FundaÃÃo Cearense de Apoio ao Desenvolvimento Cientifico e TecnolÃgico / Territorial fights between males are frequently observed among insects. However, despite the existence of theoretical models directed at explaining the possible rules adopted to decide the winner, there is little empirical support for these models. The low empirical support may be explained by difficulties in identifying traits that determine the male fighting capacity (RHP- or Resource Holding Potential) and also by the lack of simultaneous tests of different models. In this study we evaluated males traits that may determine RHP in the dragonfly Diastatops obscura. Also, we evaluated the relationship between such characteristics and duration of disputes between pairs of males to test predictions derived from the models War of Attrition (GDA), Sequential Access Information (ASI) and Cumulative Access Information (ACI). Male weight was the main feature related to the winning chances. The dispute duration decreases with increasing weight and presented the opposite pattern for the loser male, rejecting the GDA as the decision rule adopted by males to settle territorial interactions. In addition the duration of the dispute was not related to the loser male weight when we considered only pairs of rivals who presented similar weight differences. This indicates that the rivals make mutual assessment of RHP, as assumed by the model ASI. Since wings are highly melanized, we suggest that males may perform rival assessments of RHP based on wings traits, as occur in odonate species. / Apesar das brigas entre os machos serem frequentes e de existirem modelos teÃricos que postulam possÃveis regras usadas para decisÃo do vencedor, pouco suporte empÃrico direcionado ao entendimento das brigas tem sido encontrado. Provavelmente esse fato ocorre devido à dificuldade em identificar quais caracterÃsticas determinam a capacidade de luta (RHP - Resource Holding Potential) dos indivÃduos e na escassez de testes simultÃneos de diferentes modelos. Neste trabalho, usamos machos da libÃlula Diastatops obscura para identificar caracterÃsticas determinantes do RHP. Posteriormente usamos informaÃÃes sobre a relaÃÃo entre tais caracterÃsticas e a duraÃÃo das disputas entre pares de machos para testar previsÃes derivadas dos modelos Guerra de Atritos (GDA), Acesso Sequencial de InformaÃÃo (ASI) e Acesso Cumulativo de InformaÃÃo (ACI). O peso representou a principal caracterÃstica determinante da chance de vitÃria em uma disputa. A duraÃÃo da disputa diminuiu com o aumento do peso do vencedor e aumentou com o aumento de peso do perdedor, refutando o modelo GDA como explicaÃÃo das regras de resoluÃÃo das brigas. Adicionalmente a duraÃÃo da disputa nÃo esteve relacionada com o peso do macho perdedor quando consideramos apenas pares de rivais que apresentaram diferenÃas semelhantes de peso. Isso indica que os rivais fazem avaliaÃÃo mÃtua do RHP, como presumido pelo modelo ASI. Uma vez que as asas sÃo fortemente melanizadas, sugerimos que eles podem realizar avaliaÃÃes da capacidade de luta ao obter informaÃÃes sobre a coloraÃÃo das asas, como ocorre em outras espÃcies de libÃlulas.

FundaÃÃo Cearense de Apoio ao Desenvolvimento Cientifico e TecnolÃgico / Em muitas espÃcies animais, os machos brigam por territÃrios os quais aumentam as chances de cÃpulas. Os comportamentos de luta dos machos podem variar desde embates sem contato fÃsico atà lutas com injÃrias e possÃveis mortes. Existem pelo menos trÃs modelos propostos para explicar as regras usadas pelos machos para decidirem o vencedor de um confronto territorial: Guerra de Atritos (GDA), Acesso Sequencial de InformaÃÃes (ASI) e Acesso Cumulativo de InformaÃÃes (ACI). PorÃm, para sermos capazes de testar tais modelos à necessÃrio identificar primeiro quais traÃos dos machos determinam funcionalmente sua capacidade de luta (RHP). Para isso, usamos machos da libÃlula Macrothemis imitans para avaliar duas hipÃteses: 1) se as brigas ocorrerem com contato fÃsico, caracterÃsticas ligadas ao tamanho determinarÃo o RHP e 2) se os machos nÃo exibirem contato fÃsico durante as brigas, caracterÃsticas que conferem maior resistÃncia determinarÃo o RHP. Realizamos coletas de campo em campanhas de dois dias seguidos. Nessas campanhas realizamos captura, marcaÃÃo, observaÃÃo comportamental e recaptura de duas categorias de machos: os que possuÃam posse dos territÃrios (machos residentes) e os machos que ocuparam o local apÃs a remoÃÃo experimental dos machos residentes (machos substitutos). ApÃs a recaptura, levamos os machos residentes e substitutos para laboratÃrio para realizar as medidas: Ãrea alar, peso fresco, massa muscular torÃcica e quantidade de gordura. Os resultados indicaram que os machos residentes apresentaram maior quantidade de massa muscular e possivelmente gordura do que os substitutos. Uma vez que as brigas ocorreram com contato fÃsico, a maior quantidade de massa muscular e gordura dos machos residentes indica que a relaÃÃo funcional entre comportamento de luta e traÃos determinantes do RHP deve ser rejeitada. Alternativamente, à possÃvel que a maior quantidade de mÃsculo proporcione maior desempenho para causar injÃrias nos rivais ou evitar acÃmulo acelerado de danos. / In many animal species, males fight for the possession of territories that increase their mating chances. The fighting behavior of males may range from conflicts without physical contact to disputes with injuries and possible deaths. There are three models aiming to explain the rules adopted by males to decide the winner of a contest: war of attrition (WOA), sequential assessment model (SAM) and cumulative assessment model (ACM). However, to be able to test these models it is necessary to identify male traits functionally related to fighting ability (RHP). In this sense, we used males of the dragonfly Macrothemis imitans to evaluate two hypotheses: 1) if contests occur with physical contact, traits related to the size define RHP and 2) if disputes occur without physical contact, traits which confer endurance define male RHP. We did field experiments during campaigns with two consecutive days. In each campaign we captured, marked, made behavioral observations and recaptured males that were defending a territorial site (resident males) and males that stablished territories in this sites after the experimental removal of the resident male (substitute males). After recapturing, we transported all males them to lab to measure total wing area, male fresh weight, muscle mass and fat content.The results indicated that resident males presented higher muscle mass and probably fat content than substitute males. Since disputes occurred with physical contact, these differences in muscle mass and fat content indicate that the functional relationship between fight behavior and traits linked to RHP should be rejected. Alternativelly, it is possible that the higher muscle mass confers higher ability to cause injuries or to avoid damage accrual.

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Selection operates when the variability among individuals in heritable traits translates to differences in the number of offspring that survive to breed, which is a close estimate of fitness. Consequently, the outcome of sexual-selection should be higher reproductive success for individuals with a greater expression of the selected traits. In this thesis, the relationship between some male spotless starling (Sturnus unicolor) traits and reproductive success was assessed. A particular focus was given to the role of throat feathers (TF) as sexually selected trait. The study was conducted in a wild population using a correlative approach in 2004, while in 2005 and 2006 the TF of males were experimentally shortened. The genetic parentage of the offspring was required for determining the reproductive success of males. Nine highly polymorphic microsatellites (with 11.7± 3.2 alleles per locus) were developed and optimised for this species. Parentage analyses were conducted in NEWPAT XL and CERVUS 3.0.3 and confirmed using observational data. Eighty-five percent of the offspring had at least one parent assigned. The levels of intra-specific brood parasitism, extra-pair paternity and quasi-parasitism were 7%, 7% and 1% of the offspring, respectively. Polygamy levels decreased with year, as the study population matured. The correlative study showed that males with longer TF and with better condition had a higher probability of reproducing and sired more offspring, but their offspring were not of higher quality as measured by their weight and immune response to phytohaemagglutinin. Polygynous males were also in better condition. In the experimental study, males in better condition had a higher chance of reproducing and sired more fledglings. Conversely, males with reduced TF sired significantly fewer eggs and lighter fledglings than control males. Body condition and TF length are shown to be good predictors of reproductive success and TF length is shown to be under sexual selection.

Chapter 1: Relative Concerns and the Choice of Fertility Empirical research has shown that people exhibit relative concerns, they value social status. If they value their children's status as well, what effect will that have on their decisions as parents? This paper argues that parents and potential parents are in competition for status and rank in the generation of their children; as a consequence richer agents may cut back on the number of children they have and invest more in each child to prevent children of lower income agents from mimicking their own children. This effect need not be uniform so that equilibrium fertility may e.g. be a U-shaped function of income, even when agents would privately like to increase fertility when they receive greater income. These findings have wide ramifications: they may contribute to our understanding of the working of the demographic transition; they also suggest that the low fertility traps seen in some developed countries are rather strongly entrenched phenomena; and they o er a new explanation for voluntary childlessness. Chapter 2: Relative Concerns and Primogeniture While pervasive in the past, differential treatment of children, i.e. different levels of attention and parental investments into children of the same parent, has become rare in modern societies. This paper offers an explanation based on technological change which has rendered the success of a child more uncertain for a parent who is deciding on how much to invest into each of his children. Within a framework of concerns for social status (or relative concerns), agents decide on how many children to have and how much to invest in each child. When their altruism towards each child is decreasing in the total number of children, it is shown that they may solve the trade-off between low investment, high marginal return children (that come in large numbers and hence hurt parental altruism) and high investment, low marginal return children (that come in low numbers) by demanding both types and hence practice differential treatment. Uncertainty over status or rank outcomes of children reduces the range of equilibrium investment levels intro children so that the difference in the numbers they come in is reduced. Eventually the concern for return dominates and differential treatment disappears. Chapter 3: Co-Evolution of Institutions and Preferences: the case of the (human) mating market This paper explores the institutions that may emerge in response to mating preferences being constrained in their complexity in that they can only be conditioned on gender not other characteristics of the carrier of the preferences. When the cognitive capacity of the species allows a sophisticated institutional setup of one gender proposing and the other accepting or rejecting to be adopted, this setup is shown to be able to structure the mating allocation process such that preferences evolve to forms that, conditional on the setup, are optimal despite the constraint on complexity. Nature can be thought of as delegating information processing to the institutional setup. In an application to humans it is shown that the mechanism of the model can help explain why men and women may exhibit opposed preferences in traits such as looks and cleverness. The anecdotal fact that women do not marry down while men do can be interpreted as a maladaptation of female preferences to modern marriage markets.

This thesis presents two studies where natural and sexual selection have interacted to evolve sexual behaviours. The thesis uses mathematical modelling to understand how these forces have caused each behaviour to evolve. This is useful because the results allow for reflection on the potential role of sexual selection in adaptation of these species to a changing environment. The first study is of early male arrival to spring breeding grounds in migratory avian species, this is termed protandry. The study explores the main hypotheses for avian protandry and then tests the susceptibility of each hypothesis to changing environment. The second study is of convenience polyandry in species where there is conflict over mating rate. Females have multiple strategies to avoid harassive males but strategies vary in cost and success rate; she must balance her strategy use to minimise her fitness depreciation. The study identifies the main factors that cause convenience polyandry to evolve and paves the way for future studies to investigate if sexual selection over resistance strategy provides these species a future advantage in adaptation to a changing environment.

In this thesis I set out to further our understanding of two broad questions, 1) why it is that sexual selection favours the evolution of particular traits and 2) why do the patterns of sexual selection on such traits differ between groups and populations? Specifically, I focused on the role that variation in intrasexual competitive structure, the non-random distribution of socio-competitive environments across individuals, plays in shaping variation in the operation of sexual selection both within and across populations. I explore the roles of three main determinants of competitive structure, namely; population structure, polyandry and non-random variation in the distribution of the intensity of competition. To achieve this, I used a combination of empirical and theoretical tools, using the model system Red junglefowl, Gallus gallus. Throughout this thesis I both develop and employ network quantitative tools as a framework to describe variation in intrasexual competitive structure. Overall, this thesis demonstrates a complex relationship between competitive structure and the operation of sexual selection. This structure can modify the strength and direction of sexual selection operating on phenotypic traits, obscure the operation of selection at the population level and influence the relative roles of pre- and post-copulatory sexual selection. Furthermore, this thesis explores how differences in local ecology can shape competitive structure itself and in turn shape sexual selection. In doing so, this thesis sheds some light on the role that variation in competitive structure may play in shaping the operation of sexual selection both within and between populations and generating the great diversity in sexually-selected traits and behaviours between populations.

The natural world consists of a vast array of forms, some more plentiful than others, yet our understanding of the processes responsible the production of biological diversity remains surprisingly limited. Here I combine novel datasets with powerful phylogenetic modeling techniques and computer simulations to test the effects of both biotic and abiotic factors on the dynamics of species radiations and the evolution of organism traits in birds. In the first part of this thesis, I develop our understanding of the importance of abiotic factors for diversification by showing that in the early stages of lineage diversification at least, rapid adaptation to novel climatic conditions is likely to represent a prominent driver of avian diversification. In the second part I concentrate on the role of biotic factors, in particular that of sexual selection. I show that not only is sexual selection associated with accelerated rates of speciation and secondary sympatry—as well as faster rates of net diversification across the entire avian tree of life—but also that across-species variation in rates of phenotypic evolution is best understood with reference to the focus and intensity of sexual selection. Finally, given that the relative importance of such processes appears to vary predictably across latitudes, in the final part of the thesis I argue that latitudinal differences in the speciation process offers a potentially powerful explanation for conflicting viewpoints regarding the contribution of speciation to high tropical diversity. Overall, this work provides fresh insight into the processes governing broad-scale patterns in biodiversity.

Elaborate morphological ornaments can evolve if they increase the reproductive success of the bearer during competition for mates. However, ornament evolution is incredibly rare in females, and the type and intensity of selection required to develop female-specific ornamentation is poorly understood. The main goals of my thesis are to clarify the relationship between the type and intensity of sexual selection that drives the evolution of female ornamentation, and investigate alternative hypotheses that might be limiting or contributing to the development of female ornaments. I investigated the ecology and evolution of female-specific ornaments within and between species of dance flies from the subfamily Empidinae (Diptera: Empididae). The dance flies display incredible mating system diversity including those with elaborate female-specific ornaments, lek-like mating swarms, aerial copulation and nuptial gift giving. To elucidate the form of sexual selection involved in female-ornament evolution, I experimentally investigated the role of sexual conflict in the evolution of multiple female- specific ornaments in the species Rhamphomyia longicauda. Through manipulative field experiments, I found that variation in the attractiveness of two ornaments displayed by females indicates that sexual conflict, causing a coevolutionary arms race, is an important force in the evolution of multiple extravagant female ornaments. Using R. longicauda again, I tested for a role of functional load-lifting constraints on the aerial mating ability of males who paired with females displaying multiple large ornaments. I found no evidence of functional constraints influencing the mating opportunities of elaborately ornate females, but instead discovered a relationship consistent with positive assortative mating for mass. Biased sex ratios are predicted to increase the intensity of sexual selection in a population, which in turn, is predicted to influence the evolution of ornamentation. I measured the incidence and prevalence of vertically transmitted symbiotic bacteria that has been observed to distort the sex ratio in other Dipteran hosts. While my survey revealed that symbionts occur at high incidence and variable prevalence across dance fly hosts, I found no effect of symbiont infection levels on population sex ratios, or female- specific ornament evolution. Further investigation into the relationship between sex ratios and female-ornament evolution using the comparative method revealed that the operational sex ratio (OSR) of a population did not predict continuous measures of female ornamentation across species. However, female-ornament evolution did predict male relative testis investment across species indicating that female ornaments likely indicate increased levels of polyandry. My thesis reveals that sexual selection theory developed to describe male-specific ornament evolution cannot readily be translated to apply to females. I show that male mate choice, rather than functional constraints or ecological associations with bacteria, is likely driving the evolution of female-specific ornaments. I also identify sexual conflict as an important selective force in the evolution of female-specific ornaments.

<p>QCR 20161026</p>