Implications for Conservation Helena Reis Batalha
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Variation within and among populations of the endangered Cape Verde warbler: implications for conservation Helena Reis Batalha A thesis submitted for the degree of Doctor of Philosophy School of Biological Sciences University of East Anglia, UK March 2017 © This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and that use of any information derived there from must be in accordance with current UK Copyright Law. In addition, any quotation or extract must include full attribution. This thesis is dedicated to Cornelis J. Hazevoet, who did so much for ornithology in Cape Verde and for the Cape Verde warbler in particular. ii Abstract In this thesis, I investigate variation among the three remaining populations of the Cape Verde warbler Acrocephalus brevipennis, with the aim of defining conservation units and thus informing conservation. I used neutral molecular markers to assess genetic diversity and infer the evolutionary history and adaptive potential of the populations. I found a gradient of diversity from the largest population of Santiago to the smallest one of S. Nicolau, which has been isolated from the populations of Santiago and Fogo for longer than those have been isolated from each other. I measured differences in male song between the populations and whether individuals behaved differently when exposed to songs from the other populations, to determine if song might play a role in reproductive isolation. While there is high intra- population variability in male song, birds do not react differently to songs of males from other populations. I then investigated which habitat traits determined the presence of the bird and whether this differed between the three islands. Evergreen dense vegetation cover predicted the wider areas occupied by the warbler. Structural vegetation traits did not differ between sites used on the islands, but plant species composition did. Finally, I assessed morphological differences between the three populations. I found that males on S. Nicolau have shorter tarsi and longer, narrower bills. These morphological differences match the pattern of neutral genetic divergence, suggesting that the role of drift in driving these differences has not been overridden by selection. Collectively, my results show small but significant divergence between the three populations, particularly between S. Nicolau and the two other populations. This divergence has implications for this bird’s conservation, as the differences observed warrant considering the populations as different management units. iii Acknowledgements This PhD was a great adventure, both professionally and personally. It would not have been completed without the help of a huge number of people who gave their whole-hearted contribution at specific times. First of all I would like to thank Sara Pardal for sending me that email, back in 2012, telling me about a researcher at the University of East Anglia who was looking for a student to do a PhD on the Cape Verde warbler. Sara, if it weren’t for you, I would have never known about this opportunity and I would have missed what was possibly the most exciting job I had in my life. I am incredibly grateful to you! Secondly I would like to thank my excellent supervisory team: David Richardson, Iain Barr and Nigel Collar. David, you have been a great inspiration and have made me a better person and a better professional. Thank you for your persistence, determination, optimism and support. Iain, thank you for your enthusiasm, great ideas and constant support. Nigel, thank you for always finding time to answer my questions and soothing my anxiety, even though you are an incredibly busy man. All of you are really intelligent, have excellent work ethics and are incredibly professional, which contributed enormously to make this PhD thesis the best possible one. To my friend and colleague David J. Wright, what would have I done without you? You contributed officially to the chapter on genetic diversity, and unofficially to all the other chapters because you helped me with R, you reviewed a lot of my drafts and helped me made sense of incipient ideas. Thank you for your support, your optimism, for having attention to detail, and for always having time for me and an opinion about my work. Paul Donald, thank you for your help with trying to find this little brown bird on S. Nicolau and for suggesting and providing guidance on the the MaxEnt analysis that we used in the habitat chapter. Ricardo Jorge Lopes and Raquel Vasconcelos, thank you for your help and guidance in organising fieldwork and input on the genetics chapter. Gonçalo Cardoso, thank you so much for your help with song analysis and statistics, and for always trying to answer my every question. Sérgio Timóteo, thank you for always taking the time to answer my stats questions. Eduardo Garcia-del-Rey, thank you for putting me in touch with the Museum of Oslo and for allowing me to contribute to your book on the Cape Verde birds. Juan Carlos Illera, Martim Melo, Martin Haase, the University of Oslo Natural History Museum and the Seychelles Warbler Research Group for extra CVW samples; Silke Fregin, Urban Olsson, Juan Carlos Illera, Lewis Spurgin for input on molecular work for the genetics chapter. iv Cornelis J. Hazevoet, the person who most studied this bird before we did, thank you for all our meetings at Cister, where we discussed at length the Cape Verde warbler and how it is to live and work in Cape Verde, and for all the emails and papers you sent. Tommy Melo from Biosfera I and Pedro Geraldes from SPEA, thank you so much for helping me get started with my fieldwork in Cape Verde, for getting the Cape Verdean ringing scheme going, for all the advice and for finding me a place to stay – and Pedro, in your case, for letting us stay at your place in Mindelo when you were not there! Aline Rendall, Alexandre Nevsky Rodrigues, João Mascarenhas, Lindaci Oliveira and Emitério Ramos, thank you very much for supporting this project and for helping with everything you could during our fieldwork, especially for finding us places to stay that we could afford with our limited budget! Without your help, we would have been unable to do this much work in the field and this PhD would have not been finished! Also a big thank you to all the Cape Verdean biologists who helped us during fieldwork: Gilson Semedo, Evaldino Pereira, Gabriel Tavares, Herculano Dinis, Maria “Chia” Auxiliadora do Nascimento, Nelson Santana, Franciscana Sequeira, Ivani Duarte, Gilda Monteiro and others. Thank you for your interest, for your help in finding the birds, especially on S. Nicolau, for setting the mist-nets, for your advice and for your friendship and enthusiasm! Celestino Duarte, this work would have been severely impaired without your help. Thank you for letting us stay at your studio for three weeks when we first arrived in Cape Verde, for finding us and renting us that big black jeep, and for always trying to help with bureaucracy. I would like to send special thanks to the fantastic people of Mosteiros, Pai António and Monte Barro, Fogo: Ilda, Fidjim, Nene, Manu, Tony, Rui Fontes, Batinzim, Dona Christine, and so many others. You made us feel home during our time there, made our stay there much cheaper and therefore longer! Special thanks to Nene for showing us all those Cape Verde warbler territories. Also special thanks to Xexe Kifra, our “Christmas miracle”, the man who was on his balcony when we stopped at the ATM machine on our way to Chã das Caldeiras on Christmas day. Thanks to you, our fieldwork group got to spend our Christmas evening on that magical place! Now the part that always brings a silly tear to my eye: thank you very much to my incredible, wonderful, fantastic field assistants! It was a great pleasure to work with you, thank you for putting up with me, for always doing your best and for your friendship. Jaelsa Moreira and Naya Sena, my Cape Verde field assistants, two amazing women and good friends, you will always be in my heart! Jaelsa, thank you for your positive and rational spirit, for giving this project your best, for being a good friend and for partying with me on Brava! Naya, I have no v words, you are such an inspiring woman and great friend, professional, curious and motivated, and always helped me in every possible way, whether it was to find us a place to stay or an extra field assistant. Thank you to Naya’s mum Maria dos Reis “Nunui” for letting us stay at your place for such a long time when we had to work on Santiago, for your inspiring calm and wise words. Gilson Montrond, I only met you during the last field season but you were also great, with your eagle eye that never missed one bird and your tree-climbing abilities. Andrew Power, my first field assistant ever, thank you for meeting me at the airport of Praia at 2am when neither of us knew what was going to happen, for putting up with my very emotional this-is-amazing mood during two months (Torbjörn did not suffer that much haha), for always doing your best and contributing in every way possible to the success of this project. Josh Jenkins Shaw, thank you for your help and professionalism, especially on Fogo and with ageing the birds. Torbjörn Blixt aka “Golden Magnus” thank you for coming with me everywhere, even when we took that hike on Santo Antão on your birthday and did not find anything to eat (only two coca colas to drink) until night time! Thank you for helping define the methodology for the habitat chapter, for being professional, accurate, reliable and always in a good mood, even when you got sea sick, and for making us laugh by astonishing all the villagers who had never seen such long hair and so many tattoos! And finally Jeroen Arnoys who came to help twice, thank you for doing everything you could to make this project a success, from setting mist-nets to buying that thank you gift for Nunui.