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Robin Curtis The importance of resources in determining butterfly population abundance at multiple scales Robin Curtis This dissertation is submitted for a Research Degree at University College London 1 Declaration I, Robin Curtis confirm that the work presented in this thesis is my own. Where information has been derived from other sources, I confirm that this has been indicated in the thesis 2 Acknowledgements I would like to thank my supervisors, Chris Carbone, James Mallet and in particular Nick Isaac for useful advice and assistance throughout the duration of this research degree. I would also like to thank Roger Dennis and David Roy for constructive comments on draft manuscripts. For research undertaken on the Glanville fritillary on the Isle of Wight, I would like to thank members of the Isle of Wight Natural History and Archaeological Society, and in particular Andy Butler, for sharing their knowledge and enthusiasm for this amazing butterfly. I would also like to thank David Simcox for ideas and advice for research. Finally, I would like to thank Wightlink Ferries for providing free transport to and from one of the best Islands in the U.K. For the chalk grassland multisite chapter I would like to thank Butterfly Conservation staff for advice and information on sites, and especially Tom Brereton. I would also like to thank the many transect walkers, and in particular Gail Jeffcoate, for assisting me in recording the various transect routes across some of Britain’s finest nature reserves. I would like to thank Mark Botham and John Redhead at CEH Wallingford for assisting with various GIS issues. I would also like to thank Gary Powney for opening the office windows and letting in some fresh air. Finally, I would particularly like to thank Jeremy Thomas for providing advice and inspiration for butterfly research since 1992, when I initially worked as a research assistant on various butterfly projects at Furzebrook Research Station, Dorset. It was under his guidance that I started research on the Glanville fritillary butterfly in 1996, and his encouragement motivated me to keep returning to collect data on the Isle of Wight, even after the end of my employment in 1997. His knowledge and support for my research has proved invaluable, both before I registered for a PhD in 2007 and during my doctoral research. 3 Abstract Long term monitoring has revealed that butterfly populations vary dramatically between years, among sites and across species. This PhD investigates the importance of biotic and abiotic resources in determining butterfly abundance at multiple scales, using long term data from intensive studies on the Glanville fritillary and from the extensive UK Butterfly Monitoring Scheme. The most important resource at local scales is the ‘suitability’ of the larval hostplant: only plants within a certain temperature range are likely to be utilised for oviposition. Laying eggs on warmer plants maximises the probability of successful larval development within restricted seasonal temperature constraints. Analysis of Glanville fritillary population time-series reveals that populations are more abundant in those habitat types providing suitable microclimates for larval development, and in years following warm spring temperatures (i.e. during oviposition). These factors were also important when comparing abundance across sites, as were patch area and isolation. This landscape-scale analysis also revealed a significant negative interaction between habitat quality and temperature, suggesting that microsites for oviposition are less restricted in warm than cool years. Across regional scales (southeast England), the availability of both host and nectar plants are important in determining butterfly abundance, both across and within species. However, hostplant availability is a stronger predictor of abundance than nectar, and is most pronounced on sites with a northerly aspect. Trait analysis revealed that species with restricted diets, low mobility and habitat specialism are most sensitive to variation in foodplant availability. Taken together, the results suggest that many UK butterfly species are limited at small scales by the availability of hostplants in warm microclimates, but at large scales by dispersal limitation and climatic tolerance. Site management can therefore increase the abundance of key species of conservation concern by creating thermally diverse habitats and maximising the biomass of their hostplants. 4 Table of Contents Declaration .................................................................................................................................... 2 Acknowledgements ....................................................................................................................... 3 Abstract ......................................................................................................................................... 4 Table of Contents .......................................................................................................................... 5 List of Tables ................................................................................................................................. 7 List of Figures ................................................................................................................................ 8 Introduction ................................................................................................................................ 10 Insects and butterflies as study organisms ............................................................................. 12 The advent of landscape ecology ............................................................................................ 19 Thesis outline .......................................................................................................................... 25 Chapter 1) The Effects of Temperature, Microclimate and Habitat Quality on Abundance of the Glanville fritillary on the Isle of Wight. ....................................................................................... 28 Abstract ................................................................................................................................... 28 Introduction ............................................................................................................................ 29 Methods .................................................................................................................................. 35 Statistical Analysis ................................................................................................................... 39 Results ..................................................................................................................................... 41 Discussion................................................................................................................................ 48 Recommendations for Site Management ............................................................................... 54 Conclusions ............................................................................................................................. 56 Chapter 2) The Effects of Area, Isolation, Habitat Quality and Temperature on the Abundance of the Glanville fritillary along the South Coast of the Isle of Wight. ......................................... 57 Abstract ................................................................................................................................... 57 Introduction ............................................................................................................................ 58 Methods .................................................................................................................................. 62 Statistical Analysis ................................................................................................................... 66 Results ..................................................................................................................................... 68 Discussion................................................................................................................................ 71 Recommendations for Site Management ............................................................................... 76 Conclusions ............................................................................................................................. 79 5 Chapter 3) Defining the Principle Determinants of Butterfly Abundance on Chalk Grassland across Southeast England. .......................................................................................................... 80 Abstract ................................................................................................................................... 80 Introduction ............................................................................................................................ 81 Methods .................................................................................................................................. 87 Statistical Analysis ................................................................................................................... 93 Results ..................................................................................................................................... 94 Discussion.............................................................................................................................
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