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Ecology and Biogeography of Island Parasitoid Faunas

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Ecology and Biogeography of Island Parasitoid Faunas Ecology and biogeography of island parasitoid faunas Ana Margarida Coelho dos Santos Department of Life Sciences, Division of Biology, Silwood Park Campus, Ascot, Berkshire, SL5 7PY, U.K. A thesis submitted for the degree of Doctor of Philosophy of Imperial College London March 2010 1 2 Abstract Islands constitute natural laboratories for the study of evolutionary and ecological processes due to their discrete and isolated nature. Island biotas tend to be species–poor and disharmonic compared to the mainland; typically, interspecific competition is low, and entire groups of predators, parasitoids or pathogens are absent from their biotas, so the ecological space is often not fully saturated. Consequently, species from island assemblages often use a wider range of resources than their counterparts from the source mainland. Here, I investigate whether island parasitoid communities have proportionally more generalist species than their source mainland, and which factors determine island community structure. These questions were approached using data on the distribution of Ichneumonoidea (Hymenoptera) species worldwide and with data from a survey conducted in the Macaronesian region. Prior to the global analyses, I assessed whether islands and archipelagos follow the same species–area relationship, and identified which islands have comparable inventories. Globally, islands have proportionally more idiobionts (i.e. generalists) than continental areas. However, there is a latitudinal gradient in the level of generalism of island parasitoid faunas that correlates with some environmental factors and island characteristics; the species pool is the most important determinant of island community structure, together with temperature (for braconids) or biogeographical region (for ichneumonids). Host and parasitoid larvae collected in different islands of the Macaronesian region and adjacent mainland were assigned to Molecular Operational Taxonomic Units using a protocol based on host dissection and DNA barcoding. At this scale, mainland faunas have proportionally more koinobiont species and island communities have a greater proportion of idiobionts. Although overall parasitism rates were similar between islands and mainland, islands had higher idiobiont parasitism rates than expected by chance. In summary, results from this thesis indicate that indeed island parasitoid faunas are biased towards generalist species. 3 4 Declaration The work presented in this thesis is entirely my own and it has not been submitted for any other academic qualification. Co-authors on published and submitted chapters are listed as footnotes at the start of each chapter. People who provided less formal help and advice are listed in the acknowledgements. Ana Margarida Coelho dos Santos 5 6 Acknowledgements I would like to thank the many people that have helped me come this far. Unfortunately, naming all of them would be a thesis on its own, and that is something I do not want to repeat... so I will only mention some of them: My supervisors Donald Quicke and Paulo Borges, for their advice and guidance during these last years. Joaquín Hortal, whilst not officially my supervisor, provided me with many advices, and supported me through the good and the rough times. Without his love and help it would have been very difficult to “survive” this thesis. Guillaume Besnard, Colin Fontaine, Owen Jones, Kostas Triantis and Rob Whittaker were directly involved in some of the chapters, giving me invaluable ideas and advices on how to improve my manuscripts. Other people contributed with comments and suggestions for this thesis: Brad Hawkins, Jorge Lobo, Mick Crawley and Simon Leather discussed and commented on some of my ideas; Gavin Broad and Mark Shaw helped me build the database on the biology of the Ichneumonoidea; Shai Meiri helped me put together the islands database; Isabel Amorim, Marianne Elias and Hanno Schaefer gave me many advices on the practicalities of molecular work and commented on the chapter that resulted from that nightmare. Many others helped me during field work, making it an extraordinary experience: António Franquinho Aguiar, Mário Boieiro, Raimundo Cabrera, Abdelhaq Hanafi, Alberto Jiménez- Valverde, António Machado, Glória Ortega, Fernando Pereira, Carla Rego, Micra and Mustafá; also Isabel Amorim, Paulo Borges, Joaquin Hortal and Jorge Lobo. Many thanks to those who helped obtain all the permits for fieldwork: Ángel Fernández, Cristina Fuentes, Felix Medina, Cristina Padrón, Ángel Palomares, Pedro Romero and Marina Sequeira. I am in debt with Marcos Báez and Ole Karsholt for providing specimens, Joaquín Baixeras for identifying some tortricid specimens and for providing specimens, and Winnie Hallwachs, Daniel Janzen and Alex Smith for allowing me to use some of their sequences. 7 I would like to thank Rebecca Fuller, Rebecca Jeffrey and Adila Shaukat for their support in the laboratory, and Ambika and Nikos Karatolos, for helping me with the crazy experimental islands. I am also grateful to Christine Short for dealing with all the big list of orders I was always placing. My friends from Silwood have prevented me from going completely crazy. There are no words to express my gratitude to: Ana (s), Amalia (also for proof-reading my thesis), Andrés, Aurélio, Becca, Becky, Bish, Christos, Diego, Eftychis, Francesca, Guillaume, Hanno, Joana, Manju, Márcia, Meirav, Moh, Nagore, Navinder, Nikos, Omar, Oriana, Shai, Shan, Yael…. Many thanks to my friends in Lisbon for their support, the online chats, and mostly for putting up with me: Andreia, Catarina, Hugo, Inês M., Inês P., Isabel, Paulo, Rita, Sérgio, Sofia G. and Sofia L. (also for proof-reading my thesis). Special thanks to Sofia G., for all the time we spent talking online about molecular work, and to Hugo and Paulo, for drying my tears… I would like to thank my family, specially my parents and brother, for their love and understanding. Quero agradecer à minha familia, especialmente aos meus pais e ao Pedro, por todo o apoio que me deram desde que decidi ser bióloga... o doutoramento foi apenas mais uma etapa das muitas em que sei que posso contar sempre com eles, mesmo que o meu trabalho seja para eles um mistério. Finally, I want to thank again to Joaquín, for his infinite patience, his positivism and for trying to make me see “the bright side of life”. The Portuguese Foundation for Science and Technology (FCT, Fundação para a Ciência e Tecnologia) provided financial support for my project (SFRH/BD/21496/2005). The International Biogeography Society and the Royal Entomological Society provided travel grants for attending some congresses. 8 Contents Abstract ............................................................................................................................... 3 Declaration .......................................................................................................................... 5 Acknowledgements ............................................................................................................. 7 Contents ............................................................................................................................... 9 List of Tables ............................................................................................................ 13 List of Figures ........................................................................................................... 15 Appendices ............................................................................................................... 17 Chapter 1: General Introduction ...................................................................................... 19 1.1. Parasitoids .......................................................................................................... 19 1.1.1. Biology ............................................................................................... 19 1.1.2. Idiobionts versus koinobionts ............................................................. 20 1.1.3. Host range ........................................................................................... 22 1.1.4. Diversity ............................................................................................. 22 1.1.5. Dispersal, colonization and islands .................................................... 25 1.2. Islands and their communities ........................................................................... 26 1.2.1. Island types ......................................................................................... 26 1.2.2. Equilibrium model of island biogeography ........................................ 27 1.2.3. Island species–area relationship (ISAR) ............................................ 28 1.2.4. Dispersal and colonization ................................................................. 30 1.2.5. Ecological consequences of empty niche space ................................. 31 1.2.6. A model island system: the Macaronesia ........................................... 32 1.3. Thesis aims and outline ..................................................................................... 35 Chapter 2: Are species–area relationships from entire archipelagos congruent with those of their constituent islands? .................................................................. 39 2.1. Abstract .............................................................................................................. 39 2.2. Introduction ......................................................................................................
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