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Formica Lugubris The effects of forest cover change and polydomous colony organisation on the wood ant Formica lugubris Duncan Procter PhD University of York Biology March 2016 Abstract Anthropogenic land use changes, such as deforestation, generally have negative effects on ecosystems. However, in Europe recently, historic trends in deforestation are being reversed due to increases in planted forests, and it is becoming much rarer to replace native forests with plantations. British forests have undergone centuries of degradation and fragmentation, and increases in forest cover due to plantations represent a potential positive for forest specialist species struggling in isolated fragments. In this thesis, I assess forest cover change and the demographic and genetic health of populations of the wood ant Formica lugubris, a forest specialist, in the North York Moors National Park, UK. I show that, contrary to expectations, non-native conifer plantations have had incredibly beneficial effects on this forest specialist species. Populations of F. lugubris have expanded from historically isolated fragments, and show no evidence of this expansion ceasing. Furthermore expanded populations are genetically diverse in both nuclear and mitochondrial DNA, and show evidence of commercial forests connecting previously isolated population fragments. There is strong divergence within mitochondrial DNA across the landscape in F. lugubris, which suggests either a cryptic species within the study population, or an historic hybridisation event. Formica lugubris exhibits polydomous colony organisation, whereby multiple spatially separate nests display social and cooperative connections, and are therefore one colony. I show that socially connected nests are socially and cooperatively distinct from their neighbouring colony, but show no equivalent genetic distinction. The findings within this thesis support growing evidence that non-native conifer plantations can have positive effects on forest biodiversity, and that some wood ant populations within the UK are healthy and under no threat of extinction. Furthermore polydomous colonies are cooperative but not genetic units, and division of colonies in this species may be ecologically, rather than genetically determined. 2 Contents Abstract .............................................................................................................................. 2 List of Figures ...................................................................................................................... 5 List of Tables ....................................................................................................................... 7 Acknowledgements ............................................................................................................. 8 Author Declaration .............................................................................................................. 9 Chapter 1: Introduction ..................................................................................................... 10 1.1 General Introduction ................................................................................................... 10 1.2 Habitat Fragmentation ................................................................................................ 11 1.3 Forests and forest cover change ................................................................................. 13 1.4 The study species ........................................................................................................ 15 1.5 Polydomous colony organisation ................................................................................ 18 1.6 The North York Moors National Park .......................................................................... 21 1.7 Thesis outline .............................................................................................................. 22 Chapter 1A. Genetic delineation of polydomous colony boundaries ................................... 24 1A.1 Theoretical basis ......................................................................................................... 24 1A.2 Limitations................................................................................................................... 25 1A.3 Methods ...................................................................................................................... 26 1A.4 Conclusion ................................................................................................................... 28 Chapter 2: Do non-native conifer plantations provide benefits for a native forest specialist, the wood ant Formica lugubris? ........................................................................................ 29 2.1 Abstract ....................................................................................................................... 29 2.2 Introduction ................................................................................................................ 30 2.3 Materials and Methods ............................................................................................... 32 2.4 Results ......................................................................................................................... 40 2.5. Discussion .................................................................................................................... 46 2.6. Conclusion ................................................................................................................... 49 Chapter 3: The importance of spatial scale in assessing habitat quality: A case study on wood ants .................................................................................................................................. 50 3.1 Abstract ....................................................................................................................... 50 3.2 Introduction ................................................................................................................ 50 3.3 Methods ...................................................................................................................... 53 3.4 Results ......................................................................................................................... 57 3.5 Discussion .................................................................................................................... 65 3 Chapter 4: Positive effects of non-native conifer plantations: evidence of connection of previously isolated populations and high genetic diversity ................................................. 70 4.1 Abstract ....................................................................................................................... 70 4.2 Introduction ................................................................................................................ 70 4.3 Methods ...................................................................................................................... 72 4.4 Results ......................................................................................................................... 75 4.5 Discussion .................................................................................................................... 80 Chapter 5: Greater mitochondrial variation within Formica lugubris across a landscape than between species within the F. rufa group .......................................................................... 84 5.1 Abstract ....................................................................................................................... 84 5.2 Introduction ................................................................................................................ 84 5.3 Methods ...................................................................................................................... 86 5.4 Results ......................................................................................................................... 88 5.5 Discussion .................................................................................................................... 90 5.6 Conclusion ................................................................................................................... 93 Chapter 6: Does cooperation mean kinship between spatially discrete ant nests? ............... 94 6.1 Abstract ....................................................................................................................... 94 6.2 Introduction ................................................................................................................ 94 6.3 Methods ..................................................................................................................... 97 6.4 Results ....................................................................................................................... 101 6.5 Discussion .................................................................................................................. 106 Chapter 7: Discussion ...................................................................................................... 111 7.1 Thesis overview ......................................................................................................... 111 7.2 The impacts of forest cover change .......................................................................... 111 7.3 Advice for forest managers ......................................................................................
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