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Butterfly Metapopulations in Dynamic Habitats

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Butterfly Metapopulations in Dynamic Habitats BUTTERFLY METAPOPULATIONS IN DYNAMIC HABITATS JENNY HODGSON Thesis submittedfo;'the qualification of: PhD l > , , ,;.' t University of;Y~rk , ~---; Department of Biology September, 2007 Abstract Many species require habitats that are naturally patchily-distributed and ephemeral, but human activities fundamentally alter the rate and scale of habitat change. This thesis describes the development of a new metapopulation simulation model applicable to a broad range of species that depend on dynamic habitat. I apply the model to two case studies, both involving butterfly species that use early-successional habitats and that are UK Biodiversity Action Plan priority species. I describe two methods for parameterising the model for a particular metapopulation in a particular landscape. One method uses parallel disturbance and population presence data for a few consecutive years; the other derives population parameters and landscape parameters from separate data sources. In the first case study, I found that the BAP target for Heath Fritillary (Melitaea athalia) populations in the Blean Woods, Kent, could either be met by approximately doubling the coppicing effort, or by concentrating the existing effort into one of the larger woodland blocks. In the second case study, I found that the rate of heathland burning in the South Stack area of Anglesey was not enough by itself to sustain the metapopulation of Silver-studded Blues (Pleb~'us a7,us). However, the metapopulation is probably saved from extinction by the existence of permanently-suitable habitat close to the coast. I have also elucidated an important phenomenon in metapopulations with dynamic habitat: the relationship between patch occupancy and patch connectivity can be obscured by the temporal changes in habitat. This has important implications for the debate about whether many real populations actually fit the metapopulation paradigm because the existence of metapopulation dynamics is often determined by testing the connectivity-occupancy relationship. The simulation model, "MANAGE", is an important new tool for integrating landscape-scale information, and answering conservation questions, in a field which is relatively new and unexplored. 2 Contents Abstract .................................................................................................................................................... 2 Figures ...................................................................................................................................................... 6 Tables ....................................................................................................................................................... 8 Acknowledgements .............................................................................................................................. 10 Declaration ............................................................................................................................................ 12 1 General introduction ................................................................................................................... 13 1.1 Rationale .............................................................................................................................. 13 1.2 Metapopulations ................................................................................................................. 13 1.2 Habitat quality ..................................................................................................................... 17 1.3 Conserving habitats by managing succession ................................................................. 19 1.4 Study species ....................................................................................................................... 21 2 Technical introduction ................................................................................................................ 26 2.1 Review of dynamic landscape metapopulation models ................................................ 26 2.2 The dynamic landscape metapopulation model MANAGE ........................................ 33 2.2.1 Model formulation ......................................................................................................... 33 2.2.2 Population dynamics ..................................................................................................... 33 2.2.3 Habitat patch dynamics ............................................................ ·.... ·............................... 34 2.2.4 Output ............................................................................................................................. 37 2.2.5 On the differences between MANAGE and closely related models ..................... 38 3 Viability of the Heath Fritillary populations in the Blean Woods depends on the spatial distribution of coppicing..................................................................................................................... 40 3.1 Introduction ........................................................................................................................ 40 3.2 Methods ............................................................................................................................... 42 3.2.1 Data collection and digitising ....................................................................................... 42 3.2.2 Statistical analysis of colonisation and extinction ...................................................... 43 3.2.3 Simulations of management and metapopulations ................................................... 46 3.3 Results .................................................................................................................................. 49 3.3.1 Baseline simulations ....................................................................................................... 49 3.3.2 Worst case scenarios ...................................................................................................... 52 3.4 Discussion ............................................................................................................................ 54 3.5 Summary .............................................................................................................................. 56 4 Vegetation and microclimate variables affecting occurrence and abundance of Lasius niger on heathland.......................................................................................................................................... 57 3 4.1 Introduction ........................................................................................................................ 57 4.2 Methods part 1: Field study .............................................................................................. 59 4.2.1 Ant transect ..................................................................................................................... 59 4.2.2 Temperature logging ...................................................................................................... 61 4.2.3 Data analysis ................................................................................................................... 61 4.2.4 Ant-butterfly relationship ............................................................................................. 63 4.3 Results part 1: Field study ................................................................................................. 64 4.3.1 Ant transect and temperature ....................................................................................... 64 4.3.2 Ant-butterfly relationship ............................................................................................. 68 4.4 Methods part 2: Parameterisation .................................................................................... 70 4.5 Results part 2: Parameterisation ....................................................................................... 72 4.6 Discussion ............................................................................................................................ 74 4.6.1 Habitat and microclimate associations of Lanus niger ............................................... 74 4.6.2 The relationship between P. a'l,us and L niger ........................................................... 75 4.7 Summary .............................................................................................................................. 76 5 Responses of butterfly metapopulations to patch connectivity and average patch quality are masked by successional habitat dynamics ................................................................................... 78 5.1 Introduction ........................................................................................................................ 78 5.2 Methods ............................................................................................................................... 82 5.2.1 Study species and field surveys .................................................................................... 82 5.2.2 Analysis of population density ..................................................................................... 83 5.2.3 Fitting IFM parameters using SPOMSIM .................................................................. 84 5.2.4 Analysis of patch occupancy ........................................................................................ 86 5.2.5
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