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Housed in Silwood Park Library THE PHYTOPHAGOUS FAUNA OF GORSE (ULEX EUROPAEUS L.) AND HOST PLANT QUALITY RICHARD LAWRENCE HILL, M.Sc. A thesis submitted for the degree of Doctor of Philosophy of the University of London and for the Diploma of Imperial College HOUSED IN SILWOOD PARK LIBRARY Department of Pure & Applied Biology Imperial College Silwood Park Ascot Berkshire August 1982 2. ABSTRACT This study of the interactions between the fauna attacking gorse (Ulex europaeus L.) and their host-plant, was undertaken as part of a project aimed at the biological control of Ulex in New Zealand. In particular, the effects of seasonal changes in host-plant primary and secondary chemistry, plant structure and foliage morphology on the seasonal perform- ance of the phytophagous insect fauna were examined. Seasonal patterns of flowering and fruiting were described. Seasonal changes in foliage water content and toughness were summarised, and patterns of growth were described. These were discussed in relation to Lawton's (1978) living space concept and other current literature. Seasonal variation in the concentrations of 6 types of secondary compounds in the foliage were described. High concentrations of alkaloids may protect vegetative buds from herbivore attack in early spring, but otherwise foliage appeared to be unprotected. Energy content and soluble carbohydrate content of gorse foliage remained relatively high throughout the growing season. Foliage nitrogen content was high at bud-burst but declined within 6 weeks to a constant low level. The insect fauna associated with Ulex europaeus in Britain was described, compared and contrasted with the equivalent continental fauna and the faunas of related host-plants. Most species were rare, though the total number of phytophagous species was not low. Seasonal variation in abundance of the 7 commonest folivorous species was described in detail. All appeared within 3 weeks of bud-burst, no additional species appeared after the end of June, and almost all were fully developed and rare by the end of July. 3. Comparisons between the patterns of insect occurrence on gorse and seasonal variation of the plant parameters measured show that peak phyto- phagous insect feeding activity was associated with peak nutrient levels, and possibly minimum foliage alkaloid concentration. The impact of each of the measured factors on the phytophagous insect fauna was discussed in relation to current theory, and in particular the concept of "optimal defense theory". It was concluded that Ulex europaeus may be an example of a plant which has developed low nutritional quality as a defence % mechanism against herbivores. These findings were tested by two experiments. Host-plant character- istics were altered by the use of nitrogenous fertilisers in the field, and the impact of this on the phytophagous fauna was discussed. Neither the biology of the host-plant or the fauna were changed to a large extent by the fertiliser regimes used. Feeding experiments using LV larvae of Agonopterzx ulicetella showed that the nutritional quality of gorse foliage declined through summer. The relevance of such studies to the procedures used in the biological control of weeds is discussed. 4. TABLE OF CONTENTS Page ABSTRACT 2 LIST OF TABLES 8 LIST OF FIGURES 10 LIST OF APPENDICES 13 SECTION 1: INTRODUCTION 14 SECTION 2: SEASONAL CHANGES IN THE FORM AND STRUCTURE OF GORSE PLANTS 20 2.1 Introduction 20 2.2 Description of .Field Sites 21 2.3 Materials and Methods 22 2.3.1 Analysis of plant structure 22 2.3.2 Foliage water content 22 2.3.3 Foliage toughness 23 2.4 Results 23 2.4.1 Patterns of flowering and fruiting 23 2.4.2 Seasonal patterns of growth in mature Ulex spp ... 25 2.4.3 Seasonal changes in foliage water content 31 2.4.4 Seasonal variation of Ulex europaeus foliage toughness ... ... ... ... ... ... ... 34 2.5 Discussion 34 2.6 Summary ... 42 SECTION 3: THE PRIMARY AND SECONDARY CHEMISTRY OF GORSE FOLIAGE ... 44 3.1 Introduction ... 44 3.2 Sampling and Storage Methods 47 3.3 Analytical Methods 48 3.3.1 Analysis of nitrogen content 48 5. Page 3.3.2 Analysis of foliage calorific content 50 3.3.3 Analysis of soluble carbohydrate content 50 3.3.4 Analysis of alkaloids 51 3.3.5 Analysis of isoflavones 53 3.3.6 Analysis of silica content 54 3.3.7 Amino acid analysis 55 3.3.8 Analysis of foliar cyanogenesis 55 3.3.9 Analysis of foliar enzyme inhibitors ... ... ... 55 3.4 Results ... 57 3.4.1 Nitrogen content - ... 57 3.4.2 Foliage calorific content ... ... 63 3.4.3 Soluble carbohydrate content ... ... 63 3.4.4 Alkaloid content 63 3.4.5 Isoflavone content 70 3.4.6 Silica content 73 3.4.7 Unusual amino acids 73 3.4.8 Foliage cyanogenesis 75 3.4.9 Foliar enzyme inhibitor content ... 75 3.5 Discussion 77 3.6 Summary 81 SECTION 4: THE PHYTOPHAGOUS INSECT FAUNA OF GORSE 84 4.1 Introduction 84 4.2 Sampling Methods ... 88 4.3 The Fauna of Gorse Roots 91 4.4 The Fauna of Reproductive Structures 97 4.5 The Fauna Attacking Gorse Foliage 105 4.6 Discussion ... 116 6. Page 4.6.1 Differences in fauna of U. europaeus and U. minor . 116 4.6.2 Fauna 1 differences between YC and WGP 119 4.6.3 Seasonal patterns in the gorse fauna ... ... ... 121 4.6.4 Feeding generalism and specialism of the gorse fauna ... ... ... ... ... ... 125 4.6.5 Richness of the gorse fauna 127 4.7 Summary ... ... ... ... ... ... ... ... ... 130 SECTION 5: THE EFFECTS OF HOST-PLANT QUALITY ON THE INSECT FAUNA OR GORSE 131 5.1 Introduction ... 131 5.2 Methods 136 5.3 The Effect of Predators 137 5.4 Patterns of Foliage Use by Insects 140 5.5 The Effects of Plant-Borne Factors 143 5.5.1 Plant structure and living space 143 5.5.2 Plant chemistry 149 5.6 The Relative Importance of Plant-Borne Factors 153 5.7 Discussion ... 156 5.8 Summary ... 159 SECTION 6: THE EFFECTS OF FERTILISER ON GORSE AND ITS FAUNA ... 161 6.1 Introduction ... 161 6.2 Methods ... ... 162 6.3 Results 164 6.3.1 Effects on seasonal variation in N content 164 6.3.2 Effects on the growth of new shoots 167 6.3.3 Effect of fertilisation on the insect fauna ... 170 6.4 Discussion ... 180 7. Page 6.5 Summary ... ... ... ... ... ... ... 188 SECTION 7: THE FEEDING CHARACTERISTICS OF ONE GORSE SPECIES, AGONOPTERIX ULICETELLA 190 7.1 Introduction 190 7.2 Methods 193 7.3 Results 196 7.3.1 Experimental monitoring 196 7.3.2 Phytophage quality 198 7.3.3 Daily larval performance 201 7.3.4 Stadial feeding efficiencies 204 7.4 Discussion ... ... ... ... ... ... ... ••• 206 7.5 Summary 209 SECTION 8: CONCLUSIONS ... 211 2X6 ACKNOWLEDGEMENTS REFERENCES 217 APPENDICES 230 8. LIST OF TABLES Page 3.1 : Seasonal variation of gorse foliage alkaloid composition as shown by TLC 69 3.2 : Isoflavone content of Ulex europaeus and U. minor 72 3.3 : % silica content of new, dry, Ulex europaeus foliage 74 4.1 : Sources of information concerning the insect fauna of gorse in Britain ... ... ... ... ... ... ... 85 4.2 : Feeding sices of arthropod species attacking gorse in Europe . 92 4.3 : Differences in the abundance of insects on Ulex europaeus. and U. minor 98 4.4 : Differences in the abundance of insects on Ulex europaeus at Yateley Common and Windsor Great Park 99 4.5 : Infestation of reproductive structures of Ulex europaeus in spring and summer ... ... ... ... ... 101 4.6 : The relative importance of phytophagous insects collected from Ulex europaeus foliage at YC and WGP in two years ... 108 4.7 : Detritivorous and predaceous insect species encountered in gorse foliage ... ... 117 4.8 : Life histories of fauna associated with reproductives of Ulex europaeus and Ulex minor 120 4.9 : Seasonal patterns in gorse insects 122 4.10 : Feeding specialism and generalism in gorse insects 126 4.11 : Species richness of phytophagous insects on 5 host-plant ; species (Tribe: Genisteae) estimated from some standard references ... ... ... ... 129 5.1 : Correlation of faunal variables with 8 plant borne factors ... 147 5.2 : Autocorrelations between independent variables used in 9. Page regression analysis ... ... ... ... ... 152 5.3 : Assessment of the contribution of 8 independent variables to a multiple regression model 154 6.1 : A statistical comparison of the soluble and total nitrogen contents found in fertilised and unfertilised gorse foliage . 165 6.2 : Statistical comparison of gorse foliage total nitrogen content in fertilised and unfertilised plants 168 6.3 : Statistical comparison of median shoot growth (cm) on fertilised and unfertilised gorse 171 6.4 : Statistical comparison of differences in abundance between insect species on fertilised and unfertilised Ulex europaeus . 174 6.5 : Comparison of fertilisation effects on Holous Iccncctus and Ulex europaeus 185 7.1 : Incubation, storage, and rearing details for each treatment . 194 7.2 : Comparison of components in each treatment ... 197 7.3 : Measurements of parameters reflecting insect quality in each treatment ... ... ... ... ... ... 199 7.4 : Daily comparison of larval performance during LV in treat- ments 2-4 ... ... ... ... ... ... 202 7.5 : Measurement of ecological efficiencies for fifth instar in treatments 2-4 ... 205 10. LIST OF FIGURES Page 2.1 : (a) Apparatus for measuring foliage toughness and (b) Foliage nomenclature ... ... ... 24 2.2 : Patterns of flowering and fruiting in U. europaeus and U. minor 26 2.3 : Patterns of foliage growth in Ulex europaeus and Ulex minor . 29 2.4 : The relationship between shoot length and foliage biomass ... 30 2.5 : Seasonal variation of mean foliage water content in Ulex europaeus and Ulex minor 32 2.6 : Within shoot seasonal variation of foliage water content ... 33 2.7 : Seasonal variation of foliage toughness Ulex europaeus ..
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