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Behaviour and Ecology of the Primary Parasitoids Cotesia Urabae And

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Behaviour and Ecology of the Primary Parasitoids Cotesia Urabae And V/A¡TE INS IIU'I' F. 4.1 .4o LIEiT,\RV Behaviour and Parasitoids Cotesia urabae anù (HYmenoPtera: Braconidae) and (LePidoPtera: By Geoffrey Rowland Allen B.Sc. Qlons) The University of N.S-V/. A thesis submitted for the Degree of Doctor of Philosophy in the Faculty of Agricultural Science at The University of Adelaide- 11 TO MY PARENTS 111 Table of Contents Page SUMMARY xiv DECLARATION xvü ACKNOWLEDGMENTS xvüi PREFACE 1 CHAPTER 1. INTRODUCTION TO U. LUGENS AND ITS BIOLOGY. 4 Taxonomy and distribution 4 Host plants 5 General biologY 6 Phenology 8 Management and control of U- lugens 10 CHAPTER 2. BIOLOGY OF THE PARASITOIDS OF U. LUGENS AND SURVIVAL OF LARVAE OF U. LUGENS IN TIIE FIELD IN SOUTH AUSTRALIA. 11 Abstract 11 Introduction 11 Materials and methods t2 Results 15 1. The parasitoid comPlex 15 2. Survival of. U. lugens larvae in the field 25 Discussion 29 CHAPTER 3. TTIE PHENOLOGIES OF C. URABAE, D. EUCALYPTI AND THEIR HOST U. LUGENS IN TI# ADELAIDE REGION. 33 Abstr¿ct 33 Introduction 33 Materials and methods 35 1. Field monitoring 35 2.Placement of parasitized and unparasitized larvae in the field 3l 3. Analysis of data 39 lv Results 39 1. Phenology of U. lugens 39 2. Phenologies of C. urabae andD. eucalypti 42 Discussion 51 CHAPTER 4. TT{E SIZE OF ADULT C. URABA¿'AND D. EUCALYPTI IN RELATION TO TTIEIR HOST U. LUGENS. 57 Abstract 57 Introducúon 57 Materials and methods 59 1. Analysis of data 60 Results 60 Discussion 65 CHAPTER 5. TEMPERATURE AND TTIE DEVEI,OPMENT OF U. LUGENS, C. URABAE AND D. EUCALYPTI. 69 Abstract 69 Introduction 70 Materials and methods 72 1. Experiment 1: Larval development of U. Iugens, and preimaginal deveþment of D. eucalypti and C. urabae from small and mid hosts 72 a. Experimental protocol 72 b. Analysis of data 74 2. Experiment2:Eggdevelopment of U. lugens 75 3. Experiment 3: Pupal developmentof U. fugens 75 4. Experiment 4: I-ongevity of adult C. urabae and D. eucalypti 76 5. Simulation of Phenologies 76 Results 77 1. DeveloPment of U. lugens 77 a. Egg development 77 b. Larval develoPment 79 c. Pupal development 84 2. Development of C. urabae andD. eucalypti 84 a. Egg-lanral development 84 b. Pupal development 90 c. V/eight of adult parasitoids 90 v d. Longevity of adult C. urabae and D. eucalypti 91 3. Simulation of phenologies 9T Discussion 97 ¡/, CHAPTER 6. BEHAVIOURAL INTERACTIONS BETTVEEN U. LUGENS, C. URABAE AND D, EUCALYPTI. 103 Abstract 103 Introduction 104 Materials and methods 105 1. Experimental protocol 105 2. Analysis of data 108 Results 109 Discussion tzl 1. Gregariousness and defensive behaviour of U. lugens t2r 2. Experimental design , U. lugens defensive behaviour and superparasitism t25 3. Host acceptance and the success of oviposition of C. urabae andD. eucaþPti 127 CHAPTER 7. LARVAL STAGE AND RESPONSE OF U. LUGENSTO ATTACIC 130 Abstract 130 Introduction 130 Materials and methods 131 1. Experiment 1: Setal and body dimensions of larvae of U.lugens t32 2. Experiment 2: Response of U. lugens to attack. r33 3. Experiment 3: Response of U. lugetu to repeated attack 135 Results 136 1. Experiment 1: Setal and body dimensions of larvae of U.lugens t36 2. Experiment 2: Response of U. lugens to attack 138 a. Response to attack 138 b. Type of defensive response t4t c. Prior U. lugens behaviour and response to attack 148 3. Experiment 3: Response of U. lugens to repeated attack 148 Discussion 149 vi CHAPTER 8. GENERAL DISCUSSION 155 APPENDIX 1. PARASITOIDS OF U. LUGENS IN SOUTH AUSTRALIA, WITHDESCRIPTION OF TWO NEW SPECtsS OF BRACONIDAE. 160 APPENDIX 2. LABORATORY REARING OF U. LUGENS, C. URABAE AND D. EUCALYPTI. 161 APPENDX 3. TT{E REI-ATIONSHIP BETWEEN HIND TIBIAL LENGTH AND TTIE CUBE ROOT OF TTIE WEIGHT OF AN ADULT FEMALE FOR C. URABAE AND D. EUCALYPTI. 166 REFERENCES t67 vtl List of Figures Page CHAPTER 2 Fig. 1. The number of larvae of U. lugens that were found during the 1985- 1986 summer generation of U.lugens 26 Fig.2. The proportion of larvae of U. fugenr found in each instar during the 1985- 1986 summer generation 26 Fig.3. The percentage of the larval population of U. lugens remaining on29 different E. microcarparîßsduring the 1985-1986 summer genefation 27 Fig.4. The meantSE percentage of larvae remaining on E. microcarpatees at freld site 3 during the 1986 winter generation of U.lugens, when larvae were either enclosed in a fine gau7Ê sleeve cage or left uncaged 28 Fig.5. The proportion of larvae of U. Iugens in each instar of the uncaged population of larvae during ttre 1986 winter generation 28 CHAPTER 3 Fig. 1. Phenology of U. lugens from September 1985 to December 1987 inclusive 40 Fig. 2. The temporal pattern of pupation of C. urabae andD. eucalypri in each host generation as determined by weekly sampling for cocoons in the field 44 Fig. 3. Th, .,ution of final host sizes, as determined by head capsule width, from which C. urabae andD. eucalyptí emerged to pupate in the field 46 Fig. 4. The distribution of final host sizes, as determined by dry weight of host, from which C. urabae andD. eucalypti emerged to pupate in the field 47 Fig. 5. Proportion of the total number of trees with host populations of vlll [I.lugens when the first cocoon of either parasitoid species appeared, that subsequently had cocoons of C . urabae ot D . eucalypti collected upon them 50 CHAPTER 4 Fig. 1. Exit position of larvae of C. urabae and D. eucalypri along their host's body length for different sizes (head capsule widths) of host 61 Fig.2. The relationship between female parasitoid weight and the number of fully developed eggs held by a female at the time of emergence from the pupal cocoon, and upon death when left unfed at2æC andTíVo relative humidity for C. urabae and D. eucalpti 66 CHAPTER 5 Fig. 1. The percentage of U.lugens eggs that faited to develop and developed but failed to hatch over five temperatures 78 Fig.2. Distributions of the number of lanral instars for male and female U.lugens reared at three different temperatures 15o, 20o, and 25oC 81 Fig. 3. The sizes of head capsules shed by larvae of U. fugens prior to the conìmencement of head capsule stacking when reared at three different temperatures 15o, 2@, and25oC 82 Fig. 4. The mean head capsule width of each instar of U. fugens for larvae ttrat underwent from í-l4larvalinstars before pupation across all three temperatures 83 Fig. 5. The relationship between temperature and rate of egg-lanral development of D. eucalypti when developing from small hoss 87 Fig. 6. Host head capsule width of U.lugent at the time of parasitoid emergence for c. urabae deveþing from small hosts, c. urabae deveþing from mid hosts, and D. ercalypti developing from small hosts at 15o,20o, and 25oC 88 IX Fig. 7. Host dry weight of U. lugens at the time of parasitoid emergence for C. wabae developing from small hosts, C- urabae developing from mid hosts, andD. eucalypti developing from small hosts at 15o,20o, and 25o,C 89 Fig. 8. Observed and predicted durations of the life stages of U. lugens, and observed and predicted appeafances of cocoons of C. urabae and D. eucalyprj for 1985, 1986, and 1987 94 CHAPTER 6 Fig. 1. The relative sizes of the first 9 instars of U.lugens reared at 20oC 106 Fig.2. Behaviour of U. lugens and pamsitoids of U. lugens 110 Fig. 3. The mean number of visits made by C. urabae andD. eucalypti to a patch of 40 small, 40 mid, and 40 latge U- lugens over 20 min' rt2 Fig. 4. The outcomes of all visits made to patches of 40 small, 40 mid, and 4O large U.lugensby D.eucalypti and C- urabae 113 Fig. 5. The number of times each mid or large U. lugens was €ncountered by C. urabae and by D. eucalYPti 116 Fig. 6. The number of larvae displaying defensive behaviour immediately following the visit by each D. eucalypti, and each C . urabac , to a patch of U. fugens at the beginning of each quarter of the total number of visis made by that Parasitoid 119 Fig.7. The outcome of encounters between C. urabae, D. eucalypfi and the mid or large larvae of U. lugens when, immediately prior to the encountef, the larvae were either rearing or thrashing or not rearing or thrashing t20 Fig. 8. The mean number of ovipositions, of visits, and time spent in the patch, for each 5 min. of the 20 min. observation for c. urabae, and D. eucalypri with the three sizes (small, mid, large) of U- lugens t22 x CHAPTER 7 Fig. 1. MeantSE measurements of each larval instar of U. lugens for body dimensions, the longest seta surrounding the body from three directions, and the highest seta above three regions of a larva's body 137 Fig.2. Frequency of defensive, locomotory and no response recorded as immediate (Ð and ultimate (u) response to attack with a micropin to small, mid and large larvae of U. lugens r39 Fig. 3. Subdivision of immediate defensive fesponse for small, mid, and large larvae of U.lugers into pulse, rear and head curl when attacked with a micropin from the front, from the side and from behind t42 Fig. 4. Frequency of regurgitation, thrashing and both these responses occurring together when in conjunction with pulsing, rearing and head curling for small, mid and large larvae of U.
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