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RADAR Research Archive and Digital Asset Repository Geographical and temporal variation of biochemical and colour-pattern polymorphisms in the European moth, Noctua pronuba (L.) Rob Hammond (1994) https://radar.brookes.ac.uk/radar/items/12feb164-d4fa-49d0-b7f9-9d84aa604634/1/ Note if anything has been removed from thesis: Copyright © and Moral Rights for this thesis are retained by the author and/or other copyright owners. A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the copyright holder(s). The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the copyright holders. When referring to this work, the full bibliographic details must be given as follows: Hammond, R (1994), Geographical and temporal variation of biochemical and colour-pattern polymorphisms in the European moth, Noctua pronuba (L.), PhD, Oxford Brookes University WWW.BROOKES.AC.UK/GO/RADAR Geographical and temporal variation of biochemical and colour-pattern polymorphisms in the European moth, Noctua pronuba (L.). Rob Hammond A thesis submitted in partial fulfilment of the requirements of Oxford Brookes University for the degree of Doctor of Philosophy December 1994 Contents: Abstract: .......................................................................................... IV Acknowledgements: ...........................................................................v Chapter 1: Polymorphism: ..........................................•...............•...... 1 1.1: Polymorphism: .............................................................................. 1 1.2: Selective mechanisms that maintain polymorph isms: .................. 3 1.2.1: Heterozygote advantage: .................................................. 3 1.2.2: Heterogeneous environments: .......................................... 4 1.2.3: Frequency-dependent selection: ....................................... 7 1.2.4: Frequency-dependent mating: .......................................... 9 1.3: Random processes: ..................................................................... 10 1.4: Populations: .................................................................................. 12 Chapter 2: Noctua pronuba: ..............................................................14 2.1: Species range: ............................................................................. 14 2.2: Life cycle: ..................................................................................... 14 2.3: Phenology: ................................................................................... 15 2.4: Evidence for long distance dispersal: ........................................... 16 2.5: Polymorphism for forewing colour and pattern: ............................ 17 2.5.1: Description of the phenotypes: ......................................... 20 2.5.2: Genetic models: ................................................................ 22 2.5.3: Previous studies of the colour and pattern polymorphism: ............................................................................. 23 2.6: Aims: ............................................................................................ 24 Chapter 3: Background resting behaviour: ...................................... 25 3.1: Introduction ................................................................................... 25 3.2: Aims: ............................................................................................ 28 3.3: Materials and methods ................................................................. 28 3.4: Results ......................................................................................... 30 3.5: Discussion: ................................................................................... 36 Chapter 4: Variation In morph frequencies between males and females at the same site: ........................................41 4. 1.· Intro. d uct' Ion .................................................................................. .. 41 4.2: Aims: ............................................................................................ 42 4.3: Methods: ........................................................................................ 42 4.4: Results: ........................................................................................ 43 4.4.1: Comparison of phenotype frequencies in males and females ....................................................................................... 43 4.4.2: Variation between sexes over the flight season: ............... 46 4.5: Discussion: ................................................................................... 51 Chapter 5: Temporal variation In phenotype frequencies: ............. 55 5.1: Introduction: .................................................................................. 55 5.2: Aims: ............................................................................................ 56 5.3: Methods ........................................................................................ 56 5.4: Results: ........................................................................................ 57 5.4.1: Variation throughout the flight season: ............................. 57 5.4.2: Variation over a twenty five year period: ........................... 61 5.5: Discussion: ................................................................................... 74 Chapter 6: Geographic variation in the forewing polymorph ism: ................................................................ 78 6.1: Introduction: .................................................................................. 78 6.2: Aims: ............................................................................................ 79 6.3: Materials and Methods: ................................................................ 80 6.4: Results: ........................................................................................ 82 6.4.1: Variation in males: ............................................................ 91 6.4.2: Variation in females: ......................................................... 94 6.4.3: Variation in male phenotype frequency with longitude and latitude: ................................................................. 96 6.5: Discussion: ................................................................................... 100 6.5.1: Comparisons with Cook & Sarsam (1981} ........................ 100 6.5.2: Comparisons with Soli & Andersen (1990): ...................... 107 6.5.3: Comparisons with Poitout and Sues (1976): ..................... 108 6.5.4: Kentville, Nova Scotia - a recently established population: .................................................................................. 108 6.5.5: General discussion: .......................................................... 109 Chapter 7: Geographic variation in allozyme frequencies: .....•....•• 113 7.1: Introduction: .................................................................................. 113 7.2: Aims: ............................................................................................ 116 7.3: Materials and methods: ................................................................ 116 7.3.1: Gel Preparation: ................................................................ 116 7.3.2: Extract Preparation: .......................................................... 117 7.3.3: Loading of the starch gels: ................................................ 118 7.3.4: Running the gels: .............................................................. 119 7.3.5: Slicing the gels: ................................................................. 120 7.3.6: Staining the gel slices: ...................................................... 120 7.3.7: Initial screening procedure: ............................................... 121 7.3.8: Enzymes routinely screened: ............................................ 121 7.3.9: Allozyme nomenclature: .................................................... 121 7.3.10: Zymogram descriptions and interpretation: ..................... 124 7.3.11: Gel fixation and scoring of zymograms: .......................... 126 7.4: Statistical methods for analysing allozyme data ........................... 126 7.4.1: Conformance to Hardy-Weinberg: .................................... 126 7.4.2: Multilocus associations: .................................................... 127 7.4.3: F Statistics ........................................................................ 128 7.4.4: Genetic distance: .................................. ·...... ·.. ·................. 131 7.5: Results: ........................................................................................ 132 7.4.1: Allele.frequencies: ............................................................. 133 7.4.2: Conformance to Hardy-Weinberg: .................................... 133 7.4.3: Direct-count heterozygosities: ........................................... 133 7.4.4: Associations between allozyme loci: ................................. 138 .. II 7.4.5: F statistics: ........................................................................ 141 7.4.6: Genetic distance measures: ............................................. 142 7.5: Discussion: ..................................................................................