Parasites are an important component of every ecosystem, and their interactions with hosts can directly impact the fitness of the host organisms. Despite their significance, it is still largely unknown which host traits predispose a species to increased risk of parasite attack. In the present study, I investigated host and ecological correlates of parasite load in Lake Tanganyikan cichlids using phylogenetic comparative analyses. Specifically, I tested the effect of body size, brain mass, gut length, depth, trophic level, habitat complexity and mating system on parasite prevalence and diversity.  I found that the variation in both average infracommunity richness and parasite prevalence were partly explained by host body mass and habitat complexity. Total parasite diversity, on the other hand, was significantly related to mating behaviour as well as habitat complexity. The comparative analysis presented here is, to the best of my knowledge, the first to investigate determinants of parasite load in Lake Tanganyikan cichlids. As such, my results support several major hypotheses concerning the factors that underlie parasite prevalence and diversity and have broad implications for our understanding of parasite-host interactions.

The research presented here deals with inferring phylogenetic trees and their use to study the evolution of functional traits in diatoms (Heterokontophyta: Bacillariophyceae). Two chapters are concerned with the phylogeny of a mainly freshwater group, the Cymbellales, with a convoluted taxonomic history and classification. I generated a multi-gene dataset to test the monophyly of the Cymbellales and reconstruct the relationships within the order. The molecular data were equivocal with respect to the monophyly of the Cymbellales, especially when taking into account some problematic taxa like Cocconeis and Rhoicosphenia. Aside from the problem with their monophyly, my work shows that the current genus- and family-level classification of the Cymbellales is unnatural, arguing for the need of nearly wholesale re-classification of the group. The two following chapters make use of phylogenetic trees to model the evolution of functional traits. I explored the evolution of cell size across the salinity gradient finding that the opposing selective forces exerted by marine and fresh waters select for different optimal cell sizes -- larger in the oceans and smaller in lakes and rivers. Thereafter, I modelled the evolutionary histories of habitat preference (planktonic-benthic) and growth form (solitary-colonial) across the diatoms. These traits exhibit markedly different evolutionary histories. Habitat preference evolves slowly, is conserved at the level of large clades, and its evolution is generally uniform across the tree. Growth form, on the other hand, has a more dynamic evolutionary history with frequent transitions between the solitary and colonial growth forms and rates of evolution that vary through time. I hope that these empirical studies represent an incremental advancement to the understanding of the evolution diatom species and functional diversity. / text

Ever since Darwin claimed that expanding savannas were the driving force behind humanity's divergence from other apes, our understanding of human evolution has been inextricably linked to the environmental context in which our ancestors evolved. This dissertation explores various aspects of the use of one method of paleoenvironmental reconstruction -- bovid ecomorphology -- and provides new data on paleoenvironmental conditions in the Omo Shungura Formation (Plio-Pleistocene, Ethiopia). Chapter 2 uses phylogenetic simulations to explore the performance of Discriminant Function Analysis (DFA) on simulated ecomorphological data containing phylogenetic signal. DFA is shown to "over-perform" in situations in which predicted and predictor variables both contain phylogenetic signal. Phylogenetic Generalized Least Squares (PGLS) is shown to be a very useful technique for explicitly testing functional hypotheses in ecomorphology while controlling for phylogenetic signal and body size. Chapter 3 presents a functional analysis of the bovid astragalus, which is one of the most commonly preserved bones in the fossil record. Several functional hypotheses linking habitat-specific locomotor performance with the morphology of the astragalus are tested using PGLS. Strong support is found for three of these hypotheses. Thus, the astragalus is shown to be a useful ecomorphological predictor element, a point that is confirmed by the DFA analyses in Chapter 4. Chapter 5 provides new paleoenvironmental data on the Omo Shungura Formation based on habitat reconstructions from astragalar ecomorphology in addition to dietary reconstructions based on dental mesowear. Astragalar data point to a major environmental shift beginning ~2.58 Ma, which is later in time compared with some prior habitat reconstructions using different methods. Furthermore, astragalar data show environmental fluctuations of similar magnitude later in the sequence. Mesowear data on the Shungura Tragelaphini do not offer evidence for any significant grazing adaptation, in spite of relatively high carbon isotope signatures reported based on studies of tooth enamel. These data raise questions regarding the diet of fossil Tragalephini. / text

Globalization is not only a transformation of the economical world, but also of geographical space in its totality. The local territories participate in this transformation. To study and compare their evolution can help us to analyze the globalisation as well as its construction, its effects and in its limits, as the existence of social and political dynamics which are not globalized. This reflexion rests on case studies in Peru, made in collaboration with other French and Peruvian researchers. / La globalización no es solo una transformación del mundo económico sino también del espacio geográfico en su conjunto. Los territorios locales participan en esta transformación. Estudiar y comparar sus evoluciones nos puede ayudar a analizar tanto la globalización, en su construcción, en sus efectos y en sus límites, como la existencia de dinámicas sociales y políticas que no son globalizadas. La reflexión se fundamenta sobre casos estudiados en el Perú, en colaboración con otros investigadores franceses y peruanos.

The fossil record yields information on macroevolutionary patterns that remains inaccessible from the study of extant organisms alone, presenting a natural laboratory for us to test hypotheses about the long-term drivers and processes of evolution. Fossil data are therefore increasingly incorporated into evolutionary analyses, both on their own and in combination with neontological data. Phylogeny (an explicit hypothesis of the evolutionary relationships between taxa) can be used as a framework to enable direct comparison of results of comparative methods across many different timescales and taxa, and is now commonly used in investigations of fossil data. This represents an important step towards a unified approach, however, it is not yet fully understood what the effect of using fossil data is on the results of downstream phylogenetic comparative methods, which were originally developed with only living taxa in mind. In this thesis I explore the validity of phylogenetic interpretations of fossil record data. I begin with only taxonomic classification and show that this can in some cases substitute for a cladistically inferred phylogeny in phylogenetic comparative methods, without biasing results. Moving on to scenarios where a timescaled phylogeny is available I investigate the relationship between phylogeny and extinction in the geological past, show that phylogenetic clustering of extinction was common in tetrapods, and present a summary of the ways in which fossil data biases this measurement. Finally, with timescaled phylogenies and a detailed continuous trait dataset available, I interrogate the fossil record of Sauropterygia to uncover the processes of evolutionary change in this highly labile clade. By comparing the results of a suite of phylogenetic comparative methods I demonstrate that neck length evolved through changing vertebral counts rather than somite growth; that the clade experienced a release in evolutionary constraint at the Triassic-Jurassic boundary; and that evidence does not support evolution towards a stationary adaptive peak as a suitable model for phenotypic change in the clade.

No description available.

Projecting environmental niche models through time is a common goal when studying species response to climatic change. Species distribution models (SDMs) are commonly used to estimate a species' niche from observed patterns of occurrence and environmental predictors. However, a species niche is also shaped by non-environmental factors--including biotic interactions and dispersal barrier—truncating SDM estimates. Though truncated SDMs may accurately predict present-day species niche, projections through time are often biased by environmental condition change. Modeling niche in a phylogenetic framework leverages a clade's shared evolutionary history to pull species estimates closer towards phylogenetic conserved values and farther away from species specific biases. We propose a new Bayesian model of phylogenetic niche implemented in R. Under our model, species SDM parameters are transformed into biologically interpretable continuous parameters of environmental niche optimum, breadth, and tolerance evolving under multivariate Brownian motion random walk. Through simulation analyses, we demonstrated model accuracy and precision that improved as phylogeny size increased. We also demonstrated our model on a clade of eastern United States Plethodontid salamanders by accurately estimating species niche, even when no occurrence data is present. Our model demonstrates a novel framework where niche changes can be studied forwards and backwards through time to understand ancestral ranges, patterns of environmental specialization, and niche in data deficient species. / Master of Science / As many species face increasing pressure in a changing climate, it is crucial to understand the set of environmental conditions that shape species' ranges--known as the environmental niche--to guide conservation and land management practices. Species distribution models (SDMs) are common tools that are used to model species' environmental niche. These models treat a species' probability of occurrence as a function of environmental conditions. SDM niche estimates can predict a species' range given climate data, paleoclimate, or projections of future climate change to estimate species range shifts from the past to the future. However, SDM estimates are often biased by non-environmental factors shaping a species' range including competitive divergence or dispersal barriers. Biased SDM estimates can result in range predictions that get worse as we extrapolate beyond the observed climatic conditions. One way to overcome these biases is by leveraging the shared evolutionary history amongst related species to "fill in the gaps". Species that are more closely phylogenetically related often have more similar or "conserved" environmental niches. By estimating environmental niche over all species in a clade jointly, we can leverage niche conservatism to produce more biologically realistic estimates of niche. However, currently a methodological gap exists between SDMs estimates and macroevolutionary models, prohibiting them from being estimated jointly. We propose a novel model of evolutionary niche called PhyNE (Phylogenetic Niche Evolution), where biologically realistic environmental niches are fit across a set of species with occurrence data, while simultaneously fitting and leveraging a model of evolution across a portion of the tree of life. We evaluated model accuracy, bias, and precision through simulation analyses. Accuracy and precision increased with larger phylogeny size and effectively estimated model parameters. We then applied PhyNE to Plethodontid salamanders from Eastern North America. This ecologically-important and diverse group of lungless salamanders require cold and wet conditions and have distributions that are strongly affected by climatic conditions. Species within the family vary greatly in distribution, with some species being wide ranging generalists, while others are hyper-endemics that inhabit specific mountains in the Southern Appalachians with restricted thermal and hydric conditions. We fit PhyNE to occurrence data for these species and their associated average annual precipitation and temperature data. We identified no correlations between species environmental preference and specialization. Pattern of preference and specialization varied among Plethodontid species groups, with more aquatic species possessing a broader environmental niche, likely due to the aquatic microclimate facilitating occurrence in a wider range of conditions. We demonstrated the effectiveness of PhyNE's evolutionarily-informed estimates of environmental niche, even when species' occurrence data is limited or even absent. PhyNE establishes a proof-of-concept framework for a new class of approaches for studying niche evolution, including improved methods for estimating niche for data-deficient species, historical reconstructions, future predictions under climate change, and evaluation of niche evolutionary processes across the tree of life. Our approach establishes a framework for leveraging the rapidly growing availability of biodiversity data and molecular phylogenies to make robust eco-evolutionary predictions and assessments of species' niche and distributions in a rapidly changing world.

Métodos filogenéticos comparativos utilizam informações sobre as relações de ancestralidade entre as espécies para testar hipóteses evolutivas. Neste contexto, a Reconstrução de Caracteres Ancestrais (RCA) pode nos esclarecer muito a respeito dos organismos já extintos. A família Canidae apresenta variada gama de comportamentos, distribuída por quase todo o planeta. Sua rica história fóssil demonstra processos que nos dão pistas sobre a evolução e diversificação destes comportamentos ao longo de 46 Ma. Entender a importância de fatores bióticos e abióticos na evolução de carnívoros tem sido um dos grandes desafios em estudos macroevolutivos nos últimos anos. Aqui foram abordados aspectos evolutivos de Canidae com o intuito de demonstrar o papel de fatores ambientais e comportamentais, além de interações, na diversificação do grupo. Para isso, construiu-se a filogenia para todas as espécies vivas de canídeos e uma espécie recentemente extinta. No total, 37 espécies foram incluídas na árvore filogenética. Foram obtidos 23 marcadores moleculares usados na construção da filogenia. Utilizou-se também 68 caracteres morfológicos. A construção da filogenia foi feita utilizando inferência Bayesiana. O modelo evolutivo escolhido nessa etapa foi GTR + G + I. Também foi utilizado o algoritmo de Monte Carlo Markov Chain (MCMC) para obter a distribuição a posteriori, com 50 x 106 iterações. A datação da árvore filogenética foi feita através do método de Penalized Likelihood, onde foram utilizados 11 registros fósseis de nós conhecidos da filogenia. Após a filogenia feita, obteve-se os dados comportamentais para realização da RCA a respeito dos quatro atributos avaliados. As quatro reconstruções foram criadas com inferência em 1000 árvores cada. Todas análises de RCA foram realizadas com o método de parcimônia. Com o intuito de entender de que maneira os atributos se correlacionam ao longo da filogenia, foi calculada a correlação de Pagel além de Phylogenetic Generalized Least Squares (PGLS). A topologia obtida aqui foi diferente das demais árvores filogenéticas já criadas para Canidae. Além disso, a calibração temporal indica que o split entre Canini e Vulpini se deu há 12.6 Ma, diferente do que é apontado na literatura. A respeito das reconstruções, as linhagens ancestrais dos lobos e das raposas desenvolveram o hábito de viver em áreas abertas. Já os canídeos sulamericanos desenvolveram preferência por áreas florestais. Em relação à dieta, o ancestral de Caninae, assim como os ancestrais diretos das tribos Canini e Vulpini, apresentavam comportamento alimentar hipocarnívoro. O ancestral de todos os canídeos existentes hoje apresentou baixa organização social, enquanto que os lobos desenvolveram alto comportamento social, coincidindo com o surgimento do hábito hipercarnívoro. A respeito do tamanho corporal, o nó ancestral a todos os canídeos possuía tamanho médio, e as duas tribos que surgiram a partir desta linhagem divergiram seus tamanhos. O teste de Pagel demonstrou que há correlação entre dieta e socialidade, mostrando que a alimentação levou a modificações no comportamento Social. Os PGLSs mostram que três tipos de modelos evolutivos explicam as mudanças nos atributos ao longo do tempo. As mudanças no uso de habitat dos canídeos acompanharam as mudanças climáticas no planeta ao longo dos últimos 13 Ma. Já a alimentação meso e hipocarnívora dos sulamericanos se deve ao cenário encontrado na América do Sul ao chegarem, e como reflexo, não desenvolveram alto grau de socialidade. Os resultados sugerem que raposas tentaram evitar a competição com os lobos para não sobreporem seus nichos, sendo o fator fundamental para sua diversificação. / Phylogenetic comparative methods use information on ancestral relationships between species to test evolutionary hypotheses. In this context, the Ancestral Characters Reconstruction (ACR) can shed light on the already extinct organisms. The Canidae family has a wide range of behaviors, distributed throughout most of the planet. Its rich fossil history demonstrates processes that give us clues about the evolution and diversification of these behaviors over 46 Ma. Understanding the importance of biotic and abiotic factors in the evolution of carnivores has been one of the major challenges in macroevolutionary studies in recent years. Here we discuss the evolutionary aspects of Canidae with the purpose of demonstrating the role of environmental and behavioral factors, as well as interactions, in the diversification of the group. For this, the phylogeny was constructed for all living species of canids and a recently extinct species. In total, 37 species were included in the phylogenetic tree. A total of 23 molecular markers were used to construct the phylogeny. We also used 68 morphological characters. The construction of the phylogeny was done using Bayesian inference. The evolutionary model chosen in this step was GTR + G + I. The Monte Carlo Markov Chain algorithm (MCMC) was also used to obtain the posterior distribution, with 50 x 106 iterations. The phylogenetic tree was dated using the Penalized Likelihood method, where eleven fossil records of nodes known from the phylogeny were used. After the phylogeny, the behavioral data were obtained to perform the ACR in relation to the four attributes evaluated. The four reconstructions were created with inference in 1000 trees each. All ACR analyzes were performed using the parsimony method. In order to understand how the attributes correlate throughout the phylogeny, the Pagel correlation was calculated in addition to Phylogenetic Generalized Least Squares (PGLS). The topology obtained here was different from the other phylogenetic trees already created for Canidae. In addition, the time calibration indicates that the split between Canini and Vulpini occurred 12.6 Ma ago, different from what is pointed out in the literature. Concerning reconstructions, the ancestral lineages of wolves and foxes have developed the habit of living in open areas. South American canids have developed preference for forest areas. Regarding diet, Caninae's ancestor, as well as the direct ancestors of the Canini and Vulpini tribes, presented hypocampivorous feeding behavior. The ancestor of all canids present today had a low social organization, while the wolves developed a high social behavior, coinciding with the emergence of the hypercarnivore habit. Regarding the body size, the ancestral node to all canids had medium size, and the two tribes that have emerged from this lineage diverged their sizes. The Pagel test demonstrated that there is a correlation between diet and sociality, showing that diet led to changes in social behavior. The PGLSs show that three types of evolutionary models explain changes in attributes over time. The changes in the habitat use of the canids have accompanied the climatic changes in the planet during the last 13 Ma. The meso and hypocarnivorous feeding of the South Americans is due to the scenario found in South America when they arrived, and as a reflex, they did not develop high degree of Sociality. The results suggest that foxes tried to avoid competition with the wolves to avoid overlapping their niches, being the fundamental factor for their diversification.

<p>What makes our minds human? How did they evolve to be this way? This dissertation presents data from two complementary lines of research driven by these orienting questions. The first of these explores the `what' of human cognitive evolution through comparative studies with chimpanzees and bonobos. The general aim of these studies is to understand which aspects of cognition are unique to humans, and which are shared with our closest living relatives. Chapters 2-3 test the hypothesis that humans have unique cognitive skills for reasoning about the attention of other individuals (theory of mind), and unique motivation to use these skills in cooperative contexts with conspecifics. In Chapter 2 I show that understanding others' attention is unlikely to be the `small difference that makes the big difference', as some researchers have proposed. However, my data support the possibility that species differences in the ontogeny of these skills may have robust consequences for the adult cognitive phenotype. In Chapter 3 I show that (contrary to previous reports) nonhuman apes are also motivated to engage in some simple triadic social activities, which resemble those characteristic of human children. Again however, I identify important differences between humans and other apes in their spontaneous preferences for these types of activities, and their attitudes toward a partner when cooperative behaviors are interrupted. The second half of this dissertation (Chapters 4-5) explores the `why' and `how' of cognitive evolution. Chapter 4 outlines the kind of research questions and methods that comparative psychologists will need to embrace in order to use the comparative method to its full potential in the study of cognitive evolution. Chapter 5 provides a proof of principle for this approach using a dataset including 33 species tested on two cognitive tasks measuring inhibitory control. Here I show that cognitive skills for inhibitory control are closely related to phylogeny across species, and strongly predicted by absolute (but not relative) brain size. Further, I show that two of the other leading hypotheses put forth to explain primate intelligence, namely sociality and diet, do not predict cognitive skills on these tasks. These data illustrate the power of the comparative method for understanding cognitive evolution, and provide a starting point for future studies embracing this approach. Collectively, this research refines our understanding of how human cognition differs from that of other primates, and illustrates the utility of studying cognitive evolution from an explicitly phylogenetic comparative framework.</p> / Dissertation

Métodos filogenéticos comparativos utilizam informações sobre as relações de ancestralidade entre as espécies para testar hipóteses evolutivas. Neste contexto, a Reconstrução de Caracteres Ancestrais (RCA) pode nos esclarecer muito a respeito dos organismos já extintos. A família Canidae apresenta variada gama de comportamentos, distribuída por quase todo o planeta. Sua rica história fóssil demonstra processos que nos dão pistas sobre a evolução e diversificação destes comportamentos ao longo de 46 Ma. Entender a importância de fatores bióticos e abióticos na evolução de carnívoros tem sido um dos grandes desafios em estudos macroevolutivos nos últimos anos. Aqui foram abordados aspectos evolutivos de Canidae com o intuito de demonstrar o papel de fatores ambientais e comportamentais, além de interações, na diversificação do grupo. Para isso, construiu-se a filogenia para todas as espécies vivas de canídeos e uma espécie recentemente extinta. No total, 37 espécies foram incluídas na árvore filogenética. Foram obtidos 23 marcadores moleculares usados na construção da filogenia. Utilizou-se também 68 caracteres morfológicos. A construção da filogenia foi feita utilizando inferência Bayesiana. O modelo evolutivo escolhido nessa etapa foi GTR + G + I. Também foi utilizado o algoritmo de Monte Carlo Markov Chain (MCMC) para obter a distribuição a posteriori, com 50 x 106 iterações. A datação da árvore filogenética foi feita através do método de Penalized Likelihood, onde foram utilizados 11 registros fósseis de nós conhecidos da filogenia. Após a filogenia feita, obteve-se os dados comportamentais para realização da RCA a respeito dos quatro atributos avaliados. As quatro reconstruções foram criadas com inferência em 1000 árvores cada. Todas análises de RCA foram realizadas com o método de parcimônia. Com o intuito de entender de que maneira os atributos se correlacionam ao longo da filogenia, foi calculada a correlação de Pagel além de Phylogenetic Generalized Least Squares (PGLS). A topologia obtida aqui foi diferente das demais árvores filogenéticas já criadas para Canidae. Além disso, a calibração temporal indica que o split entre Canini e Vulpini se deu há 12.6 Ma, diferente do que é apontado na literatura. A respeito das reconstruções, as linhagens ancestrais dos lobos e das raposas desenvolveram o hábito de viver em áreas abertas. Já os canídeos sulamericanos desenvolveram preferência por áreas florestais. Em relação à dieta, o ancestral de Caninae, assim como os ancestrais diretos das tribos Canini e Vulpini, apresentavam comportamento alimentar hipocarnívoro. O ancestral de todos os canídeos existentes hoje apresentou baixa organização social, enquanto que os lobos desenvolveram alto comportamento social, coincidindo com o surgimento do hábito hipercarnívoro. A respeito do tamanho corporal, o nó ancestral a todos os canídeos possuía tamanho médio, e as duas tribos que surgiram a partir desta linhagem divergiram seus tamanhos. O teste de Pagel demonstrou que há correlação entre dieta e socialidade, mostrando que a alimentação levou a modificações no comportamento Social. Os PGLSs mostram que três tipos de modelos evolutivos explicam as mudanças nos atributos ao longo do tempo. As mudanças no uso de habitat dos canídeos acompanharam as mudanças climáticas no planeta ao longo dos últimos 13 Ma. Já a alimentação meso e hipocarnívora dos sulamericanos se deve ao cenário encontrado na América do Sul ao chegarem, e como reflexo, não desenvolveram alto grau de socialidade. Os resultados sugerem que raposas tentaram evitar a competição com os lobos para não sobreporem seus nichos, sendo o fator fundamental para sua diversificação. / Phylogenetic comparative methods use information on ancestral relationships between species to test evolutionary hypotheses. In this context, the Ancestral Characters Reconstruction (ACR) can shed light on the already extinct organisms. The Canidae family has a wide range of behaviors, distributed throughout most of the planet. Its rich fossil history demonstrates processes that give us clues about the evolution and diversification of these behaviors over 46 Ma. Understanding the importance of biotic and abiotic factors in the evolution of carnivores has been one of the major challenges in macroevolutionary studies in recent years. Here we discuss the evolutionary aspects of Canidae with the purpose of demonstrating the role of environmental and behavioral factors, as well as interactions, in the diversification of the group. For this, the phylogeny was constructed for all living species of canids and a recently extinct species. In total, 37 species were included in the phylogenetic tree. A total of 23 molecular markers were used to construct the phylogeny. We also used 68 morphological characters. The construction of the phylogeny was done using Bayesian inference. The evolutionary model chosen in this step was GTR + G + I. The Monte Carlo Markov Chain algorithm (MCMC) was also used to obtain the posterior distribution, with 50 x 106 iterations. The phylogenetic tree was dated using the Penalized Likelihood method, where eleven fossil records of nodes known from the phylogeny were used. After the phylogeny, the behavioral data were obtained to perform the ACR in relation to the four attributes evaluated. The four reconstructions were created with inference in 1000 trees each. All ACR analyzes were performed using the parsimony method. In order to understand how the attributes correlate throughout the phylogeny, the Pagel correlation was calculated in addition to Phylogenetic Generalized Least Squares (PGLS). The topology obtained here was different from the other phylogenetic trees already created for Canidae. In addition, the time calibration indicates that the split between Canini and Vulpini occurred 12.6 Ma ago, different from what is pointed out in the literature. Concerning reconstructions, the ancestral lineages of wolves and foxes have developed the habit of living in open areas. South American canids have developed preference for forest areas. Regarding diet, Caninae's ancestor, as well as the direct ancestors of the Canini and Vulpini tribes, presented hypocampivorous feeding behavior. The ancestor of all canids present today had a low social organization, while the wolves developed a high social behavior, coinciding with the emergence of the hypercarnivore habit. Regarding the body size, the ancestral node to all canids had medium size, and the two tribes that have emerged from this lineage diverged their sizes. The Pagel test demonstrated that there is a correlation between diet and sociality, showing that diet led to changes in social behavior. The PGLSs show that three types of evolutionary models explain changes in attributes over time. The changes in the habitat use of the canids have accompanied the climatic changes in the planet during the last 13 Ma. The meso and hypocarnivorous feeding of the South Americans is due to the scenario found in South America when they arrived, and as a reflex, they did not develop high degree of Sociality. The results suggest that foxes tried to avoid competition with the wolves to avoid overlapping their niches, being the fundamental factor for their diversification.