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From Individuals to Ecosystems From Individuals to Ecosystems A Study of the Temporal and Spatial Variation in Ecological Network Structure Submitted by Dominic Charles Henri to the University of Exeter as a thesis for the degree of Doctor of Philosophy in Biological Sciences In June 2014 From Individuals to Ecosystems This thesis is available for Library use on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I certify that all material in this thesis which is not my own work has been identified and that no material has previously been submitted and approved for the award of a degree by this or any other University. Signature: Dominic C Henri 2 D.C.Henri: Thesis for the degree of Doctor of Philosophy in Biological Sciences Abstract Ecological network theory has developed from studies of static, binary trophic relationships to the analysis of quantitative, dynamic communities consisting of multiple link-types. Particularly, work has focused on the dynamic nature of ecological networks, which maintains stability in complex communities. However, there are few in situ network-level studies of the determinants of temporal and spatial variation in community structure. This thesis utilises data from a 10-year study of a host-parasitoid network and a collaborative study in an applied ecological setting to identify individual level factors important to network structure. The work aims towards an empirical, predictive framework linking adaptive foraging behaviour to ecological network structure. The results show that condition-dependent foraging behaviours structure host- parasitoid networks. The realised niches of the studied parasitoid species were generally biased towards larger host species and condition-dependent sex ratio allocation increased the likelihood that females would eclose from relatively larger hosts and males from relatively smaller hosts, which resulted in sex ratios deviating from Hamiltonian (50:50) predictions. Further, both of these aspects of behaviour are plastic, where parasitoid behaviour responded to environmental heterogeneity. Particularly, host preference behaviour conformed to an egg-/time-limitation framework, where the size dependency of the behaviour is greater when individuals have a greater likelihood of being egg-limited. Both the size-dependency and the plasticity of these behaviours differed significantly between secondary parasitoid species. This species identity effect interacted with landscape heterogeneity, which 1 From Individuals to Ecosystems may explain some inter- and intra-specific variation in network structure. With respect to applied ecology, the results show that the benefits of natural vegetation for pest control are dependent upon the dispersal capabilities and the diet breadth of the pest and its natural enemies. The findings are evaluated towards a predictive framework for understanding the effects of future climate change on community structure and stability. We consider this framework in terms of applied ecology, particularly pest control ecosystem services provided by natural vegetation in an agricultural environment. The synergistic nature of the multiple determinants of network structure found in this thesis suggest that future studies should focus on the whole network, which is not necessarily the sum of its parts. 2 D.C.Henri: Thesis for the degree of Doctor of Philosophy in Biological Sciences Acknowledgements Firstly, I would like to thank Frank van Veen for his guidance and trusting me with important academic opportunities that have truly enriched my PhD process and for giving me the freedom to pursue some ambitious projects. I would like to thank the other academics who collaborated on the reports herein: Colleen Seymour, Carolina Reigada, Dave Hodgson, Dave Shuker, Elisa Thébault and Sabrina Araújo. Each of you has dramatically improved the quality of this thesis and has been a pleasure to work with. I count you among the best persons and scientists that I know and I apologise for increasing your work-load with constant manuscript redrafts. Further, thanks to Juliet Osborne and Rebecca Morris for agreeing to be my examiners. I appreciate the aid of all of the students who helped with the data collection for this thesis. The methodologies ranged from the truly repetitive to the nauseating, but each of you showed utmost character and has earned your authorships on the various chapters. Thanks to my colleagues, Chris Coles, Jenni McDonald, Sally Luker, Dirk Sanders and Katy Scott, whose empathy with the process has ensured my sanity. I would also like to thank Rob Wilson for his mentorship and friendship during my short-tenure as a lecturer. Thanks to NERC UK for funding this project and to the University of Exeter, the British Entomological Society and the British Ecological Society for various funding for the untold travels I have made during this process. Thanks to my friends and family who managed to visit me in the beautiful corner of the world and from whom my character, ambitions and self-assurance manifest. 3 From Individuals to Ecosystems My dearest Louisa, although your name is not on any author list this is as much your success as it is mine. Now that it is finally finished with; I have time to do something that I have wanted to do for a very long time... To my foundation For each reason here and moral compass, and so many more, empathic lodestone everything you are and sense of purpose. and that you stand for. So full of beauty, I ask one question, within and without. of my hoped-for wife. I love you truly Can my profession and without doubt and one task for life you are the strongest be to be your star, person that I know. forever aglow? Louisa Rose Briggs, will you marry me? 4 From Individuals to Ecosystems “It always seems impossible until it is done” 2 D.C.Henri: Thesis for the degree of Doctor of Philosophy in Biological Sciences Table of Contents Abstract ...................................................................................................................... 1 Acknowledgements .................................................................................................... 3 Table of Contents ....................................................................................................... 3 Table list ..................................................................................................................... 7 Figure List .................................................................................................................. 8 Publications resulting from this thesis ...................................................................... 12 Chapter 1 Body Size, Life History and the Structure of Host-Parasitoid Networks ... 13 1.1 Abstract ........................................................................................................... 13 1.2 An Introduction To Ecological Network Theory And Host-Parasitoid Networks .............................................................................................................................. 14 1.3 The Aims of This Review ................................................................................ 22 1.4 The Structuring Of Trophic Networks .............................................................. 23 1.5 Limitations On Host Range ............................................................................. 30 1.6 Life-History and Host Range ........................................................................... 32 1.7 Realised Niche In Parasitoids ......................................................................... 35 1.8 Parasitoid Life-History And Host Electivity ...................................................... 44 1.9 Sex Allocation and Host Quality ...................................................................... 57 1.10 Conclusions .................................................................................................. 60 1.11 Thesis Chapter Descriptions ......................................................................... 76 1.12 Ackowledgements ......................................................................................... 78 Chapter 2 Potential for climate effects on the size-structure of host-parasitoid indirect interaction networks ................................................................................................. 79 2.1 Chapter Concept ............................................................................................. 79 3 From Individuals to Ecosystems 2.2 Abstract ........................................................................................................... 79 2.3 Introduction ..................................................................................................... 80 2.4 Methods .......................................................................................................... 86 2.5 Results ............................................................................................................ 91 2.6 Discussion....................................................................................................... 95 2.7 Acknowledgments ......................................................................................... 100 Chapter 3 Egg-/time- limitation and condition-dependent foraging behaviour: Mechanisms driving the dynamic nature of ecological networks ............................ 102 3.1 Chapter Concept ..........................................................................................
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