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Podarcis Muralis): Population Dynamics, Interactions and Adaptations

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Podarcis Muralis): Population Dynamics, Interactions and Adaptations THE INVASION ECOLOGY OF COMMON WALL LIZARD (PODARCIS MURALIS): POPULATION DYNAMICS, INTERACTIONS AND ADAPTATIONS Robert James Williams Submitted in accordance with the requirements for the degree of Doctor of Philosophy (Ph.D.) The University of Leeds Faculty of Biological Sciences School of Biology August 2019 I The candidate confirms that the work submitted is his own, except where work which has formed part of jointly-authored publications has been included. The contribution of the candidate and the other authors to this work has been explicitly indicated below. The candidate confirms that appropriate credit has been given within the thesis where reference has been made to the work of others. The work in Chapter 6 of the thesis has appeared in publication as follows: Williams, R*, Dunn AM, Quinn C, Hassall C (2019) Stakeholder discourse and opinion towards a charismatic non-native lizard species: potential invasive problem or a welcome addition? People and Nature. 1: 152-166. Myself and C.H. conceived the ideas and designed methodology; I collected and analysed the data; C.Q., A.D. and C.H. provided guidance on the final Q set. I led the writing of the manuscript. All co-authors contributed critically to the drafts and gave final approval for publication. This copy has been supplied on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. The right of Robert Williams to be identified as Author of this work has been asserted by him in accordance with the Copyright, Designs and Patents Act 1988. © 2019 The University of Leeds and Robert James Williams II Acknowledgements Firstly I would like to express huge gratitude to all those who gave me the opportunity to undertake this Ph.D. Special thanks go to Dr Chris Hassall and Dr Alison Dunn, firstly for allowing me the freedom to undertake this particular project and to disappear off into the field, and secondly, for all the invaluable support, guidance, knowledge, advice, and encouragement they have imparted to me. Thanks to Dr Gavin Hanke and his family for their friendship and hospitality in hosting me on Vancouver Island. Thank you Ellie Field, Hannah Grice, Miguel Martinez, Joel Dixon, Lily Mendes da Costa, Jack Carney, and Elizabeth Cooper for their assistance, hard work, and good times in the field. Thanks to Thomas Moor, staff, and pupils of Bury C of E Primary School for letting me take over their shed; Ray and Sue Hamilton for their hospitality and local knowledge; Nick Dobbs, Martin Noble, Rob Pilley, Jim Foster, and Chris Davis for assistance navigating through the herpetologist community, and Liam Russel for access to sand lizards. Many thanks to Steve Langham, Dr Sozos Michailides and Dr Tobias Uller for invaluable information and their previous work on UK P. muralis; Dr Colin Donihue for advice on collecting bite force data, and special thanks to Dr Anthony Herrel for the kind use of his bite force equipment. I perhaps would not have taken on this challenge without the early encouragement received from Dr Angelo Pernetta, who assured me I was up to the task, thank you. Huge thanks to my parents and family for their unwavering support, understanding, and patience in allowing me to do what I have to do, and to the friends who haven’t seen me for 3 ½ years. Lastly, thanks to friends and colleagues in the Hassall and Dunn labs, and to NERC for funding this research and providing the opportunity for me to benefit from their Doctoral programme. III Abstract The potentially damaging outcomes of species introductions to areas outside of their natural range are well known, and invasive non-native species are regarded as one of the most significant threats to biodiversity worldwide. Social perceptions of non-native species are open to subjective influence, and in a rapidly changing world the implications of species introductions are becoming less defined. Understanding the ecology of invasions and the human perceptions of them, is therefore fundamentally important for managing all stages of species introductions, relying on a multidisciplinary approach to understanding the invasion process. Invasions can also provide opportunities to study adaptive responses of organisms to novel or changing environments, which in turn can provide insight into mechanistic workings of the invasion process and range expansion. In this study I investigate the ecology of introduced populations of the Common Wall lizard (Podarcis muralis). Using a multidisciplinary approach including population ecology, functional ecology, predictive modelling, social science, and behavioural experiments, I focus on the species’ invasion potential, adaptive responses, and implications for ecological impacts on native lizards in the UK and on Vancouver Island, British Columbia. I found that morphological differences between populations were associated with ancestral origins and infer a greater invasion potential for animals of Italian origin over French. Comparison of physical and performance traits, and a broad dietary niche of P. muralis, indicate considerable overlap between P. muralis and native lizards, suggesting high potential for competitive interaction. However, the varied behavioural responses observed towards scent cues in P. muralis and native lizards suggests an interplay between naivety and threat sensitivity may influence interspecific interaction. Models of predicted range expansion suggest P. muralis populations are likely to remain localised, but that potential for secondary translocation is likely to increase with increasing population size, particularly in urban habitats. I also found clear indication that charismatic non-native species such as P. muralis, may have use as model species with which to raise awareness and minimise the subjectivity shaping perceptions of invasive species in general. IV Table of Contents Acknowledgements.................................................................................................................... II Abstract ..................................................................................................................................... III List of figures ............................................................................................................................. IX List of Tables ........................................................................................................................... XIII Abbreviations .......................................................................................................................... XV Chapter 1: Alien reptile and amphibian introductions in Europe: a review .............................. 1 1.1 Abstract ............................................................................................................................ 2 1.2 Introduction ..................................................................................................................... 3 1.2.1 Alien species in a changing world ............................................................................. 3 1.2.2 A European perspective ............................................................................................ 4 1.2.3 Historical pattern of amphibian and reptile introductions in Europe ...................... 5 1.2.4 Patterns of establishment and invasion ................................................................... 7 1.2.5 Pathways to introductions ........................................................................................ 9 1.2.6 Biogeography ............................................................................................................ 9 1.2.7 Climate matching and species traits ....................................................................... 10 1.2.8 Propagule pressure ................................................................................................. 11 1.2.9 The role of native biodiversity ................................................................................ 12 1.2.10 Impacts in Europe ................................................................................................. 13 1.2.11 Disease .................................................................................................................. 14 1.2.12 Competition and predation .................................................................................. 14 1.2.13 Genetic pollution .................................................................................................. 15 1.3 Case study: Alien lizards in the UK ................................................................................. 15 1.3.1 Podarcis muralis ...................................................................................................... 16 1.3.2 Lacerta bilineata ..................................................................................................... 17 1.3.3 Direct impacts of introduction ................................................................................ 17 V 1.3.4 Range expansion ..................................................................................................... 18 1.4 Conclusions, future directions ....................................................................................... 20 1.5 Thesis outline ................................................................................................................. 21 Chapter 2: Population and individual fitness as indicators of invasion potential in multiple
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