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Download (8MB) https://theses.gla.ac.uk/ Theses Digitisation: https://www.gla.ac.uk/myglasgow/research/enlighten/theses/digitisation/ This is a digitised version of the original print thesis. Copyright and moral rights for this work are retained by the author A copy can be downloaded for personal non-commercial research or study, without prior permission or charge This work cannot be reproduced or quoted extensively from without first obtaining permission in writing from the author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Enlighten: Theses https://theses.gla.ac.uk/ [email protected] Studies on the Biology of Tipula paludosa Meigen (Diptera: Tipulidae) with special reference to Mortality Factors NURI MILAD BARBASH, M.A. (W. Michigan) A thesis submitted for the degree of Doctor of Philosophy in the Faculty of Science at the University of Glasgow Department of Zoology University of Glasgow January 1988 ProQuest Number: 10997921 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 10997921 Published by ProQuest LLC(2018). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 Declaration I hereby declare that this thesis represents, except where a note is made to the contrary, work carried out by myself. It has not been previously submitted for any degree. N.M. Barbash January 1988 Table of Contents SECTION PAGE Acknowledgements .. .. .. xix Summary .. .. .. xx 1. Introduction .. .. .. 1 1.1. The life cycle and economic importance of Tipula paludosa .. .. .. 1 1.2. The present work .. .. .. 3 2. General Review of Literature .. .. .. 4 2.1. Taxonomy and descriptions .. .. - 4 2.2. Bionomics .. .. .. 5 2.3. The effects of climate on the immature stages .. 6 2.3.1. Water relations .. .. .. 6 2.3.2. Effects of temperature .. .. .. 8 2.3.3. Natural mortality factors .. .. 9 3. The Study Area .. .. .. 10 4. Quantitative Studies on Field Population of T.paludosa 13 4.1. Studies on adult flies .. .. .. 13 4.1.1. Introduction .. .. .. 13 4.1.2. Methods and Materials .. .. .. 13 4.1.3. Emergence cages .. .. .. 14 4.1.4. Results from observations with emergence cages 15 4.2. Sex ratios in T.paludosa .. .. .. 30 4.2.1. Introduction .. .. .. 30 4.2.2. Materials and Methods .. .. .. 31 4.2.3. Results and Discussion .. .. .. 32 iii 4.3. Studies on eggs .. .. .. 37 4.3.1. Materials and Methods .. .. .. 37 4.3.2. Results .. .. .. 42 5. Test of Technique of Studying Egg Oviposition .. 49 5.1. Ovipositional preference of T.paludosa .. 49 5.1.1. Introduction .. .. .. 49 5.1.2. Materials and Methods .. .. .. 49 5.1.3. Results .. .. .. 50 5.2. The effects of saturated solutions of magnesium sulphate on newly-laid eggs of T.paludosa .. 56 5.2.1. Introduction .. .. .. 56 5.2.2. Materials and Methods .. .. .. 57 5.2.3. Results .. .. .. 57 6 . Effects of Temperature on the Development and Hatching of Eggs .. .. .. 59 6.1. Introduction .. .. .. 59 6.2. Materials and Methods .. .. .. 59 6.3. Results .. .. .. 60 6.4. Hatching rates of eggs .. .. .. 63 7. Effects of Reduced Humidity on Eggs and First Instar Larvae of T.paludosa .. .. .. 69 7.1. Introduction .. .. .. 69 7.2. Materials and Methods .. .. .. 70 7.2.1. Observations on eggs .. .. .. 70 7.2.2. Observations on larvae .. .. .. 73 iv 7.3. Results .. .. .. 7 3 7.3.1. Eggs .. .. .. 73 7.3.2. Larvae .. .. .. 75 8 . Studies on Larvae .. .. .. 81 8.1. Extraction of larvae from the soil .. .. 81 8.2. Sampling for larvae in the field .. .. 8 6 8.2.1. Studies on population densities of larvae in the field .. .. .. 87 8.2.2. Results .. .. .. 87 8.3. Test of technique - The efficiency of the extracting equipment .. .. .. 89 8.4. Rearing T.paludosa in the laboratory .. 90 8.4.1. Introduction .. .. .. 90 8.4.2. Materials and Methods .. .. .. 90 8.4.3. Results .. .. .. 92 8.5. The effects of population density on survival of larvae of T.paludosa .. .. .. 92 8.5.1. Introduction .. .. .. 92 8.5.2. Materials and Methods .. .. .. 93 8 .5.2.1. Experiment 1 .. .. .. 93 8 .5.2.2. Experiment 2 .. .. .. 94 8.5.3. Results .. .. .. 94 8 .5.3.1. Experiment 1 .. .. .. 94 8 .5.3.2. Experiment 2 .. .. .. 95 8.5.4. Discussion .. .. .. 101 8 .6 . The effects of submergence and oxygen depletion on the third instar larvae of T.paludosa .. 101 v 8.6.1. Introduction .. .. .. 101 8.6.2. Materials and Methods .. .. .. 102 8 .6 .2.1. Survival in aerated water .. .. 102 8 .6 .2 .2 . Oxygen-deficient water .. .. 103 8.6.3. Results .. .. .. 103 8 .6 .3.1. Aerated water .. .. .. 103 8 .6 .3.2. Oxygen-deficient water .. .. 107 8.6.4. Discussion .. .. .. 112 8.7. Effects of cold on third instar larvae of T.paludosa 113 8.7.1. Introduction .. .. .. 113 8.7.2. The ability of T.paludosa third instar larvae to survive exposure to sub-zero temperatures 116 8 .7.2.1. Materials and Methods .. .. 116 8 .7.2.2. Results .. .. .. 117 8.7.3. The ability of the larvae to move from freezing conditions .. .. .. 117 8 .7.3.1. Materials and Methods .. .. 117 8 .7.3.2. Results .. .. .. 122 8 .8 . Studies on the growth of larvae .. .. 124 8.8.1. Studies on field populations .. .. 124 8 .8 .1.1. Materials and Methods .. .. 124 8 .8 .1.2. Results .. .. .. 124 8.8.2. The effects of temperature on the growth of larvae in the laboratory .. .. 131 8 .8 .2.1. Materials and Methods .. .. 131 8 .8 .2.2. Results .. .. .. 132 8.9. Proportion of larvae pupating when cultured at different temperatures .. .. .. 143 vi 8.10. The weights of mature larvae and pupae in relation to their sex and the fecundity of the females 144 8.10.1. Introduction .. .. .. 144 8.10.2. Materials and Methods .. .. .. 145 8.10.3. Results .. .. .. 145 9. Diseases, Parasites and Invertebrate Predators of Tipulidae .. .. .. 155 9.1. Introduction .. .. .. 155 9.2. Virus diseases ... .. .. 157 9.2.1. Previous work .. .. .. 157 9.2.2. Present observations .. .. .. 158 9.3. Bacterial diseases .. .. .. 163 9.4. Infections by fungi .. .. .. 163 9.4.1. Previous work .. .. .. 163 9.4.2. Present observations .. .. .. 164 9.5. Protozoal infections .. .. .. 164 9.5.1. Previous work .. .. .. 164 9.5.2. Present observations .. .. .. 165 9.6. Parasitic Nematoda .. .. .. 182 9.6.1. Previous work .. .. .. 182 9.6.2. Present observations .. .. .. 182 9.7. Insect parasitoids .. .. .. 188 9.7.1. Previous observations .. .. .. 188 9.7.1.1. Tachinidae .. .. .. 188 9.7.1.2. Phoridae .. .. .. 189 9.7.1.3. Muscidae .. .. •• 189 9.7.2. Present observations .. .. .. 198 vii 9.8. Insect predators .. .. .. 198 9.8.1. Previous work .. .. .. 198 9.8.2. Present observations .. .. .. 199 9.9. Parasites and predators of adult Tipulidae .. 199 9.9.1. Previous work .. .. .. 199 9.9.2. Present observations .. .. .. 199 9.10. Other abnormalities .. .. .. 200 9.10.1. Affecting larvae .. .. .. 200 9.10.2. Affecting adults .. .. .. 201 9.11. Discussion .. .. .. 201 10. Predation of Larvae by Birds .. .. .. 203 10.1. Introduction .. .. .. 203 10.2. Materials andMethods .. .. .. 204 10.2.1. The 1985-86 experiment .. .. 204 10.2.2. The 1986-87 experiment .. .. 207 10.3. Results .. .. .. 208 10.3.1. Results - 1985-86 .. .. .. 208 10.3.2. Results - 1986-87 .. .. .. 217 11. Effects of the Larvae of Tipula paludosa on the Grass of the Bird-Exclusion Experiments .. .. 224 11.1. Introduction .. .. .. 224 11.2. Yield of grass in bird-exclusion experiments .. 225 11.2.1. Materials and Methods .. .. .. 225 11.3. Results .. .. .. 226 11.4. Discussion .. .. .. 230 References .. .. • • 233 Appendices .. • • • • 249 viii LIST OF APPENDICES PAGE Appendix I. The numbers, accumulative numbers and accumulative percentages of males and females extracted daily from emergence traps in 1984, 1985 and 1986 249 Appendix II. Number of eggs collected weekly from the field in 3 years and the percentage hatched in the laboratory with their detailed plot counts in 1985 and 1986 253 Appendix III. Analysis of variance of oviposition preference of T.paludosa 260 Appendix IV. Detailed results of the eggs hatched in different cultures using different ages of eggs 265 Appendix V. Individual larvae collected from each soil core in each plot in 1984-85 with transformed data .. .. .. 272 Appendix VI. The effects of population density on survival of larvae (Experiment 1) 283 Appendix VII. The number of larvae surviving prolonged freezing in the laboratory 285 Appendix VIII. Temperature and rainfall records from the period October 1983 to October 1986 287 Appendix IX. A trial with TIV 292 Appendix X. Data of the two years experiment on predation of larvae by birds 294 IX LIST OF TABLES PAGE 1. Data from emergence cages summarised for 1984, 1985 & 1986 15 2. Estimates of mean population density of adults on Lawmuir field in 1984, 1985 & 1986 30 3. Numbers of adult T.paludosa observed and their sex ratios 32 4. Weekly totals of eggs recorded on Lawmuir plots 43 5. Numbers of eggs/m^ calculated after square root transformation 44 6 . The numbers of eggs found in the three oviposition preference experiments 53 7.
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