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Effects of Altitude and Canopy Cover on the Nest Size and Colony Size of the Red Wood Ants Formica Lugubris and Formica Paralugubris

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Effects of Altitude and Canopy Cover on the Nest Size and Colony Size of the Red Wood Ants Formica Lugubris and Formica Paralugubris Effects of altitude and canopy cover on the nest size and colony size of the red wood ants Formica lugubris and Formica paralugubris Yi-Huei Chen PhD University of York Department of Biology January 2015 1 Abstract Variations in life-history characteristics across geographic gradients may have implications for the impact of environmental change on animals. Linking one of the most important life-history characteristics and a geographic gradient, Bergmann’s rule describes body size increase with increasing latitude. Due to comparable thermal patterns between latitude and altitude, a similar process is expected to apply across altitude. For social insects, the colony could be biologically analogous to the body of a unitary organism. This study investigates the relationship between altitude and colony size in social insects. The model species used were wood ants Formica lugubris and F. paralugubris. These species have a flexible nesting strategy known as polydomy. I therefore considered both nest size and colony size. Initially, I developed an accurate mark-release-recapture method to estimate nest size, and found that mound volume can be a useful nest size index. A detailed case-study focused on canopy cover effects and showed that nests were larger in shadier areas. Informed by the results, I finally assessed the relationship between altitude, canopy cover, polydomy, nest size and colony size. The results reveal that colony size follows Bergmann’s rule along altitude when canopy cover is controlled for: microclimatic factors can be more significant than geographic factors in determining colony size. A systematic review in the Appendix shows that F. lugubris populations in different locations differ in mean nest size, but shows no evidence of a trade-off between nest size and multi-nest organisation. This thesis not only provides the first intra-specific evidence of Bergmann’s rule acting at the colony level across altitude, but also indicates the prominent role of microclimate on a key life-history characteristic. The work therefore sheds light on the evolution of an eco-geographic cline and the effects which climate change may have on the cline. 2 Table of Contents Abstract .............................................................................................................................. 2 Table of Contents ............................................................................................................. 3 List of Tables ..................................................................................................................... 5 List of Figures ................................................................................................................... 7 Acknowledgements ........................................................................................................... 9 Declaration ...................................................................................................................... 10 Chapter 1 – General Introduction ............................................................................... 11 Body Size ......................................................................................................... 11 Bergmann’s Rule ............................................................................................. 14 Microclimate ................................................................................................... 20 The Social Insect Colony ................................................................................ 21 Red Wood Ants and Polydomy ...................................................................... 24 Rationale for the Thesis and Aims ................................................................. 27 Chapter 2 – A comparison of mark-release-recapture methods for estimating colony size in the wood ant Formica lugubris ............................................................ 30 Abstract ............................................................................................................ 30 Introduction ..................................................................................................... 31 Materials and Methods.................................................................................... 37 Results .............................................................................................................. 41 Discussion ........................................................................................................ 45 Acknowledgements ......................................................................................... 49 Chapter 3 – Preliminary tests and power analyses for the relationship between altitude and ant colony size ........................................................................................... 50 Abstract ............................................................................................................ 50 Introduction ..................................................................................................... 51 Materials and Methods.................................................................................... 53 Results .............................................................................................................. 57 Discussion ........................................................................................................ 63 3 Chapter 4 – The relationship between canopy cover and colony size of the wood ant Formica lugubris - implications for the thermal effects on a keystone ant species ............................................................................................................................... 68 Abstract ............................................................................................................ 68 Introduction ..................................................................................................... 69 Materials and Methods.................................................................................... 72 Results .............................................................................................................. 76 Discussion ........................................................................................................ 84 Acknowledgements ......................................................................................... 92 Chapter 5 – Is Bergmann’s rule applicable to the relationship between altitude and ant colony size? Geographic gradient versus microclimate ............................. 93 Abstract ............................................................................................................ 93 Introduction ..................................................................................................... 94 Materials and Methods.................................................................................... 99 Results ............................................................................................................ 103 Discussion ...................................................................................................... 112 Acknowledgements ....................................................................................... 118 Appendices .................................................................................................... 118 Chapter 6 – General Discussion ................................................................................. 120 Summary of Chapters ................................................................................... 120 Limitations and Future Work ....................................................................... 123 Conclusion ..................................................................................................... 127 Appendix – Polydomy and Nest Size ......................................................................... 129 Introduction ................................................................................................... 129 Materials and Methods.................................................................................. 130 Results ............................................................................................................ 134 Discussion ...................................................................................................... 136 List of References ......................................................................................................... 140 4 List of Tables Table 1.1. Original definition and suggested redefinitions of Bergmann’s rule from the review of Watt et al. (2010) ........................................................................... 16 Table 1.2. Number of studies in insects with Bergmann’s, converse-Bergmann’s, and no clines, compared inter- and intra-specifically within the type of range examined (latitude or altitude) (adapted from Shelomi, 2012) .......................... 17 Table 1.3. Summary of previous research on Bergmann’s rule in social insects, broken down into inter-specific or intra-specific Bergmann’s rule between latitude (or altitude) and body size (or colony size) of social insects ............... 28 Table 2.1. Summary of the results in studies which compared the estimates by non- destructive methods with real parameters of ant colony size ............................ 34 Table 2.2. Results of the linear regression for the relationships between the estimated colony size and the actual colony size from five methods
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