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The Termite by Ogden Nash Biological studies on two European termite species: establishment risk in the UK Laetitia Virginie Laine B.Sc. M.Sc. D.I.C. A thesis submitted for the degree of Doctor of Philosophy of the University of London November 2002 Department of Biological Sciences, Imperial College, Silwood Park, Ascot, SL5 7PY, Berkshire Abstract The discovery of an accidental introduction of termites into Devon in 1994 generated great interest as termites were previously thought to be unable to establish in the UK due to unfavourable climatic conditions. Information about the species present in Devon, Reticulitermes grassei, was found to be lacking and the present study was undertaken to determine the importance of various abiotic and biotic factors in establishment of this species. The factors included in the study were the minimum termite number for establishment, the consumption of wood and its effect on survival and temperature and soil type. A review of the literature was also conducted, detailing the problems with the taxonomy of this termite genus, their present distribution pattern and the life cycle of Reticulitermes species. Two populations of both R. grassei and R. santonensis were studied. The effect of the minimum termite number was found to be significant in both laboratory and field conditions. However, survival decreased in the laboratory and increased in the field with increased number of termites. Consumption experiments were performed using blocks of Scots pine, beech and oak. In most cases termite populations were found to consume and survive best on oak. Consumption was also tested on live seedlings but these results were inconclusive. Survival was observed to increase with increased temperature. Soil type experiments provided very mixed results for termite survival and these did not seem to be linked to soil moisture content. The trigger for alate production was not determined but fluctuating temperature seemed to have a positive effect on fecundity. The two most important factors for termite establishment appear to be the actual termite number and temperature. The final chapter looks at climate change and the various factors involved in termite establishment in order to put the results of this study into context. 2 Acknowledgements There are many people that have helped me during this PhD project. I would firstly like to thank my supervisor Denis Wright for all his help and advice, The Forestry Commission who funded this project and Robert Verkerk who initiated the project. Simon Leather, Hugh Evans and Geoff LePatourel gave me much appreciated guidance during this research project. Thanks to Janet Phipps, for always knowing Denis' whereabouts (or not!). I am also indebted to Mick Crawley for his help (and patience) as I waded through the various problems I encountered with the interminable 'S TAT S'. I would like to thank Dave Yeoman (Rothamstead) for providing the soil samples for Chapter 7 and Ann Kelly and Richard Hartley (University of Plymouth) for doing the particle analysis of the soils. I would like to thank the various members of the CTBA who allowed me to use the laboratory in Bordeaux and the field site on L'ile d' Oldron. More specifically, Ivan Paulmier, especially for helping me to lug half a tonne of soil to Bordeaux and back, and Sandra Girardi for keeping an eye on my various experiments. I would like to thank Mr. Chaigneau for allowing me the use of the 'Office National des Forets' facilities and Paul Beasley, for producing various pieces of equipment, especially the field equipment (Chapter 4). Thanks also to Andy Wilby for proof reading part of the thesis. I am also grateful to Florent Vieau for the interesting discussions about termite life cycles and Brian Forschler for my week in Athens (Georgia) and for initiating me to 'house inspections'. I would like to thank all the people that made Silwood a little sunnier (even on a wet and windy winter's morning). Ek, Anna, Dan and Lou (the inseparable pair of pranksters!)... as well as Paul B., Kate, Rob B., Aunty Anne, Pauline and Christine. I would like to thank my family for all their support and love. Finally I would like to thank Fabien, my husband to be, without whom I would probably never had got through it. I thank him for all his support, encouragement, patience (the halo is in the post) and love. 3 And last but not least... thank you to all those termites that gave their lives for my research. This is for you... The Termite by Ogden Nash Some primal termite knocked on wood And tasted it, and found it good! And that is why your Cousin May Fell through the parlor floor today. 4 Contents ABSTRACT 2 ACKNOWLEDGEMENTS 3 CONTENTS 5 LIST OF FIGURES 8 LIST OF TABLES 13 CHAPTER 1 14 INTRODUCTION 1.1 Pest Status 14 1.2 Control tactics 15 1.2.1 Preventative 15 1.2.2 Remedial treatment 16 1.2.3 Conclusion 17 1.3 Project background 17 1.4 Aims and Objectives 18 CHAPTER 2 19 LITERATURE REVIEW 2.1 Introduction 19 2.2 Species studied 19 2.3 Taxonomy 20 2.4 Distribution 23 2.4.1 Genus 23 2.4.2 Reticulitermes grassei 25 2.4.3 Reticulitermes santonensis 26 2.4.4 Reticulitermes flavipes 26 2.5 Movement of termite species in France 27 2.6 Differences between the species studied 29 2.7 The life cycle of Reticulitermes spp. (Isoptera, Rhinotermitidae): what do we know? 32 2.7.1 Abstract 32 2.7.2 Introduction 33 2.7.3 Life history 34 2.7.4 Comparisons of life cycles 48 2.7.5 Conclusion 56 CHAPTER 3 58 GENERAL MATERIALS AND METHODS 3.1 Termite species and strains 58 3.2 Insect culture 58 3.3 Experiments using vermiculite 58 3.1 Wood block quality 59 3.4 Data analysis 59 5 CHAPTER 4 60 MINIMUM NUMBER FOR SUCCESSFUL TERMITE ESTABLISHMENT 4.1 Background 60 4.2 Materials and Methods 61 4.2.1 Survival of termites on Scots pine 61 4.2.2 Survival of termites in the field 62 4.2.3 Survival of termites in different substrate types 65 4.2.4 Survival of termites in different substrate volumes 66 4.3 Results 66 4.3.1 Survival of termites on Scots pine 66 4.3.2 Survival of termites in the field 69 4.3.3 Survival of termites in different substrate types 70 4.3.4 Survival of termites in different substrate volumes 71 4.4 Discussion 72 4.4.1 Survival of termites on Scots pine 72 4.4.2 Survival of termites in the field 74 4.4.3 Survival of termites in different substrate types 74 4.4.4 Survival of termites in different substrate volumes 75 4.5 Conclusion 75 CHAPTER 5 77 STUDIES ON TERMITE CONSUMPTION 5.1 Background 77 5.1.1 Consumption 78 5.2 Materials and Methods 80 5.2.1 Consumption on different wood species - no choice 80 5.2.2 Consumption on different wood species - choice 81 5.2.3 Consumption and survival on Scots pine over time 81 5.2.4 Survival on Corsican pine seedlings 81 5.2.5 Survival on Scots pine seedlings and in combination with wood blocks 82 5.2.6 Survival on Scots pine and cherry seedlings 83 5.3 Results 83 5.3.1 Consumption on different wood species - no choice 83 5.3.2 Consumption on different wood species - choice 85 5.3.3 Consumption and survival on Scots pine over time 87 5.3.4 Survival on Corsican pine seedling 89 5.3.5 Survival on Scots pine seedlings and in combination with wood blocks 90 5.3.6 Survival on Scots pine and cherry seedlings 92 5.4 Discussion 93 5.4.1 Consumption on different wood species - no choice 93 5.4.2 Consumption on different wood species - choice 94 5.4.3 Consumption and survival on Scots pine over time 94 5.4.4 Survival on Corsican pine seedlings 95 5.4.5 Survival on Scots pine seedlings and in combination with wood blocks 95 5.4.6 Survival on Scots pine and cherry seedlings 95 5.4.7 Overview 96 5.5 Conclusion 97 CHAPTER 6 99 REPRODUCTIVE STRATEGIES 6.1 Introduction 99 6.2 Materials and Methods 100 6.2.1 Observation Apparatus 100 6.2.2 Preliminary observations on R. grassei UK and R. santonensis BRE 101 6 6.2.3 Observational study on two populations of R. grassei and R. santonensis.101 6.2.4 Effect of cold treatment on reproductive strategy 102 6.3 Results 103 6.3.1 Preliminary observations on R. grassei UK and R. santonensis BRE 103 6.3.2 Observational study on two populations of R. grassei and R. santonensis104 6.3.3 Effect of cold treatment on reproductive strategy 109 6.4 Discussion 116 6.4.1 Preliminary observations on R. grassei UK and R. santonensis BRE 116 6.4.2 Observational study on two populations of R. grassei and R. santonensis 116 6.4.3 Effect of cold treatment on reproductive strategy 117 6.5 Conclusion 118 CHAPTER 7 120 TEMPERATURE AND SOIL TYPE AND THEIR EFFECT ON TERMITE SURVIVAL 7.1 Background 120 7.2 Materials and Methods 122 7.2.1 Survival of termites on different soil types 122 7.2.2 Survival of termites at different temperatures 124 7.3 Results 125 7.3.1 Survival of termites on different soil types 125 7.3.2 Survival of termites at different temperatures 126 7.4 Discussion 127 7.4.1 Survival of termites on different soil types 127 7.4.2 Survival of termites at different temperatures 128 7.5 Conclusion 129 CHAPTER 8 131 SUMMARY AND GENERAL DISCUSSION 8.1 Summary 131 8.2 Factors affecting termite establishment 132 8.3 Climate and termite distribution 134 8.4 Climate change 137 8.5 Conclusion 138 GLOSSARY 139 REFERENCES 140 APPENDIX 159 7 List of Figures CHAPTER 2 LITERATURE REVIEW FIGURE 2.1 WORLD-WIDE DISTRIBUTION OF RETICULITERMES SPECIES (AFTER PEARCE (1997)).
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