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Ulsd732423 Td Ines Dias.Pdf UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS The role of conspecific social information on male mating decisions “Documento Definitivo” Doutoramento em Biologia Especialidade Etologia Inês Órfão Dias Tese orientada por: Professor Doutor Paulo Jorge Fonseca Doutora Susana Araújo Marreiro Varela Professora Doutora Anne Elizabeth Magurran Documento especialmente elaborado para a obtenção do grau de doutor 2018 UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS The role of conspecific social information on male mating decisions Doutoramento em Biologia Especialidade Etologia Inês Órfão Dias Tese orientada por: Professor Doutor Paulo Jorge Fonseca Doutora Susana Araújo Marreiro Varela Professora Doutora Anne Elizabeth Magurran Júri: Presidente: ● Doutor Rui Manuel dos Santos Malhó, Professor Catedrático e Presidente do Departamento de Biologia Vegetal da Faculdade de Ciências da Universidade de Lisboa Vogais: ● Doutora Anne Elizabeth Magurran, Professor School of Biology da University of St Andrews (Orientadora) ● Doutor Paulo Jorge Gama Mota, Professor Associado Faculdade de Ciências e Tecnologia da Universidade de Coimbra ● Doutor Gonçalo Canelas Cardoso, Pós-Doutoramento CIBIO - Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do Porto ● Doutor Manuel Eduardo dos Santos, Professor Associado ISPA - Instituto Universitário de Ciências Psicológicas, Sociais e da Vida ● Doutora Sara Newbery Raposo de Magalhães, Professora Auxiliar Faculdade de Ciências da Universidade de Lisboa ● Doutora Maria Clara Correia de Freitas Pessoa de Amorim, Professora Auxiliar Faculdade de Ciências da Universidade de Lisboa Documento especialmente elaborado para a obtenção do grau de doutor Fundação para a Ciência e Tecnologia (SFRH / BD / 90686 / 2012) 2018 This research was founded by Fundação para a Ciência e Tecnologia through a PhD grant (SFRH/BD/90686/2012). With the institutional and logistical support of: This thesis should be cited as: Órfão, I. (2018). The role of conspecific social information on male mating decisions. PhD Thesis, Universidade de Lisboa, Portugal, 2018. NOTA PRÉVIA A presente tese contém capítulos já publicados (capítulo 4) e outros preparados para ser submetidos (capítulos 2, 3, e 6) de acordo com o Regulamento de Estudos Pós-Graduados da Universidade de Lisboa, E no Despacho N.º 4624/2012 do Diário da República II série nº65 de 30 de Março de 2012. A candidata realizou os trabalhos em colaboração, mas liderou e participou integralmente na concepção dos mesmos, desde a obtenção dos dados, análise estatística, discussão dos resultados e redação dos manuscritos. Lisboa, 26 de Abril de 2018 Inês Órfão Dias Acknowledgments Agradecimentos Ao longo do meu doutoramento tive a sorte de ter ao meu lado e de encontrar pessoas incríveis. A eles, agradeço todo o apoio! Em primeiro lugar agradeço aos meus orientadores. Ao Joca que me desafiou a entrar nesta aventura e que me apresentou ao mundo fascinante dos guppies. Agradeço- lhe também por ter sempre acreditado em mim. À Susana que esteve sempre atenta e disponível para discutir ideias, começando sempre por dar a sua opinião realçando o melhor de um trabalho, ideia ou projeto e só depois referindo os pontos a melhorar (isto fez toda a diferença!). To Anne I would like to thank, in first place, for accepting to be my supervisor even if she didn’t know me, for giving me the opportunity to work in her amazing research group in St Andrews where I learned SO MUCH, for questioning my initial ideas making me truly think and to leave preconceived ideas behind, for not giving me the answers and expecting me to search for them, for challenging me to do simpler but better. Por último, agradeço ao Paulo que aceitou tão simpáticamente “embarcar” nesta aventura e ajudar-me. Tive a grande sorte de ter como meus colegas o Miguel e o Alfredo. Estou eternamente grata por toda a disponibilidade que tiveram desde o início, quer fosse para me socorrerem no laboratório quando estava em apuros (lembras-te Miguel daquela vez que me esclareceste em relação àquela “fêmea” que estava ter um comportamento estranho?), quer para discutirem ideias, quer para “falar da vida”. Ao Alfredo agradeço ter sempre a porta do gabinete aberta (não me lembro uma única vez que me tenhas dito que estavas ocupado e que não me podias ouvir) e um café pronto a fazer, por me ter ensinado TANTO sobre apresentações e sobre escrita, pelo seu espírito crítico e pela sua i constante procura por fazer melhor. Ao Miguel agradeço também por ter estado sempre disponível para me ouvir e ajudar, para ler e reler (e ainda reler) os meus textos, por estar sempre a incentivar-me a fazer mais, por elogiar as minhas apresentações, e por me transmitir que um bom trabalho não pode ser só bom, tem antes que estar feito (escrito). À malta da EcoComp (Ana Lopes, Ângela, Antonieta, Constança, Daniela, Daniel, Joana, João, Manuel, Marilda, Rita) agradeço as perguntas feitas e as sugestões dadas durante as reuniões, pela boa disposição intervalada com discussões intensas acerca do tudo e do nada, e muito muito obrigada por estarem sempre dispostos a ajudar, especialmente nos “bloqueios” – agradeço em especial ao Manuel e ao Daniel. Ao pessoal de St Andrews (Maria João, Esme, Laura, Ale, Miguel Pinheiro, Luciele, Jess, Will, Shoko, Sónia, Miguel Neves, Sara, Gui, Mano, Maria, Tiago, Nuno, Manuel, Martim, Andreia, Claúdia) agradeço por me terem feito sentir em casa. Agradeço ainda a todas as pessoas com quem passei tantos bons momentos, quer fosse na pausa para o almoço, quer para o café e sem as quais não seria a mesma coisa ir à FCUL: à Ainara, ao Ricardo Rocha, ao Luis Borda D´Água, ao Sr. André, à Dona Elvira, ao Pedro e à Andreia do bar do c2, à Bé, à minha prima Carla, à Sara Lobo, à Cristina e ao Professor Francisco. À Professora Maria Turkman agradeço por se ter sempre mostrado disponível para me ajudar com a estatística. Finalmente agradeço àqueles que têm um papel fundamental na minha vida e que sempre se interessaram por o que faço e por mim (quer eu estivesse contente porque “tenho muita sorte em fazer o que gosto” ou infeliz “porque isto nunca mais anda”). À minha querida mãe que me ensinou a lutar pelo que quero, que sempre me apoiou e que teve paciência nos momentos mais difíceis. Ao Noel que esteve sempre ao meu lado, ii apoiando as minhas decisões e aceitando que estar ao lado de uma bióloga implica muitas vezes ter uma vida instável com garantias apenas por curtos períodos de tempo. À minha prima Paula e Rita, aos meus amigos Bruno e Rita Mateus que sempre se preocuparam acima de tudo com o meu bem-estar. Ao meu pai, tia Leonor, tio Heitor, prima Tânia e João por estarem sempre presentes na minha vida e por transmitirem que posso sempre contar com eles. Aos animais que não estão cá para me satisfazer a curiosidade, que simplesmente andam nas suas vidas, e por sorte eu gosto de observar. Ao Valter Hugo Mãe, ao Sebastião Salgado, e a tantos outros artistas que me fizeram sair do campo do racional, para voltar a ele. À ciência e lógica, e à humanidade e emoção que, de formas diferentes, nos permitiram chegar aqui. Que triste seria (…) se nós humanos viéssemos a perder a noção de mistério, todo o conceito de reverência. Se a parte esquerda do cérebro dominar um dia a direita, de forma a que a lógica e a razão triunfem sobre a intuição, o homem ficará totalmente alienado da sua mais profunda essência, do seu coração e da sua alma. Janne Goodall Em Motivo de Esperança iii Abstract Behavioural plasticity occurs when animals adjust their behaviour to current environmental conditions. Research suggests that this ability helps animals cope with changeable environments, especially in the social domain, where social information is highly variable and unpredictable. In this thesis I evaluated the contribution of the complexity of social information to: 1) male mating behaviours, 2) evolution of courtship display, and 3) evolution of dishonest communication. For male mating behaviours, I performed experiments with guppies (Poecilia reticulata). I predicted that males would invest more on mating attempts if they spend more time without mating (chapter 2), and when there was a higher probability of mating success or fertilization success (chapters 3 and 4). I found that time between encounters with females (not time between actual mating opportunities) was determinant to male investment. Moreover, males did not avoid encountering competitors but invested more when they were the first to arrive near females (not the last), and when competing against more attractive males (but only for orange colouration). These evidences support that males produce complex plastic responses in face of diverse social information. For the evolution of courtship display and dishonest communication, my hypothesis was that competitor (bystander) males use the courtship performance of other males as information about their competitive ability, and that displayer males adjust their behaviour accordingly. For this, I performed systematic reviews (chapter 5), where I found that male-male courtship display is, indeed, frequently associated with intrasexual competition, suggesting that courtship display has evolved a dual utility: attract females and intimidate competitors. A corollary of this, which I developed in a conceptual study (chapter 6), is that males can display dishonestly to deter competitors. If so, the iv interference of bystanders in communication systems could have a non-negligent role in the evolution of dishonest signalling. Keywords: animal communication, behavioural plasticity, male mating behaviour, sexual selection, social information. v Resumo A constatação de que animais de diversas espécies apresentam, em menor ou maior grau, a capacidade de ajustarem o seu comportamento face às condições ambientais encontradas (plasticidade comportamental) tem fascinado a comunidade científica. Consequentemente, vários estudos têm sido realizados no âmbito desta temática, nomeadamente partindo de três questões: como é que os indivíduos ajustam o seu comportamento face a diferentes componentes ambientais, se essa plasticidade comportamental traz benefícios para o seu fitness e quais as consequências evolutivas da plasticidade comportamental.
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