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Climate Matching in the Colonisation of Biological Control Agents Against Chrys Anthemoides Moniliferø and Murrubium Vulgure

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Climate Matching in the Colonisation of Biological Control Agents Against Chrys Anthemoides Moniliferø and Murrubium Vulgure Climate Matching in the Colonisation of Biological Control Agents against Chrys anthemoides moniliferø and Murrubium vulgure Craig R. Clarke Thesis submitted for the degree of Doctor of Philosophy Department of Applied and Molecular Ecology Facuþ of Agriculture and Natural Resource Sciences University of Adelaide March, 200I DECLAIL{TION This work contains no material which has previously been accepted for the award of any other degree or diploma in any university or other tertiary institution. To the best of my knowledge and beliet it does not contain any material previously published or written by another person, except where due reference has been made, I give consent that this work may be photocopied or loaned from the University library. Signed Date 30 to ot Craig R. Clarke 2 ACKNOWLEDGEMENTS I would like to express my sincere thanks to: o Ltzfor her support, love, encouragement, enthusiasm, understanding, proof reading and assistance with the counting of a seemingly endless number of insects. ¡ Dr Mike Keller and Dr Rick Roush for their support, enthusiasm and academic guidance a The CRC for Weed Management Systems and the South Australian Pastoral Board for funding the project. o Professor Steve Powles for encouraging me to undertake the PhD. a Petra Mueller, Penny Edwards and Jean-Louis Sagliocco for the collection of the insects from their native range. a Helena Oakey and Dr Ari Verþla for their assistance with the analysis of the plume moth field data. a Dr Marianne Horak from CSIRO, Entomology for her investigation into the taxonomy of the two bitou tip moth populations. a Robin Adair, AltneBruzzese, John'Weiss and Emma Wills from the Keith Turnbull Research Institute for advice about the rearing of the insects, assistance with the importation of the insect populations and information about past releases. a Mark Adams from the South Australian Museum for his assistance with the allozyme electrophoresis study, enthusiasm and academic advice. a Tom for his assistance evaluating the Finders Ranges plume moth release sites. o Professor Dudþ Pinnock fromthe University of Adelaide for his advice regarding the plume moth disease. o The Animal and Plant Control Commision, Monarto Zoo andNatinal Parks and Wildlife for assistance in selecting release sites. a Terry Feckner, Dr Garry Taylor and Ernie Nagy for their management and organisation of facilities. a Other members of the Department of Applied and Molecular Ecology and South Australian Research and Development Institute, Entomology for their advice and making the work environment pleasant and inspiring. a My trusty sawn-offshovel for warding offthe snakes. J TABLE OF CONTENTS Abstract I Chapter I Introduction and project aims 10 1.1 Introduction 10 t.2 Climate matching l4 1.2.I Reviews of past bio-control programs - importance of climate 1.2.2 Climatic adaptation- importance to bio-control 1.2.3 Climatemodelling 1.2.4 Other key attributes that vary between populations 1.2.5 Conclusions drawn fromthe literature 28 1.3 Project aims Chapter 2 The target weeds and the history of the biological control 30 progr¡ms against them 2.1 Target no. 1: horehound 30 2.1.1 Growth and development 2.1.2 Seed dispersal 2.1.3 Importance 2.1.4 Control 2.2 Biological control of horehound 34 2.3 Target no. 2z Chrysanthemo¡des monilífera 36 2.3.1 Growth and development 2.3.2 Seed dispersal 2.3.3 Importance 2.3.4 Control 2.4 Biological control of C hrysanthemoìdes monìlifurø 40 2.4.1 Comostolopsisgermana Chapter 3 Insect population origins, importation and maintenance 43 3.1 Introduction 43 3.2 Origins of the horehound plume moth populations 44 3.3 Maintenance of the plume moth laboratoqy cultures 47 3.3.1 Rearing 3.3.2 Disease 3.4 Origins of the bitou tip moth populations 51 3.4.1 Rearing Chapter 4 Climate matching in the plume moth: how close is close enough? 57 4.1 Introduction 57 4.2 Materials and methods 57 4.2.1 Experiment 1: the influence of temperature on egg development time 4.2.2 Experiment 2: the influence of temperature on larval development 4 4.2.3 Experiment 3: the influence of temperature on fecundity 4.2.4 Experiment 4: effect of short bursts of a stressful temperature on larval survival 4.2.5 Experiment 5: effect of water stress on host plant quality 4.2.6 Experiment 6: diapause 4.2.7 Experiment 7: field releases 4.3 Results 73 4.3.1 Experiment 1: the influence of temperature on egg development time 4.3.2 Experiment 2: the influence of temperature on larval development 4.3.3 Experiment 3: the influence of temperature on fecundity 4.3.4 Experiment 4: effect of short bursts of a stressful temperature on larval survival 4.3.5 Experiment 5: effect of water stress on host plant quality 4.3.6 Experiment 6: diapause 4.3.7 Experiment 7: comparison of populations in field releases 4.4 Discussion 92 Chapter 5 Effects of lab rearing and inbreeding in the plume moth 97 5.1 Introduction 97 5. 1 . 1 Genetic variation within populations 5.1.2 Laboratory rearing 5.2 Materials and methods 10s 5.2.1 Experiment 1: recþocal crosses 5.2.2 Experiment 2: fecundity of field-collected and lab- collected pupae 5.2.3 Experiment 3: inbred lines 5.2.4 Experiment 4: allozyme electrophoresis studies 5.3 Results 111 5.3.1 Experiment l: reciprocal crosses 5.3.2 Experiment 2: fecundity of field-collected and lab- collected pupae 5,3.3 Experiment 3: inbred lines 5.3.4 Experiment 4: alToryme electrophoresis studies 5.4 l)iscussion 117 Chapter 6 Plume moth release technique: caged ven¡us free release t2r 6.1 Introduction tzt 6.2 Materials and methods 123 6.3 Results 124 6.4 I)iscussion 125 5 Chapter 7 Climate matching in the bitou tip moth: how close is not close 128 enough? 7.1 Introductlon r28 7.2 Materials and methods r29 7.2.1 Experiment l: the influence of temperature on egg development time 7.2.2 Experiment 2: the influence of temperature on larval development 7.2.3 Experiment 3: effect of short bursts of a stressful temperatute on larval survival 7.2.4 Experiment 4: survival during more realistic "heatwave" conditions 7.2.5 E4periment 5: effect of plant sub-species and plant qualitY 7.2.6 Experiment 6: fecundity of moths reared on healtþ and water stressed boneseed and bitou bush 7.2.7 Experiment 7: field releases 7.2.8 Experiment 8: predation by ants 7.3 Results t4t 7.3.1 Experiment 1: the influence of temperature on egg development time 7.3.2 Experiment 2:the influence oftemperature on larval development 7.3.3 Experiment 3: effect of short bursts of a stressful temperature on larval survival 7.3.4 Experimerrt 4: survival during more realistic "heatwaveo' conditions 7.3.5 Experiment 5: effect of plant sub-species and plant quality 7.3.6 Experiment 6: fecundity of moths reared on healthy and water stressed boneseed and bitou bush 7.3.7 Experiment 7: field releases 7.3.8 Experiment 8: predationbyants 7.4 I)iscussion 157 Chapter 8 Speciation in the bitou tip moth 163 8.1 Introduction 163 8.1.1 Importance of taxonomy to biological control 8.1.2 Genetic variation and the mixing or hybridisation of populations 8.2 Materials and methods 166 8.2.1 Experiment 1 : reciprocal crosses - fecundity, %o egg hatch and Fl survival 8.2.2 Experiment 2: alToryme electrophoresis studies 5.2.3 Experiment 3: isolation between insect populations under semi-realistic conditions 8.2.4 Experiment 4: host specificity testing 8.2.5 Taxonomy 6 8.3 Results 172 8.3.1 Experiment 1: reciprocal crosses - fecundity,%o egg hatch and Fl survival 8.3.2 Experiment 2: alloryme electrophoresis studies 8.3.3 Experiment 3: isolation between insect populations under semi-realistic conditions 8.3.4 Experiment 4: host specificity testing 8.3.5 Taxonomy 8.4 I)iscussion 180 Chapter 9 Conclusions and general discussion 183 9.1 Reasons for the failure of the original plume moth and 183 bitou tip moth populations 9.2 Climate matching 185 9.3 Insect quality and genetic variation of the agent 187 9.4 Release technique 188 9.5 Predation 190 9.6 Further research 19t Appendix 1 Plume moth field evaluation dates 193 Àppendix 2 Plume moth field data anaþsis 194 Appendix 3 Dispersion of the plume moth from the release point 23s References 244 7 ABSTRACT Approximately 40Yo of releases of classical biological control agents targeting weeds fail to establish in the field and 70Yo fatlto have a significant impact on the target weed (Julien et ol,, l9S4). Many theories have been proposed to explain these failures but, to date, little experimental research has been conducted to test them. In this project, several theories relating to the establishment ofbiological control agents were tested using two model systems, the bitou tip moth, Comostolopsis germana. for the control ofthe weed Chrysanthemoides monílifera and the plume moth, Wheeleria spilodactylus, for the control of horehound, Marrubium vulgare. The importance of climate matching, predation, release technique, laboratory adaptation and inbreeding to the establishment ofthese agents was investigated. In both examples, it was believed that thç moth populations failed to establish in South Australia because they were poorly adapted to South Australia's Mediterranean climate, particularly its dry, hot summers. The original imported populations ofthe bitou tip moth (from Durban, South Africa) were successfully established in coastal areas ofNew South 'Wales, which have a sub-tropical to warm temperate climate.
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