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Effects of Tree Species Diversity on Insect Herbivory

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Effects of Tree Species Diversity on Insect Herbivory Effects of tree species diversity on insect herbivory A thesis submitted to the University of London in partial fulfilment of the requirements for the degree of Doctor of Philosophy By Simon Morath School of Biological Sciences Royal Holloway, University of London Egham, Surrey, TW2O 0EX United Kingdom March 2013 1 Declaration of Authorship This thesis and the work presented herein is my own work, conducted whilst enrolled in the School of Biological Sciences as a candidate for the degree of Doctor of Philosophy. This work has not been submitted for any other degree or award in any other University or educational establishment. Where I have consulted the work of others, this is clearly stated. Signed: ______________________ SIMON MORATH Date: ________________________ 2 Acknowledgments I would like to thank my supervisor, Professor Julia Koricheva for support and providing her scientific opinion over the last 4 years; the time she has spent reading and commenting on Chapter drafts has helped improve the quality of the thesis and is greatly appreciated. I’d also like to thank Professor Alan Gange for his enthusiastic and motivating interjections. A debt of gratitude is owed to David Lloyd, Naomi Pankhurst, Craig Williams, Victoria Goad, Gerlinde Schaeffter, Shaimalli Roy and Paul Rassell who all helped out to varying degrees with field work; to this end thanks also go to, Iiris Kallajoki, Juho Yli-Rosti, and Kai Ruohomäki. The research reported here was conducted as part of the European BACCARA and FunDiv projects, which received funding from the European Commission’s Seventh Framework Programme (FP7/2007-2013), under grant agreement number 226299. 3 Abstract It is generally believed that tree species growing in mixed forest stands are less susceptible to insect herbivore damage than if grown in monocultures, but previous studies have been largely observational and focussed mainly on tree species richness effects. In this thesis, I examined effects of three components of forest diversity (tree species richness, intraspecific genotypic diversity and functional diversity) on insect herbivores using three long-term forest diversity experiments in Finland and Germany. I have also explored the sources of variation in and the mechanisms behind the effects of tree diversity on insect herbivores. I found that all three components of forest diversity significantly influenced insect herbivore abundance and damage. Tree species richness effects depended on the insect herbivore feeding guild, but also changed within season and between years. As a result, silver birch (Betula pendula) experienced both associational resistance (reduced damage in mixed stands) and associational susceptibility (higher damage in mixed stands) to different insect herbivores and in some instances this altered temporally. In contrast, tree species richness effects on insect herbivory were spatially consistent and not mediated by tree size (physical apparency), physical properties of leaves or natural enemies. Interestingly, tree species richness and genotypic diversity had opposite effects on leaf miners; leaf miner abundance and species richness were lower in species-rich stands, but higher in mixtures containing several genotypes of silver birch. To test the effects of tree functional diversity, I created a functional diversity index based on constitutive emissions of monoterpenes and isoprene by different tree species and showed that tree species which emitted low levels of volatiles experienced associational resistance in stands with high diversity of volatile emissions. This suggests that increasing chemical complexity in mixed stands may interfere with host finding ability of herbivores. 4 Title page................................................................................................................... 1 Declaration................................................................................................................ 2 Acknowledgements................................................................................................... 3 Abstract...................................................................................................................... 4 Contents..................................................................................................................... 5 List of Tables............................................................................................................. 8 List Figures................................................................................................................ 9 List of Plates.............................................................................................................. 11 List of Abbreviations................................................................................................ 11 Chapter 1 …………………………………………………………………………. 12 1.1 General introduction............................................................................................. 12 1.1.1 Approaches to study forest diversity effects on insect herbivory............... 14 1.1.2 Components of forest diversity and their role in control of insect herbivory.................................................................................................... 14 1.1.3 Mechanisms of forest diversity effects on herbivores................................. 17 1.1.4 Thesis aims.................................................................................................. 23 1.1.5 Structure of the thesis.................................................................................. 23 Chapter 2 Experimental sites and insect monitoring……………………….. 25 2.1 Experimental sites and experimental design........................................................ 25 2.1.1 Tree species diversity experiment, (Satakunta, Finland)............................ 25 2.1.2 Genotypic forest diversity experiment (Satakunta, Finland)...................... 28 2.1.3 BIOTREE experiment (Thuringia, Germany)............................................. 29 2.2 Insect herbivory monitoring protocol................................................................... 31 2.2.1 Satakunta tree species diversity experiment................................................ 31 2.2.2 Satakunta genotypic diversity experiment.................................................. 33 2.2.3 Herbivory monitoring protocol at Bechstedt............................................... 33 2.3 Tree growth and leaf traits................................................................................... 34 Chapter 3 Tree species diversity effects on insect herbivory on silver birch in the boreal forest zone................................................................... 35 3.1 Introduction.......................................................................................................... 35 3.2 Methods................................................................................................................ 38 3.2.1 Data analysis............................................................................................... 40 3.3 Results.................................................................................................................. 42 3.3.1 Chewing damage......................................................................................... 44 3.3.2 Skeletonisers................................................................................................ 45 3.3.3 Leaf galls..................................................................................................... 45 3.3.4 Leaf rollers.................................................................................................. 47 3.3.5 Silver birch aphid (Euceraphis betulae)..................................................... 47 3.3.6 Total insect herbivore damage.................................................................... 49 3.3.7 Predators..................................................................................................... 50 3.3.8 Leaf characteristics...................................................................................... 51 5 3.3.9 Tree growth characteristics......................................................................... 52 3.4 Discussion............................................................................................................ 52 3.4.1 Tree species diversity effects on herbivores............................................... 52 3.4.2 Mechanisms of forest diversity effects on insect herbivores, the influence of predators, physical leaf traits and tree apparency................................... 55 3.5 Conclusions.......................................................................................................... 59 Appendix 3.1-ANCOVA: Birch herbivory with leaf trait covariates...................... 61 Appendix 3.2A -ANCOVA: Tree growth traits as covariates (2010)...................... 62 Appendix 3.2B -Assumptions of independence (tree growth traits)....................... 63 Chapter 4 Effects of tree species diversity and tree species composition on leaf miner abundance and species richness.................................. 64 4.1 Introduction.......................................................................................................... 64 4.2 Methods................................................................................................................ 66 4.2.1 Data analysis..............................................................................................
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