gradients, the analysis of phylogenetic signal at metacommunity level and to explore the association between clades and ecological gradients. In the chapter 3, we access the relationship between phylogenetic signal in traits and species co-occurrence patterns in the community levels. This approach allows one to test whether phylogenetic close related species cooccurring in metacommunities express their niche dimensions more similarly than would be expected by neutral expectation. We tested the statistical properties of these methods in relation to two null models, which incorporate these different aspects of the community structure and evolution of species traits. The three chapters represent different works that are interconnected in order to elucidate the concept of phylogenetic signal and phylogenetic niche conservatism.

Compreender os fatores que afetam a distribuição das espécies tem sido um dos principais objetivos dos ecólogos. Atualmente, sabe-se que os processos ecológicos e evolutivos moldam a dinâmica de especiação e extinção de espécies, e determinam a distribuição e abundância das mesmas. Ao longo dos últimos anos, tem havido um aumento no número de estudos que utilizam informação filogenética para explicar as dinâmicas populacionais e as distribuições de espécies, e que buscam identificar os mecanismos responsáveis pela montagem das comunidades. Interações das espécies, sejam elas intraespecífica, interespecífica ou com o ambiente, ocorrem baseadas nas diferenças e semelhanças fenotípicas. Essas variações fenotípicas tem origem na evolução das espécies, e com isso espera-se que as espécies proximamente relacionadas tendam a ser ecologicamente mais semelhantes entre si do que as espécies distantemente relacionadas. Esta concepção tem dado origem a um conceito importante, com implicações para estudos tanto ecológicos quanto evolutivos: o conceito de conservação filogenética de nicho, isto é, quando as espécies relacionadas mantêm seus nichos ancestrais ao longo do tempo evolutivo. Esse padrão tem importância para diversas áreas de ecologia, permitindo a ligação das espécies aos processos ecológicos e auxiliando na maior compreensão da ecologia evolutiva das diferentes linhagens. Devido à sua importância, é fundamental o desenvolvimento de métodos estatísticos adequados para quantificar esses padrões e inferir os processos que o subjazem. Atualmente, os métodos utilizados para inferir conservação filogenética de nicho são, em sua maioria, incompatíveis com determinados conceitos ecológicos e não abrangem todos os tipos de dados e esse fato explica uma visão incompleta dos processos presentes nas comunidades e conflitante com o objetivo de muitos estudos ecológicos e conservacionistas que buscam vincular as espécies aos processos ecológicos e evolutivos. Desta forma, o principal objetivo desta tese é propor novos métodos para quantificar o sinal filogenético que integrem diferentes aspectos do conceito de nicho ecológico. Apresentamos aqui os novos métodos em detalhes e avaliamos suas propriedades estatísticas (erro tipo I e poder estatístico) por meio de dados simulados. No capítulo 1, nós propomos um método para medir sinal filogenético utilizando o teste de Mantel, incorporando modelos evolutivos para testar hipóteses específicas da evolução dos atributos. No capítulo 2, descrevemos um conjunto de funções e um novo pacote estatístico para explorar os padrões filogenéticos no nível de metacomunidade. Este pacote permite explorar a distribuição de linhagens filogenéticas através de gradientes ecológicos, a análise de sinal filogenético no nível da metacomunidade e explorar a associação entre clados e gradientes ecológicos. No capítulo 3, investigamos a relação entre sinal filogenético dos atributos com os padrões de coocorrência das espécies nos níveis da comunidade. Esta abordagem permite testar se espécies filogeneticamente relacionadas que coocorrem expressam as suas dimensões de nicho com maior semelhança do que seria esperado por modelos neutros de evolução. Por fim, testamos as propriedades estatísticas destes métodos em relação dois modelos nulos, que incorporam diferentes aspectos da estrutura da comunidade e evolução dos atributos das espécies. Os três capítulos representam diferentes trabalhos que se interconectam no sentido de elucidar o conceito de sinal filogenético e conservação filogenética de nicho. / Understanding the factors that can affect species distributions has been a main goal of ecologists. Currently, it is known that evolutionary and ecological processes shape the speciation dynamics, species extinction and determine the distribution and abundance of species. Over the last years, there has been an increase in the number of studies using phylogenetic information to explain the dynamics of population, species distribution and identifying the mechanisms of community assembly. Species interactions – intraspecific, interspecific or with the environment – occur based on their phenotypic differences and similarities. As phenotypic variation has a basis in evolutionary history, it is expected that closely related species tend to be more ecologically similar to each other than distantly related ones. This notion has given rise to an important concept, with implications for both evolutionary and ecological studies: the concept of phylogenetic niche conservatism, that is, when related species maintain their ancestral niches over evolutionary time. This pattern is important for several areas of ecology, and allows to link species to ecological processes and to understand the evolutionary ecology of different lineages. Despite its importance, it is crucial the development of appropriate statistical method to measure this pattern and to infer the processes behind it. The methods currently available to infer phylogenetic niche conservatism are sometimes incompatible with some ecological concepts and do not cover all kind of data, this fact leads to an incomplete view of the process acting in the currents communities and conflict with the goal of many ecological and conservation studies that need to link species to ecological and evolutionary processes. The main goal of this dissertation is to propose novel methods to measure phylogenetic signal incorporating different aspects of ecological niche. We introduce novel methods in detail and evaluate its statistical properties (type I error and statistical power) by means of simulated data with known structure. In chapter 1 we propose a method to measure phylogenetic signal using the Mantel test, incorporating evolutionary model to test specific hypothesis of trait evolution. In chapter 2, we describe a set of function and a new statistical package for exploring the phylogenetic patterns at the metacommunity level. This package allows the exploration of distribution of phylogenetic lineages across ecological gradients, the analysis of phylogenetic signal at metacommunity level and to explore the association between clades and ecological gradients. In the chapter 3, we access the relationship between phylogenetic signal in traits and species co-occurrence patterns in the community levels. This approach allows one to test whether phylogenetic close related species cooccurring in metacommunities express their niche dimensions more similarly than would be expected by neutral expectation. We tested the statistical properties of these methods in relation to two null models, which incorporate these different aspects of the community structure and evolution of species traits. The three chapters represent different works that are interconnected in order to elucidate the concept of phylogenetic signal and phylogenetic niche conservatism.

Compreender os fatores que afetam a distribuição das espécies tem sido um dos principais objetivos dos ecólogos. Atualmente, sabe-se que os processos ecológicos e evolutivos moldam a dinâmica de especiação e extinção de espécies, e determinam a distribuição e abundância das mesmas. Ao longo dos últimos anos, tem havido um aumento no número de estudos que utilizam informação filogenética para explicar as dinâmicas populacionais e as distribuições de espécies, e que buscam identificar os mecanismos responsáveis pela montagem das comunidades. Interações das espécies, sejam elas intraespecífica, interespecífica ou com o ambiente, ocorrem baseadas nas diferenças e semelhanças fenotípicas. Essas variações fenotípicas tem origem na evolução das espécies, e com isso espera-se que as espécies proximamente relacionadas tendam a ser ecologicamente mais semelhantes entre si do que as espécies distantemente relacionadas. Esta concepção tem dado origem a um conceito importante, com implicações para estudos tanto ecológicos quanto evolutivos: o conceito de conservação filogenética de nicho, isto é, quando as espécies relacionadas mantêm seus nichos ancestrais ao longo do tempo evolutivo. Esse padrão tem importância para diversas áreas de ecologia, permitindo a ligação das espécies aos processos ecológicos e auxiliando na maior compreensão da ecologia evolutiva das diferentes linhagens. Devido à sua importância, é fundamental o desenvolvimento de métodos estatísticos adequados para quantificar esses padrões e inferir os processos que o subjazem. Atualmente, os métodos utilizados para inferir conservação filogenética de nicho são, em sua maioria, incompatíveis com determinados conceitos ecológicos e não abrangem todos os tipos de dados e esse fato explica uma visão incompleta dos processos presentes nas comunidades e conflitante com o objetivo de muitos estudos ecológicos e conservacionistas que buscam vincular as espécies aos processos ecológicos e evolutivos. Desta forma, o principal objetivo desta tese é propor novos métodos para quantificar o sinal filogenético que integrem diferentes aspectos do conceito de nicho ecológico. Apresentamos aqui os novos métodos em detalhes e avaliamos suas propriedades estatísticas (erro tipo I e poder estatístico) por meio de dados simulados. No capítulo 1, nós propomos um método para medir sinal filogenético utilizando o teste de Mantel, incorporando modelos evolutivos para testar hipóteses específicas da evolução dos atributos. No capítulo 2, descrevemos um conjunto de funções e um novo pacote estatístico para explorar os padrões filogenéticos no nível de metacomunidade. Este pacote permite explorar a distribuição de linhagens filogenéticas através de gradientes ecológicos, a análise de sinal filogenético no nível da metacomunidade e explorar a associação entre clados e gradientes ecológicos. No capítulo 3, investigamos a relação entre sinal filogenético dos atributos com os padrões de coocorrência das espécies nos níveis da comunidade. Esta abordagem permite testar se espécies filogeneticamente relacionadas que coocorrem expressam as suas dimensões de nicho com maior semelhança do que seria esperado por modelos neutros de evolução. Por fim, testamos as propriedades estatísticas destes métodos em relação dois modelos nulos, que incorporam diferentes aspectos da estrutura da comunidade e evolução dos atributos das espécies. Os três capítulos representam diferentes trabalhos que se interconectam no sentido de elucidar o conceito de sinal filogenético e conservação filogenética de nicho. / Understanding the factors that can affect species distributions has been a main goal of ecologists. Currently, it is known that evolutionary and ecological processes shape the speciation dynamics, species extinction and determine the distribution and abundance of species. Over the last years, there has been an increase in the number of studies using phylogenetic information to explain the dynamics of population, species distribution and identifying the mechanisms of community assembly. Species interactions – intraspecific, interspecific or with the environment – occur based on their phenotypic differences and similarities. As phenotypic variation has a basis in evolutionary history, it is expected that closely related species tend to be more ecologically similar to each other than distantly related ones. This notion has given rise to an important concept, with implications for both evolutionary and ecological studies: the concept of phylogenetic niche conservatism, that is, when related species maintain their ancestral niches over evolutionary time. This pattern is important for several areas of ecology, and allows to link species to ecological processes and to understand the evolutionary ecology of different lineages. Despite its importance, it is crucial the development of appropriate statistical method to measure this pattern and to infer the processes behind it. The methods currently available to infer phylogenetic niche conservatism are sometimes incompatible with some ecological concepts and do not cover all kind of data, this fact leads to an incomplete view of the process acting in the currents communities and conflict with the goal of many ecological and conservation studies that need to link species to ecological and evolutionary processes. The main goal of this dissertation is to propose novel methods to measure phylogenetic signal incorporating different aspects of ecological niche. We introduce novel methods in detail and evaluate its statistical properties (type I error and statistical power) by means of simulated data with known structure. In chapter 1 we propose a method to measure phylogenetic signal using the Mantel test, incorporating evolutionary model to test specific hypothesis of trait evolution. In chapter 2, we describe a set of function and a new statistical package for exploring the phylogenetic patterns at the metacommunity level. This package allows the exploration of distribution of phylogenetic lineages across ecological gradients, the analysis of phylogenetic signal at metacommunity level and to explore the association between clades and ecological gradients. In the chapter 3, we access the relationship between phylogenetic signal in traits and species co-occurrence patterns in the community levels. This approach allows one to test whether phylogenetic close related species cooccurring in metacommunities express their niche dimensions more similarly than would be expected by neutral expectation. We tested the statistical properties of these methods in relation to two null models, which incorporate these different aspects of the community structure and evolution of species traits. The three chapters represent different works that are interconnected in order to elucidate the concept of phylogenetic signal and phylogenetic niche conservatism.