UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS DEPARTAMENTO DE BIOLOGIA ANIMAL PREDATOR-PREY INTERACTIONS BETWEEN AN INVASIVE CRAYFISH PREDATOR AND NATIVE ANURANS: ECOLOGICAL AND EVOLUTIONARY OUTCOMES ANA LUÍSA SUMARES DA CRUZ NUNES DOUTORAMENTO EM BIOLOGIA ESPECIALIDADE DE ECOLOGIA 2011 UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS DEPARTAMENTO DE BIOLOGIA ANIMAL PREDATOR-PREY INTERACTIONS BETWEEN AN INVASIVE CRAYFISH PREDATOR AND NATIVE ANURANS: ECOLOGICAL AND EVOLUTIONARY OUTCOMES ANA LUÍSA SUMARES DA CRUZ NUNES TESE ORIENTADA POR: DR. RUI MIGUEL REBELO DR. ANSSI LAURILA DOUTORAMENTO EM BIOLOGIA ESPECIALIDADE DE ECOLOGIA 2011 This study was funded by Fundação para a Ciência e a Tecnologia (FCT) through a PhD grant (SFRH/BD/29068/2006) and by the research project POCI/BIA-BDE/56100/2004. This dissertation should be cited as: Nunes AL (2011) Predator-prey interactions between an invasive crayfish predator and native anurans: ecological and evolutionary outcomes. PhD Thesis, Universidade de Lisboa, Portugal. Nota Prévia A presente tese apresenta resultados de trabalhos já publicados ou em preparação para publicação (capítulos 2 a 5), de acordo com o previsto no nº 1 do artigo 41º do Regulamento de Estudos Pós-Graduados da Universidade de Lisboa, publicado no Diário da República II série nº 209 de 30 de Outubro de 2006. Tendo os trabalhos sido realizados em colaboração, a candidata esclarece que participou integralmente na concepção dos trabalhos, obtenção dos dados, análise e discussão dos resultados, bem como na redacção dos manuscritos. Lisboa, Novembro de 2011 Ana Luísa Sumares da Cruz Nunes Aos papás e à mana ACKNOWLEDGMENTS After almost five years of hard work, finally this thesis comes to an end. I guess one of the things that it made me realise is my unbelievable capacity of never being able to achieve deadlines. However, it also allowed me to become a mini-scientist and to get to experience things that will forever be in my mind and that make me who I am today. I start by thanking my supervisors, Rui Rebelo and Anssi Laurila. To Rui, for convincing me to apply for a PhD grant, even though I was very reluctant to start a life as a researcher at the time. I turned out to be right, science is way too scientific for me, but I cannot be happier for having done this PhD. Thank you for teaching me so many things about the natural world, for going with me to the field every time I needed, for your constant good mood but especially for, other than a supervisor, being a true good friend. To Anssi, the nicest Finnish person I have ever met, thank you for promptly accepting me as your half PhD student and for allowing me to work in Uppsala for two years. Thank you for all the chats in your office about work and many other things, from wolves and bears to the horrible Swedish weather. And for always being so patient about me handing in work late. Thank you both so much for helping me make this thesis a nice scientific work, but especially for being always present, available and extremely funny supervisors. Throughout the discussion of the thesis I use the personal pronoun “I”, but this work is really “ours”. To Fundação para a Ciência e Tecnologia I thank for my PhD grant (SFRH/BD/29068/2006) and for funding through the Project POCI/BIA-BDE/56100/2004. I also thank Stiftelsen för Zoologisk Forskning for funding. To Dr. Miguel Tejedo for having received us so well in Seville and having allowed us to perform fieldwork in Doñana. To my master students, Erika, Susana and Cátia, for making the huge experiment in the Herdade a doable thing and for going to Grândola, even at weekends, to change waters and feed the predators. Thank you for all your help in the field and in the lab. To Pedro Andrade, the best field assistant ever, for keeping me company in Grândola so many times, for being so good in fixing problems and in building weird necessary things and for talking about crazy nonsense sometimes. You made my fieldwork extremely pleasant and easy. To Sandra Amaral for making me go to Rui’s office many years ago and help trace a way into studying amphibians. For going with me through the hard process of learning about amphibian larvae (“Is this an Alytes or a Pelobates?”). To Maria João, for teaching me almost everything I know about amphibians and for calling me to Sweden when I needed help with statistics. Both you and Mário have been really nice friends. To Helder, an amphibian companion, for the help with the phylogenetic trees, but mostly for sharing the love for amphibians and always being ready to talk about them. Thank you for always being ready to help and for hosting me in Seville during the congress. To all the friends I made in Sweden, thank you so much for having made life in the north a, not only bearable, but extremely nice thing. You have allowed me to experience many incredibly nice and different situations and to live the real grad student life. To everyone in the department, thank you for making work a very nice place, filled with friends. Thank you for the thursday pancakes, sushi lunches, friday fikas and Christmas dinners. To Marianne, for all the help in solving ‘money issues’, you were great! To Jan L, for always asking me about the thesis and for giving me nice hints on how a paper should be written. To the frog group, Emma, Björn, Magnus, Rike, Sandra H, Katja, German and Alex, thanks for sharing your knowledge with me and for being nice friends. To Rike, thanks for being such a lovely office mate and for letting me feed your snails. To the people from the Iberian (not Spanish) office, Violeta, Maria, Germán and Alex, for always speaking in english with me and for all the healthy gossip. To Germán and Alex, thank you for all the help in my papers and for showing me the wonders of doing fieldwork with frogs in Sweden. To Alex, for all the chocolates in the fika room (no more oranges for you!), for the company in the office when no one else was there and, especially, for having become a very good friend. To the Costa Rica crew, Biao, Jonathan, Jenny and Caro, for an amazing trip to the other side of the world. Thank you, Biao, for being such an easy going person and for always being in a good mood. Thank you, Jonathan, for being my friend, despite I though you didn’t like me in the beginning and for showing me so many birds in Costa Rica. You are lovely! Jenny, the atypical Swedish girl, thank you for teaching me how to do kannelbulle and for all the shared lunches outside. To Caro, thank you for all the time spent together, during and after work, for always liking the things I cooked, for letting me get to see a black grouse (or many) and, especially, for the very nice friendship that we have. To other people from EBC, Jürg, Maria, Omar, Yoshi, Christina, Kasia, Anna, Richard and Andreas for all the dinners, parties and beers in the Nations. To Kasia, the fortunately more PopBio than AnimEcol girl, thanks for being so nice and caring and for making me understand how much a real scientist can love her job. To my Portuguese family in Uppsala, Tiago and Vanda, for being very close and good friends and for allowing me to not forget my mother tongue while I was away. To my ‘Colombian’ family, Daniel C, Daniel O, Tobias, Angie, Caro, Jessica, Ricardo, Manda, Tanai, Sahar, Tobi, Anna, Ola, Claire and Sara for all the friendship and for making my stay in Uppsala much better than I could wish for. Thanks for all the shared moments, especially in our amazing trips to Sälen and Colombia. Muitossshhh obrigadossshhh, from radiosssshh portuguessshhhh. To Caro, Angie and Tobias, for always being there for me. To my dearest friends from the “Sala dos terminais”, Ana Leal, Nuno, Tatá, Mafalda, Joana, Miguel and Luís, thank you for making it really nice to go to work, even at the final stages of this thesis (especially when it was a cake day). Thank you for giving me all the strength I needed to finish this, you were the ones who made it possible for me to hand this in now. I especially thank you for constantly asking if I needed help and for the last minute reviewing of the thesis. To all my korfball people, for often making me think of things other than work, for all the holidays spent together and for, after all these years, still not understanding what my job was. Ralha and David, now you can finally see what ‘your’ money was used for. To my twins, Sara and Ana, thank you for coming to Grândola to help me in the field. Xiquinho, thank you for, even when I was far, always being close through the internet. To Joana Vinagre, Henrique, Natércia and Sónia for coming all the way from Portugal to experience the ‘warm’ Swedish summer, making my first year in Uppsala much nicer, at a time I wanted to come back to Portugal. To Natércia, for always asking me about the thesis and for being there, even if through a phone call, every time I needed. To Sónia, thank you for your help changing waters and feeding tadpoles and for all the times you stayed at home keeping me company while I had to work. To Joaninha and Ricardo, the ever present university friends, for the solid friendship and for always encouraging me to finish this fast.