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Seed Capsules.M 1800 Durham E-Theses Aspects of the ecology of coleophora alticolella zeller (lepidoptepa) with particular reference to altitude Randall, Martin G. M. How to cite: Randall, Martin G. M. (1980) Aspects of the ecology of coleophora alticolella zeller (lepidoptepa) with particular reference to altitude, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/7424/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk 2 ASPECTS OF THE ECOLOGY OF COLEOPHORA ALTICOLELLR ZELLER (LEPIDOPTERA) WITH PARTICULAR REFERENCE TO ALTITUDE MARTIN G.M. RANDALL B.Sc. (Dunelm) The copyright of this thesis rests with the author. No quotation from it should be published without his prior written consent and information derived from it should be acknowledged. .... being a thesis presented in candidature for the degree of Doctor of Philosophy in the University of Durham, 1980 "Yesterday afternoon set in misty and cold. I had half a mind to spend it by my study fire, instead of wading through heath and mud Emily Bronte 1847. iii ABSTRACT Aspects of the ecology of the moth Coleophora alticolella have been investigated at a low-altitude site on the Cumbrian coast (15m above sea-level) and along a transect from 215m to 6lOm on the western escarpment of the northern Pennines. The larvae of this moth feed on the seeds of the rush Juncus squarrosus. Seed production by the food-plant is reduced at higher altitudes, where the climate is severe. The oviposition period is delayed with increasing altitude and the eggs are laid singly, if there are sufficient oviposition sites on the developing inflorescences. Survival of the larvae, between hatching and establishment inside the food supply, is directly related to the proportion of J. squarrosus florets developing into seed capsules. Consequently, as a result of the progressive reduction in seed capsule production with increasing altitude, there is greater mortality during larval establishment at the higher sites. Larval case production and subsequent migration to overwinter in the leaf- litter are both retarded with increasing altitude, provided that the food supply is adequate. Both the number of species of parasitic Hymenoptera, attacking the larvae, and the percentage parasitization are reduced with increasing altitude. .Hyperparasitoids were present at the lowest site. Starvation of the larvae, as a result of the reduced J. squarrosus seed production, was the most important mortality factor in the population dynamics of Coleophora alticolella at the highest altitudes. Parasitoids controlled the population at the lowest altitude. Between these two extremes, competition for food by the fourth instar larvae is most important. This acts as a density-dependent factor, reducing natality in the following spring. The larvae often eat all of the seeds produced by the food-plant in this middle region, but not at sites of higher or lower altitude. iv ACKNOWLEDGMENTS It is a pleasure to thank Dr John Coulson for his guidance and supervision during this study. I am grateful to Professor D.Barker for providing facilities in the Department of Zoology at the University of Durham. 1 should also like to thank Mr M.Rawes for permission to work at the Moor House National Nature Reserve, Lord Carlisle for granting access to the Drigg study area and many farmers in Cumbria for allowing me to conduct my research on their land. I am indebted to Dr Z.R.j.Boucek, Dr M.G. Fitton, Mr I.D. Gauld, Dr M.W.R. de V. Graham, Dr J.S. Noyes and Dr G.J. Robinson for identifying insect specimens. Many thanks also go to: Tony Baker, John Chardine, Steven Randall and Ray Williams for helping with fieldwork; Len and Nic Snowball for hospitality at Dufton,- Dr A.H. Seheult for mathematical advice and Mrs R.L. Reed for typing the manuscript. I would also like to thank the staff and research workers at Durham for helpful and constructive discussion, particularly Dr Lewis Davies, Tusi Butterfield, Callum Thomas, David Brearey and Steve Courtney. Finally, many thanks go to Amanda Jennings for proof-reading the manuscript and for her patience and encouragement. This work was financed by an N.E.R.C. research studentship. CONTENTS Page ABSTRACT ... ... ... ... ... iii ACKNOWLEDGMENTS iv CONTENTS v 1. INTRODUCTION 1 2. SAMPLE SITES AND GENERAL METHODS 5 2.1 Location of the sample areas ... ... ... 5 2o2 Sample site descriptions ... ... ... ... 5 2.3 General methods ... ... ... ... ... 9 2.4 The logistic equation — ... ... ... 12 3. STUDIES ON JUNCUS SQUARROSUS, THE LARVAL FOOD-PLANT ... 14 3.1 Introduction ... ... ... ... ... ... 14 3.2 The density of Juncus squarrosus inflorescences 16 3.3 Floret production ... ... 20 3.4 The effect of altitude on flowering phenology 21 3.5 Seed capsule production ... ... ... ... 22 3.6 The number of seeds per capsule ... ... 30 3.7 Grazing of inflorescences ... ... ... 31 3.8 The density of seeds produced ... ... ... 33 3.9 Discussion ... ... ... ... ... ... 34 4. STUDIES ON THE EGGS AND OVIPOSITION BEHAVIOUR OF COLEOPHORA ALTICOLELLA 37 4.1 Introduction ... ... ... ... ... ... 37 4.2 The size of the eggs ... ... ... ... 38 4.3 The colour of the eggs ... ... ... ... 38 4.4 Oviposition sites ... ... ... ... ... 38 4.5 The density of eggs in relation to the Juncus squarrosus florets ... ... ... 40 4.6 The rate of egg-laying „ „, ... ... ... 44 4.7 The median date cf the oviposition period in relation to e.ltituds ... ... ... 47 vi Page 4.8 Ovipositlon behaviour in relation to food-plant development ... ... ... ... 48 4.9 The effect of inflorescence density on oviposition behaviour ... ... ... ... 59 4„10 Discussion ... ... ... ... ... ... 63 5. STUDIES ON THE LARVAE OF C0LE0PH0RA ALTICOLELLA ... 68 5.1 Introduction ... ... ... ... ... ... 68 5.2 Larval density in relation to the food supply 69 5.3 Larval distribution in relation to the seed capsules ... ... ... ... ... ... 71 5.4 The survival of Coleophora alticolella from egg to establishment in the seed capsule ... 74 5.5 The upper altitudinal limit of larval distribution ... ... ... ... ... 78 5.6 A comparison of larval development at 15m and 455m in 1978 ... ... ... ... ... 81 5.7 Larval case production and larval migration .. 86 5.8 Larval feeding after case production ... ... gg 5.9 Discussion ... ... ... ... ... ... lol 6. STUDIES ON THE PARASITOIDS ATTACKING COLEOPHORA ALTICOLELLA 105 6.1 Introduction ... ... ... ... ... ... io5 6.2 The species of parasitoid attacking Coleophora alticolella ... ... ... 106 6.3 Elachertus olivaceus ... ... ... ... 108 6.4 Scambus brevicornis ... ... ... ... 119 6.5 Euderus viiidis ... ... ... ... ... 121 6.6 Pteromalus semotus ... ... ... ... ... 121 6.7 Gelis op. ... ... ... ... ... ... 122 6.8 Hyperparasitization by Tetrastichus endemus .. 123 Page vii 6.9 The parasitoid sex-ratios ... ... ... 125 6.10 The phenology of parasitoid attack ... 129 6.11 Parasitization of Coleophora alticolella larvae in relation to altitude ... 131 6.12 The diversity of the parasitoid guild in relation to altitude ... ... ... 136 6.13 A summary of the relationships between Coleophora alticolella and its larval parasitoids ... ... ... ... 140 6.14 Discussion ... ... , .. 140 POPULATION STUDIES ON COLEOPHORA ALTICOLELLA 146 7 a 1 Introduction ... ... ... ... 146 7„2 The life-tables 146 7.3 Analysis of the life-tables ... ... 157 7.4 Changes in the mortality in relation to altitude ... ... ... ... ... 170 7.5 A comparison of the survivorship curves at 15m and 455m ... ... ... ... ... 171 7.6 Discussion ... ... ... ... ... 172 THE EFFECT OF COLEOPHORA ALTICOLELLA ON THE SURVIVAL OF JUNCUS SQUARROSUS SEEDS 177 8.1 Introduction ... ... ... ... ... 177 8.2 Seed capsule production in 1978 ... 178 8.3 Seed capsule predation in 1978 ... ... ... 179 8.4 Seed capsule survival in 1978 ... ... ... 182 8.5 Seed survival in other years ... ... ... 184 8.6 Discussion ... ... ... ... ... 186 GENERAL DISCUSSION 188 SUMMARY 197 REFERENCES 203 APPENDICES ... 218 CHAPTER 1 INTRODUCTION Animal populations fluctuate around a mean level of abundance. However, this mean level is not the same in all parts of the animal's range; it is generally lower towards the edge of the distribution (Klomp 1962). Despite this, insects, and other animals, are usually studied where they are most abundant and conditions are favourable (Wilson 1968). Huffaker (1958), among others, has stressed that insects should be considered in both salubrious and hazardous environments. Few studies have actually achieved this, a notable exception being that of Whittaker (1971) . This thesis describes the results of an investigation into the ecology of the moor rush moth Coleophora alticolella Zeller, over a large altitudinal range in northern England; from 15m to 610m above sea-level. Previous ecological studies of this species have mainly concentrated on the upper part of this range (Pearsall 1950;
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