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The Impact and Spread of the Invasive Garden Ant: an Alien Invasive Species in the UK Phillip Buckham-Bonnett Phd Thesis Univer

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The Impact and Spread of the Invasive Garden Ant: an Alien Invasive Species in the UK Phillip Buckham-Bonnett Phd Thesis Univer The impact and spread of the invasive garden ant: an alien invasive species in the UK Phillip Buckham-Bonnett PhD Thesis University of York Biology December 2019 Abstract Alongside climate change and habitat loss, invasive non-native species are a major threat to the natural world. Ants are amongst the most widespread and damaging invasive species. The invasive garden ant, Lasius neglectus, has only recently been detected in the United Kingdom and is the country’s first invasive ant species. This thesis aims to assess the impact and spread of this species in the UK. In this thesis I carry out a UK-wide risk assessment for the species and develop a protocol for experimentally assessing its potential impact on an economically important crop plant. I investigate behaviours that may contribute to its success as an invasive species. I evaluate the feasibility of commonly used pesticides for the control of Lasius neglectus, and conclude that granular products, while convenient for large- scale application, are not suitable for this species due to low palatability; this may hinder future control attempts. I carry out a large-scale survey to assess the distribution of this easily-overlooked species, and conclude that while it is not as widely distributed in botanic gardens as expected, the number of urban sites where it occurs is increasing rapidly. In conclusion Lasius neglectus is difficult to detect and hard to eradicate once established, so efforts should be made to reduce its spread from the areas in which it currently occurs. 2 Contents Abstract ............................................................................................................. 2 Contents ............................................................................................................ 3 Acknowledgements ........................................................................................... 4 Declaration ........................................................................................................ 6 Chapter 1: General Introduction ........................................................................ 8 Chapter 2: GB Non-native Species Rapid Risk Assessment (NRRA) for Lasius neglectus ......................................................................................................... 26 Chapter 3: Tactile communication in Lasius ants ............................................. 49 Chapter 4: Integrating foraging ecology informs approaches to invasive ant eradication ....................................................................................................... 71 Chapter 5: Assessing the UK distribution of Lasius neglectus ........................... 96 Chapter 6: General Discussion ....................................................................... 117 Appendix A (supplement 1 to Chapter 3) ....................................................... 129 Appendix B (supplement 2 to Chapter 3) ....................................................... 133 Appendix C (supplement to Chapter 4 - Methods) ......................................... 141 Appendix D (supplement to Chapter 4 - Results) ........................................... 147 Appendix E (supplement to Chapter 4 – Hydrogel Experiment) ..................... 184 Appendix F (supplement to Chapter 5) .......................................................... 187 Appendix G (supplement to Chapter 5) .......................................................... 195 Appendix H (Ant-plant interactions) .............................................................. 197 Appendix I (Lasius neglectus informational outputs) ..................................... 204 References ..................................................................................................... 210 3 Acknowledgements This thesis has been the supported by the contributions of a great many people for which I am very grateful. First and foremost, I would like to thank my supervisor, Elva Robinson. Without her help, input, inspiration, guidance, encouragement and even some video analysis, the project would not have been as rewarding and I would not have finished this thesis. I am also very grateful to Paul Lee as a supervisor, to Dan Franks as a TAP member and supervisor, and Peter Mayhew as a TAP member for all their contributions and support. I am very grateful to NERC and Hymettus for funding my PhD and to the Non-Native Species Secretariat for providing additional funds for research assistants. I also would like to thank the NNSS and the National Trust for providing and dispatching the traps and trapping materiel used in the botanic garden survey. I am grateful to Bayer, Bábolna Bio and DuPont for supplying the pesticides used in the experiments described in chapter 4. The placements with Hymettus have been one of the highlights of my PhD and I would like to thank all those involved, especially Paul Lee, Murdo Macdonald, Mike Edwards, Rowan Edwards, Scotty Dodd, Paddy Saunders, Brian Banks, Stephen Carroll, Nikki Gammans, Stewart Taylor, Athayde Tonhasca and Hayley Wiswell. My fieldwork would not have been possible without the support of the National Trust and the staff at Hidcote Manor, especially Chris Charman, Ann Davies and Glynn Jones. I am also grateful to John Righton for allowing me to run part of my experiment in his sheep field and for advice on how to stop the sheep eating it. My field work was also supported by the Cambridge University Botanic Gardens and I would like to thank all their staff especially Sally Petitt, Beverly Glover and Peter Sheld. My Cambridge field work was also very generously supported by Mary Vickers who provided me with accommodation. I am grateful to the staff at the sites I surveyed, for the permission to look for ants and also for the occasional cups of tea! I would like to thank past and present members of the Robinson Lab, for all their help, support, interesting discussions, red pens of justice and camaraderie. I would especially like to thank Duncan Proctor, Sam Ellis, Yi-Huei Chen, Eleanor Drinkwater, Dom Burns, Josie Monaghan, Meg Holgate, Valentin Lecheval and Joshua Sammy. I would also like to thank members of the Biology Department, YCCSA, GReBE, and the Theoretical Ecology and Whole Organism groups, especially Jon Pitchford, Jamie Woods, Leda Cai, Tom Hartley, Emma Rand, Melanie Smee, Stuart Nattrass, Ellie 4 Heyworth, Richard Law, Sarah Collings, Andy Suggitt and Gail Stride. I am grateful to Monica Bandeira and Sarah Christmas for their administrative support, Phil Roberts for permission to use his ant photographs and Mark Bentley for his fantastic workshop creations. I would like to thank Clive Boase for all his contributions and advice (even once answering the phone whilst at the top of a ladder!) on the pesticide and survey work and for first showing me around the Hidcote site. I am grateful to Alba Aguion for all her work on the first pesticide experiment and enthusiasm for learning survival analysis. I am grateful to Mike Fox for his ant identification assistance. I would like to thank Sebastian Mellor for discussions about and help with statistical analysis and coding. I am grateful to Tommy Czaczkes, Stefan Popp and Sophie Evison for being so willing to collaborate on tactile communication and all their work. I would like to thank Niall Moore, David Bullock and Simon Toomer for all their contributions and input through the Lasius neglectus task group. I am very grateful to all the volunteers who helped with the video analysis for chapter 4: Alice Bedois, Leda Cai, Laura Dunsby, Maria Guaita, Kate Hutchinson, Robyn Kinnersley, Alexander Kyte, Anna Middlemiss, Sarah Pickering, Charlotte Sauer and Lakesha Smith. I would like to thank all the staff at sites which took part in the survey for their help deploying and returning the traps for the botanic garden survey. I would like to thank Sarah Nicholson, Sarah Pickering and Tom Rudd, for processing the samples; and Sarah Pickering and Tom Rudd for their help with sample analysis for the botanic garden survey. I also would like to thank the staff in stores for tolerating all the dubious looking parcels labelled “ants” that I have received over the years. I would like to thank Helen Roy and Peter Mayhew for an enjoyable and stimulating viva and their very useful comments and suggestions for improving this thesis. Finally, I would like to thank my parents for all their support during my PhD. 5 Declaration I declare that this thesis is a presentation of original work and my contribution is detailed below. This material has not previously been presented for an award at this, or any other, university. All sources are acknowledged as references. Author contribution statements Chapter 1 (General introduction) Written by Phillip Buckham-Bonnett (PBB) with input from Elva Robinson (EJHR) and Paul Lee (PL) Chapter 2 (Risk Assessment) Written by PBB with input from EJHR and PL. This is published on the Non-Native Species Secretariat website at the following links: http://www.nonnativespecies.org/index.cfm?pageid=143 https://secure.fera.defra.gov.uk/nonnativespecies/downloadDocument.cfm?id=2065 Chapter 3 (Tactile Communication) This chapter represents a collaborative project to which PBB made significant contributions. More detail on the origin and contributions to this project is provided in the preface to Chapter 3, but in summary: experimental design involved Sophie Evison (SEFE), Tomer Czaczkes (TC), Stefan Popp (SP), EJHR and PBB. Data collection was performed by SP and PBB. Data analysis
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