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DOCTOR of PHILOSOPHY Bottom-Up Effects of Aboveground Biota, Invasive Species and Drought on Soil Animal Communities Ilardi

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DOCTOR of PHILOSOPHY Bottom-Up Effects of Aboveground Biota, Invasive Species and Drought on Soil Animal Communities Ilardi DOCTOR OF PHILOSOPHY Bottom-up effects of aboveground biota, invasive species and drought on soil animal communities Ilardi, Marco Award date: 2020 Awarding institution: Queen's University Belfast Link to publication Terms of use All those accessing thesis content in Queen’s University Belfast Research Portal are subject to the following terms and conditions of use • Copyright is subject to the Copyright, Designs and Patent Act 1988, or as modified by any successor legislation • Copyright and moral rights for thesis content are retained by the author and/or other copyright owners • A copy of a thesis may be downloaded for personal non-commercial research/study without the need for permission or charge • Distribution or reproduction of thesis content in any format is not permitted without the permission of the copyright holder • When citing this work, full bibliographic details should be supplied, including the author, title, awarding institution and date of thesis Take down policy A thesis can be removed from the Research Portal if there has been a breach of copyright, or a similarly robust reason. If you believe this document breaches copyright, or there is sufficient cause to take down, please contact us, citing details. Email: [email protected] Supplementary materials Where possible, we endeavour to provide supplementary materials to theses. This may include video, audio and other types of files. We endeavour to capture all content and upload as part of the Pure record for each thesis. Note, it may not be possible in all instances to convert analogue formats to usable digital formats for some supplementary materials. We exercise best efforts on our behalf and, in such instances, encourage the individual to consult the physical thesis for further information. Download date: 07. Oct. 2021 Bottom-up effects of aboveground biota, invasive species and drought on soil animal communities Marco Onofrio Ilardi MSc. (Hons.) Natural Sciences (University of Palermo) A thesis submitted to the School of Biological Sciences, Queen’s University Belfast for the degree of Doctor of Philosophy February 2019 “We know more about the movement of celestial bodies than about the soil underfoot” - Leonardo da Vinci Acknowledgements Apart from the Department of Employment and Learning (DEL), which provided the funding for this research, I would like to thank first of all my principal supervisor Dr Tancredi Caruso, without whom this doctoral work would not have been possible. It is quite a task to find the right words to properly express my infinite gratitude. His help and guidance, backed by his deep knowledge and great enthusiasm, but also his moral support and friendship have been invaluable throughout the years, and well over the expectations, especially in the times when the work load seemed too challenging and overwhelming, and all that I needed was some dose of encouragement. Something which he never failed to provide. Thanks also to my second and third supervisors, Prof. Mark Emmerson and Prof. Jaimie Dick, who gave me important suggestions and advice, especially at the start of my research. But in particular, I owe Jaimie for the favour of making his driveway and garden available for me to visit several times, to collect amphipod specimens which were very much needed for the experiments. Special thanks go to Dr Sheena Cotter for her help and expertise in the design of the carrion beetle experiment, for which she also provided some of the primary resources; to Prof. Mark Maraun and Dr Sarah Zieger for welcoming me at the University of Gӧttingen, where they also trained me and guided me to the discovery of the wonders of stable isotopes; to Prof. Richard Bardgett and Dr Ellen Fry for having given me the opportunity to contribute to their NERC funded project, which in turn contributed to the realization of this work; and to the personnel at AFBI (Belfast) for having so kindly made themselves available to provide demonstrations and training on nematodes’ extraction and identification. 3 I am very much grateful to my fellow PhD-ers Matthew and Flor, and also to Alana and Patrick, for their help during field sampling and lab work, but specially for the craic in and out of the lab. Finally, but not by order of importance, I wish to thank my family. My wife Ying, for her infinite patience and support, even when my PhD study routines were at times too demanding on the family life. My children Erin, Ethan and Eugene, as they are my joy and my energy boost, always capable of erasing any residues of stress or tiredness, maintaining my focus and motivation. To them I dedicate this thesis. 4 Contents Summary……………………………………………………………………………..8 List of figures……………………………………………………………………….10 List of tables………………………………………………………………………...15 CHAPTER 1: General introduction………………………………………………17 1.1 Research Background………………………………………...……..18 1.2 Origin of soil and soil ecosystem………………………………….…20 1.3 Ecosystem functions and services under threat……………………22 1.4 The soil food web………………………………………………….…25 1.5 Modern approaches…………………………………………………29 1.6 The conceptual structure of this thesis, research aims and hypotheses…………………………………….…………………………….30 CHAPTER 2: Scavenging beetles regulate and stabilise the response of soil animal communities to pulse input of nutrients…………………………………..34 2.1 Abstract……………………………………………………………….35 2.2 Introduction…………………………………………………………..36 2.3 Methods……………………………………………………………….40 2.3.1 Soil sampling and microcosm assemblage…………………….40 2.3.2 Microcosms sampling and samples analysis……...……………43 2.3.3 Statistical analyses…………………………………………….44 2.4 Results………………………………………………………………..45 2.4.1 Microarthropods abundances…………………………………45 5 2.4.2 Organic matter and pH………………………………………...48 2.4.3 Community structure…………………………………………..51 2.5 Discussion……………………………………………………………..55 2.6 Conclusions…………………………………………………………...57 2.7 Acknowledgements……………………………………….…………..58 CHAPTER 3: Response of native forest floor microarthropods to the invasive macroinvertebrate Arcitalitrus dorrieni (Crustacea; Amphipoda)……………..59 3.1 Abstract……………………………………………………………….60 3.2 Introduction…………………………………………………………..61 3.3 Methods……………………………………………………………….64 3.3.1 Experiment on community structure, environmental variables and trophic responses…………………………………..……………64 3.3.2 Secondary experiments: trials on leaf litter consumption rates (leaf litter quality and competition)………..…………...……………67 3.3.3 Stable isotopes analysis………………………………………..69 3.3.4 Statistical analyses…………………………………………….70 3.4 Results………………………………………………………………..71 3.4.1 Main experiment……………………………………………….71 3.4.2 Secondary experiments………………………………………...79 3.4.3 Summary of key results………………………………………..85 3.5 Discussion…………………………………………………………….86 3.6 Conclusions…………………………………………………………..89 3.7 Acknowledgements…………………………………………………..90 6 CHAPTER 4: Effects of extreme drought on the soil animal communities of manipulated grassland associations……………………………………………….91 4.1 Abstract……………………………………………………………….92 4.2 Introduction…………………………………………………………..93 4.3 Methods………………………………………………………………96 4.3.1 Study area and field experiment……………………………….96 4.3.2 Sampling, extraction and sorting………………………………99 4.3.3 Statistical analyses…………………………………………...101 4.4 Results……………………………………………………………….102 4.4.1 Microarthropods……………………………………………..102 4.4.2 Nematodes……………………………………………………108 4.5 Discussion……………………………………………………………113 4.6 Conclusions………………………………………………………….115 4.7 Acknowledgements………………………………………………….116 CHAPTER 5: General discussion………………………………………………..117 5.1 Theory and general hypotheses…...…….………………………….118 5.2 The role of fungi and bacteria in soil ………….……………………119 5.3 The experimental evidence …….…………...………………………121 5.4 Novel hypothesis and future directions…………….………………123 References…………………………………………………………………………125 Appendix A (supplemental material – Chapter 2)……………………...……….151 Appendix B (supplemental material – Chapter 3)……...……………………….156 Appendix C (supplemental material – Chapter 4)………………………….…...165 Appendix D (R scipts)……………………………………………………………..170 7 Summary Soils around the world, and the ecosystem functions and services they provide, are facing the common threat of a growing human population and the consequent changes currently observed from local to global scale. Soil food webs, which drive the ecological functions of soils, are being exposed to unprecedented levels of disturbance. This disturbance has the potential to disrupt the regulation of the interactions between components of the soil biota, and the role these components play in maintaining the stability of the belowground-aboveground linkages that support terrestrial ecosystems. This thesis investigates the ecological responses of the soil animal community to some sources of disturbance. The work focused on an array of perturbations that have their origins in the above-ground subsystem and that, directly or indirectly, lead to global environmental changes. These perturbations exert their primary impact on the resources entering the system, engendering a series of reverberating effects that travel from the bottom of the food web to the different levels of the decomposer community, from the basal consumers all the way up to the level of top predators (bottom-up perturbation). The perturbations investigated here were applied to the soil food web through experimental manipulations that ranged from microcosm to field experiments. It was possible to investigate disturbance in a controlled manner and so detect systematic effects on the soil animal communities, specifically soil microarthropods and nematodes,
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