Rodents exhibit diverse biology, mating systems and diverse sperm morphology. Different levels of sperm competition have therefore been found in this group. Generally, it is accepted that sperm competition in animals causes the prolongation of sperm tails. However, what other sperm morphological adaptations in rodents are affected by sperm competition? Species-specific sexual selection likely optimizes sperm morphology. Sperm physiology, morphology and competition may be influenced, ultimately, by environmental factors. This Ph.D. thesis focuses on sperm competition in murine rodents. The thesis contains three research articles (including one as a submitted manuscript). In addition, another published article, partialy related to the topic of this thesis, is represented in the Appendix. The sperm morphology in rodents plays an important role in sperm competition. Longer tails and apical hooks are usually found in the species with relatively larger testes and with higher percentage of multiple paternity (i.e. rate of promiscuity), that is in species with the higher risk of sperm competition. The goal of the first study was to investigate the relationships between relative testis weight and sperm traits (apical hook and tail lengths and variance in this traits). The apical hook length was positively...

O objetivo geral desta tese foi investigar como a distribuição espacial dos indivíduos influencia a organização dos sistemas de acasalamento e a intensidade da seleção sexual. A tese contém dois capítulos em que analisamos dados empíricos e dois capítulo em que abordamos questões mais teóricas mais gerais. No primeiro capítulo, avaliamos como a distribuição espacial de haréns defendidos por machos territoriais influencia a rede de competição espermática que existe entre machos do opilião Serracutisoma proximum. No segundo capítulo, propusemos um modelo probabilístico de escolha de parceiros que leva em consideração o fato de que as fêmeas estão restritas a amostrar apenas alguns machos da população, e que essa restrição é imposta principalmente pela distribuição espacial dos indivíduos. Nosso modelo foi mais acurado do que modelos alternativos que ignoram o espaço. No terceiro capítulo, investigamos os movimentos de busca de parceiros do besouro Leptinotarsa undecimlineata. Encontramos que tanto machos quanto fêmeas se movimentam estrategicamente, saindo de onde estão quando não obtém cópulas buscando áreas próximas e com muitos parceiros em potencial. Finalmente, no quarto capítulo, voltamos a explorar o efeito de restrições espaciais sobre a escolha de parceiros. Usamos simulações baseadas em indivíduos para investigar como a restrição espacial influencia a seleção sexual e a evolução de ornamentos sexualmente selecionados. Encontramos que quanto maior o número de parceiros que as fêmeas podem amostrar durante a escolha, mais intensa é a seleção sexual, o que permite a evolução de ornamentos mais exagerados nos machos. Além disso, analisamos um conjunto de dados publicados sobre a intensidade de seleção sexual e observamos que estes se ajustam às previsões do modelo teórico. Concluímos que o espaço é muito mais importante para a organização de sistemas de acasalamento e para a intensidade da seleção sexual do que se acreditava previamente. Esperamos que as idéias propostas aqui encontrem terreno fértil na mente do leitor e que gerem novos e estimulantes desdobramentos no campo teórico e empírico / The main goal of this thesis was to investigate how the spatial distribution of individuals influences the organization of the mating systems and the intensity of sexual selection. The thesis contains two empirical chapters and two chapters with a theoretical scope. In the first chapter, we investigate how the spatial distribution of harems defended by territorial males influences the sperm competition network among males of the harvestman Serracutisoma proximum. In the second chapter, we proposed a probabilistic model of mate choice that includes the spatial constraint in the analyses of mate choice. This model takes into account the fact that females are restricted to sample only some males available in the population, and that this restriction is imposed primarily by the spatial distribution of individuals. In the third chapter, we investigate the mate searching movements of the leaf beetle Leptinotarsa undecimlineata. We found that both males and females move strategically, leaving the host plant when they do not obtain copulations and seeking neighboring areas with many potential mates. Finally, in the fourth chapter, we explore the effect of spatial restrictions on mate choice. We used individual based simulations to investigate how spatial restrictions influence sexual selection and the evolution of sexually selected ornaments. We found that the higher the number of potential partners female can sample during mate choice, the more intense is sexual selection, which leads to the evolution of more exaggerated male ornaments. In addition, we analyzed a published dataset of intensity of sexual selection and found that the data adjust to the predictions made by our simulations. We conclude that the space is much more important than previously thought to mating systems organization and the intensity of sexual selection. We hope the ideas proposed here can flourish in the minds of the readers, stimulating both empirical and theoretical follow ups

A seleção e produção de touros para o mercado pecuário tornam-se foco de muitos estudos, e um grande desafio para o segmento em se tratando da constante busca de melhorias nos desempenhos produtivos dessa categoria animal. A tese teve por objetivo simular o que poderá acontecer com a produção de reprodutores, em termos de quantidades produzidas e custos operacionais efetivos, sob a influência dos métodos de acasalamentos e das biotécnicas reprodutivas disponíveis no mercado nacional. A hipótese em questão é a de testar se acasalar matrizes, classificadas por genótipo, suas produções serão melhores do que quando comparada à produção de matrizes classificadas por fenótipo. Foram simulados 42 cenários produtivos, divididos em 21 cenários com acasalamento genotípico e 21 com acasalamento fenotípico. Em cada divisão constam simulações com uso de Monta Natural, Inseminação Artificial padrão e com sêmen sexado para machos, Transferência de Embriões padrão e com sêmen sexado para machos, Fecundação in vitro padrão e com sêmen sexado para machos, sendo que em todos os casos três níveis diferentes de taxas de concepção foram testados. Os resultados apontaram que o sistema de acasalamento teve influência direta na produção de tourinhos, sendo que o acasalamento por genótipo foi mais eficiente do que o acasalamento por fenótipo. As taxas de concepção influenciaram negativamente mais os resultados dos sistemas de acasalamento fenotípico. Há um grande aumento no custo operacional efetivo dos sistemas que utilizaram as biotécnicas reprodutivas de Transferência de Embriões e Fecundação in vitro, e consequentemente uma redução na lucratividade destes sistemas. As simulações com Monta Natural apresentaram as maiores Margens Brutas e as simulações com Inseminação Artificial com uso de sêmen sexado para machos apresentaram os maiores Lucros Brutos. / The selection and production of bulls for the cattle market have both become the goal of many studies, as well as a great challenge for the segment, providing the constant search for performance improvement of that sort of animal. The target of this thesis is to simulate what may happen to the production of stud, in terms of quantity and effective operational costs, under the influence of the mating methods and the available reproduction biotechniques in the national market. The hypothesis under analysis consists of testing if mating matrices, classed by their genotype, their produce will be better than when compared to the produce of phenotype-classed matrices. 42 productive-scenes, divided into 21 genotype mating and 21 phenotype mating were simulated. In each of the scene divisions there were the following simulations: Natural Breeding, standard Artificial Insemination, Artificial Insemination with male-gendered semen, standard Embryo Transference, Embryo Transference with male-gendered semen, standard In-vitro Fecundation and In-vitro Fecundation with male-gendered semen, so that in all the cases, three different conception rates were tested. The results displayed that the mating system directly influenced the young bull production, once the genotype mating was more efficient than the phenotype mating. The conception rates negatively influenced the results of the phenotype mating mainly. There was a dramatic increase in the effective operational cost of the systems that used the reproductive biotechniques of Embryo Transference and In-vitro Fecundation, and, consequently, profitability reduction of those systems. The Natural Breeding simulations presented the highest Gross Margin, and the simulations of Artificial Insemination with male-gendered semen showed the highest Gross Profit.

<p>Natural selection mediated by pollinators has influenced the evolution of floral diversity of the flowering plants (angiosperms). The scope of this thesis was to study: 1) phenotypic selection, 2) mating systems, and 3) floral shifts involved in plant speciation. Model plant species were <i>Platanthera bifolia</i> and <i>P. chlorantha</i> (Orchidaceae). These orchids are moth-pollinated, strictly co-sexual (bisexual flowers), and produce a spike that displays 10-20 white flowers.</p><p>I explored the influence of characters on plant fitness by using multiple linear regressions. Pollen removal (male fitness) and fruit set (female fitness) increased with more flowers per plant in three <i>P. bifolia</i> populations. There was selection towards longer spurs in a dry year when average spur length was shorter than in normal-wet years. Female function was sensitive to drought, which enabled an application of the male function hypothesis of floral evolution (Bateman's principle). The results show that selection may vary between populations, years, and sex-functions.</p><p>I examined inbreeding by estimating levels of geitonogamy (self-pollination between flowers of an individual) with an emasculation method in two <i>P. bifolia</i> populations. Geitonogamy did not vary with inflorescence size. Levels of geitonogamy was 20-40% in the smaller, but non-significant in the larger population. This may relate to lower number of possible mates and pollinator activity in the smaller population.</p><p><i>Platanthera bifolia</i> exhibits the ancestral character state of tongue-attachment of pollinia on the pollinator. Its close relative <i>P. chlorantha</i> attaches its pollinia onto the pollinator's eyes. To explore the mechanism of a floral shift, pollination efficiency and speed was compared between the two species. The results showed no differences in pollination efficiency, but <i>P. chlorantha</i> had faster pollen export and import. Efficiency of pollination in terms of speed may cause floral shifts, and thus speciation.</p>

Natural selection mediated by pollinators has influenced the evolution of floral diversity of the flowering plants (angiosperms). The scope of this thesis was to study: 1) phenotypic selection, 2) mating systems, and 3) floral shifts involved in plant speciation. Model plant species were Platanthera bifolia and P. chlorantha (Orchidaceae). These orchids are moth-pollinated, strictly co-sexual (bisexual flowers), and produce a spike that displays 10-20 white flowers. I explored the influence of characters on plant fitness by using multiple linear regressions. Pollen removal (male fitness) and fruit set (female fitness) increased with more flowers per plant in three P. bifolia populations. There was selection towards longer spurs in a dry year when average spur length was shorter than in normal-wet years. Female function was sensitive to drought, which enabled an application of the male function hypothesis of floral evolution (Bateman's principle). The results show that selection may vary between populations, years, and sex-functions. I examined inbreeding by estimating levels of geitonogamy (self-pollination between flowers of an individual) with an emasculation method in two P. bifolia populations. Geitonogamy did not vary with inflorescence size. Levels of geitonogamy was 20-40% in the smaller, but non-significant in the larger population. This may relate to lower number of possible mates and pollinator activity in the smaller population. Platanthera bifolia exhibits the ancestral character state of tongue-attachment of pollinia on the pollinator. Its close relative P. chlorantha attaches its pollinia onto the pollinator's eyes. To explore the mechanism of a floral shift, pollination efficiency and speed was compared between the two species. The results showed no differences in pollination efficiency, but P. chlorantha had faster pollen export and import. Efficiency of pollination in terms of speed may cause floral shifts, and thus speciation.

Sicyos deppei G. Don (Cucurbitaceae) is an invasive monoic annual tendril-bearing vine; it is endemic to Mexico, adapted to the rainfall cycle (June through the end of September) and produces spiny, single seeded fruits. Under serious infestation conditions, S. deppei grows extensively, covering native plants, crops and tree trunks. This study determined the breeding system and pollinators of S. deppei growing in three study sites at urban gardens and disturbed sites at the Campus of the Universidad de las Américas, Puebla in the city of San Andrés Cholula, Puebla (Mexico). Each female inflorescence had an average of 17 flowers at the three study sites. As a step prior to experiments, the timing for stigma receptiveness and pollen viability was determined with the Peroxtesmo KO test (PKO) and Diaminobenzidine test (DAB),respectively. Stigmas of female flowers reach maximum receptivity when flowers are at anthesis; receptivity decreases as the flower reaches the succeeding floral stages. Viability of pollen grains increases with flower development; viabilitypercentages at early stages of floral development are very low, as opposed to the higher percentages of pollen viability found in flowers at anthesis and following developmental stages. Breeding systems were tested for xenogamy and geitonogamy by hand pollinating female flowers. Breeding systems results demonstrated that S. i deppei has a mixed-mating system, being able to set fruit and seed when pollinated with pollen from different plants and from pollen of the same plants. The pollinators were determined with the single-visit method. The diversity of floral visitors in this study was low; the most effective pollinator for S. deppei in this study was Apis mellifera. Throughout the rainy season, this study also described the phenology of S. deppei at the plant and flower level, as well as total plant length. Each stage of development in male and female flowers lasts one day. At the end of the rainy season (N2010) individual plants measured between 947 and 270 cm. / CONACYT, NSERC-CANPOLIN

O objetivo geral desta tese foi investigar como a distribuição espacial dos indivíduos influencia a organização dos sistemas de acasalamento e a intensidade da seleção sexual. A tese contém dois capítulos em que analisamos dados empíricos e dois capítulo em que abordamos questões mais teóricas mais gerais. No primeiro capítulo, avaliamos como a distribuição espacial de haréns defendidos por machos territoriais influencia a rede de competição espermática que existe entre machos do opilião Serracutisoma proximum. No segundo capítulo, propusemos um modelo probabilístico de escolha de parceiros que leva em consideração o fato de que as fêmeas estão restritas a amostrar apenas alguns machos da população, e que essa restrição é imposta principalmente pela distribuição espacial dos indivíduos. Nosso modelo foi mais acurado do que modelos alternativos que ignoram o espaço. No terceiro capítulo, investigamos os movimentos de busca de parceiros do besouro Leptinotarsa undecimlineata. Encontramos que tanto machos quanto fêmeas se movimentam estrategicamente, saindo de onde estão quando não obtém cópulas buscando áreas próximas e com muitos parceiros em potencial. Finalmente, no quarto capítulo, voltamos a explorar o efeito de restrições espaciais sobre a escolha de parceiros. Usamos simulações baseadas em indivíduos para investigar como a restrição espacial influencia a seleção sexual e a evolução de ornamentos sexualmente selecionados. Encontramos que quanto maior o número de parceiros que as fêmeas podem amostrar durante a escolha, mais intensa é a seleção sexual, o que permite a evolução de ornamentos mais exagerados nos machos. Além disso, analisamos um conjunto de dados publicados sobre a intensidade de seleção sexual e observamos que estes se ajustam às previsões do modelo teórico. Concluímos que o espaço é muito mais importante para a organização de sistemas de acasalamento e para a intensidade da seleção sexual do que se acreditava previamente. Esperamos que as idéias propostas aqui encontrem terreno fértil na mente do leitor e que gerem novos e estimulantes desdobramentos no campo teórico e empírico / The main goal of this thesis was to investigate how the spatial distribution of individuals influences the organization of the mating systems and the intensity of sexual selection. The thesis contains two empirical chapters and two chapters with a theoretical scope. In the first chapter, we investigate how the spatial distribution of harems defended by territorial males influences the sperm competition network among males of the harvestman Serracutisoma proximum. In the second chapter, we proposed a probabilistic model of mate choice that includes the spatial constraint in the analyses of mate choice. This model takes into account the fact that females are restricted to sample only some males available in the population, and that this restriction is imposed primarily by the spatial distribution of individuals. In the third chapter, we investigate the mate searching movements of the leaf beetle Leptinotarsa undecimlineata. We found that both males and females move strategically, leaving the host plant when they do not obtain copulations and seeking neighboring areas with many potential mates. Finally, in the fourth chapter, we explore the effect of spatial restrictions on mate choice. We used individual based simulations to investigate how spatial restrictions influence sexual selection and the evolution of sexually selected ornaments. We found that the higher the number of potential partners female can sample during mate choice, the more intense is sexual selection, which leads to the evolution of more exaggerated male ornaments. In addition, we analyzed a published dataset of intensity of sexual selection and found that the data adjust to the predictions made by our simulations. We conclude that the space is much more important than previously thought to mating systems organization and the intensity of sexual selection. We hope the ideas proposed here can flourish in the minds of the readers, stimulating both empirical and theoretical follow ups

A seleção e produção de touros para o mercado pecuário tornam-se foco de muitos estudos, e um grande desafio para o segmento em se tratando da constante busca de melhorias nos desempenhos produtivos dessa categoria animal. A tese teve por objetivo simular o que poderá acontecer com a produção de reprodutores, em termos de quantidades produzidas e custos operacionais efetivos, sob a influência dos métodos de acasalamentos e das biotécnicas reprodutivas disponíveis no mercado nacional. A hipótese em questão é a de testar se acasalar matrizes, classificadas por genótipo, suas produções serão melhores do que quando comparada à produção de matrizes classificadas por fenótipo. Foram simulados 42 cenários produtivos, divididos em 21 cenários com acasalamento genotípico e 21 com acasalamento fenotípico. Em cada divisão constam simulações com uso de Monta Natural, Inseminação Artificial padrão e com sêmen sexado para machos, Transferência de Embriões padrão e com sêmen sexado para machos, Fecundação in vitro padrão e com sêmen sexado para machos, sendo que em todos os casos três níveis diferentes de taxas de concepção foram testados. Os resultados apontaram que o sistema de acasalamento teve influência direta na produção de tourinhos, sendo que o acasalamento por genótipo foi mais eficiente do que o acasalamento por fenótipo. As taxas de concepção influenciaram negativamente mais os resultados dos sistemas de acasalamento fenotípico. Há um grande aumento no custo operacional efetivo dos sistemas que utilizaram as biotécnicas reprodutivas de Transferência de Embriões e Fecundação in vitro, e consequentemente uma redução na lucratividade destes sistemas. As simulações com Monta Natural apresentaram as maiores Margens Brutas e as simulações com Inseminação Artificial com uso de sêmen sexado para machos apresentaram os maiores Lucros Brutos. / The selection and production of bulls for the cattle market have both become the goal of many studies, as well as a great challenge for the segment, providing the constant search for performance improvement of that sort of animal. The target of this thesis is to simulate what may happen to the production of stud, in terms of quantity and effective operational costs, under the influence of the mating methods and the available reproduction biotechniques in the national market. The hypothesis under analysis consists of testing if mating matrices, classed by their genotype, their produce will be better than when compared to the produce of phenotype-classed matrices. 42 productive-scenes, divided into 21 genotype mating and 21 phenotype mating were simulated. In each of the scene divisions there were the following simulations: Natural Breeding, standard Artificial Insemination, Artificial Insemination with male-gendered semen, standard Embryo Transference, Embryo Transference with male-gendered semen, standard In-vitro Fecundation and In-vitro Fecundation with male-gendered semen, so that in all the cases, three different conception rates were tested. The results displayed that the mating system directly influenced the young bull production, once the genotype mating was more efficient than the phenotype mating. The conception rates negatively influenced the results of the phenotype mating mainly. There was a dramatic increase in the effective operational cost of the systems that used the reproductive biotechniques of Embryo Transference and In-vitro Fecundation, and, consequently, profitability reduction of those systems. The Natural Breeding simulations presented the highest Gross Margin, and the simulations of Artificial Insemination with male-gendered semen showed the highest Gross Profit.

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The drivers of fidelity and divorce of pair-bonded individuals, along with their fitness consequences, are of great interest as they influence mating systems, population structure and productivity, and gene flow. Socially monogamous birds offer an ideal opportunity to study divorce since they show great variability in the extent to which pair bonds are maintained. However, there has been little consensus as to whether divorce is a behavioural adaptation to improve a mating situation, or a consequence of other processes. Moreover, the biological and ecological correlates of fidelity are difficult to address because previous work has been based on indirect and potentially biased methods. Finally, in terms of process, the link between the process of mate choice and subsequent mating decisions has been largely inaccessible to study. My doctoral thesis addressed these significant gaps in our understanding of cause, process and consequence in the formation and dissolution of pair bonds in socially monogamous birds. I accomplished this in three principal ways. First, I conducted a robust phylogenetic meta-analysis on 84 studies across 64 species to assess the existing empirical evidence that divorce in socially monogamous birds is adaptive (in terms of breeding success). This analysis revealed that divorce is, in general, adaptive as it is both triggered by relatively low breeding success and leads to improvement in success. Next, I developed a novel probabilistic multievent capture–mark–recapture framework that provides joint estimates of survival and fidelity while explicitly accounting for imperfect detection, capture heterogeneity, and uncertainty in pair status. By applying this model to breeding data on a wild great tit population I showed that birds that remain faithful to their partner exhibit higher survival rates and are more likely to remain faithful in the next breeding season than do birds that change partners. Subsequently, I confirmed the generality of a survival benefit by applying the model to breeding data on other tit populations. Then, by applying the model to data from a population of mute swans, I showed that fidelity decreases the likelihood of skipping breeding and mortality in this long-lived species, and that these effects depended on age, individual quality, and immigration status. Finally, I investigated how the timing of pair formation influences breeding success and divorce probability using five years of data on the over-winter social behaviour of great tits. I showed that early pair formation had a positive effect on fitness components, influencing the likelihood of divorce only indirectly, through breeding success. Further, my work revealed that males, but not females, with higher numbers of the female associates in winter, and males whose future breeding partners were ranked low amongst these, divorced more often. My research makes a significant contribution to our understanding of divorce and fidelity, and generates a number of important implications for future studies. First, my work establishes that divorce is adaptive for breeding success. Second, my results highlight that survival is an important (and likely, widespread) fitness consequence of pairing decisions. Third, I provide a novel statistically rigorous modelling framework for estimating fidelity-rates and testing hypothesis about fidelity that overcomes many of the inherent biases in traditional estimates. Fourth, it provides the first evidence for a selective advantage of early pair formation in wild, thus highlighting that there are benefits to pair familiarity that manifest via social associations of individuals prior to breeding. Finally, my work reveals the selective pressures operating via the social environment can ultimately influence the mating strategies individuals adopt.