Diptera: Agromyzidae)
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È ,\l []. li,ri I II-tjTH 3 .í .93 LIiJRA}TY COMPARATIVE HOST STAGE UTILIZATION oF TWO PARASITOIDS OF Liriomyza brassicae (DIPTERA : AGROMYZIDAE) BY ROSS MICHAEL LARDNER B. Sc. (Hons) Macquarie University A thesis submitted for the Degree of Doctor of Philosophy r¡r the Faculty_of ;gr;îtu;l -¿ Nutural Resõurce Sciences at The Universiry of Adelaide. November 1991 To My FamilY ru TABLE OF CONTENTS Pa ge List of Figures vlll xii List of Tables xvi Summary xviii Declaration Acknowledgments xix CHAPTER ONE. INTRODUCTION 1 CHAPTER TIryO. THE BIOLOGY AI{D PHENOLOGY oF Liriomyza brassícae (DIPTERA : AGROMYZIDAE) IN THE ADELAIDE REGION 8 2.L. Abstract 8 2.2. Introduction 9 2.3. Materials and Methods 11 2.3.1. Instar Determination ll 2.3.2. DeveloPment 11 2.3.3. Fecundity and l-ongevity of Mated Females t2 2.3.4. Longevity of Unmated Flies t2 2.3.5. Field SurveY 13 2.4. Results 18 2.4.1. Instar Determination 18 2.4.2. DeveloPment 20 2.4.3. FecunditY and l,ongevitY 20 2.4.4. Field SurveY 25 2.5. Discussion 39 CHAPTER THREE. THE BIOLOGY OF PARASITOIDS ASSOCIATED WITH Líriomyza brassicae IN THE ADELAIDE REGION 44 44 3.1. Absract 45 3.2. Introduction 47 3.3. Materials and Methods 49 3.4. Results lv 3.4.1. OPius atricornis 53 3.4.2. OPius cinerariac 53 3.4.3. Chrysocharis Pubicornis 56 3.4.4. ChrysonotomYia sP. 56 3.4.5. Closterocerus sP. 57 3.4.6. H emiPtarseruts varicornís 57 3.4.7. Zagrammosomo sP. 58 3.4.8. Culturing Parasitoids 60 3.5. Discussion 60 CHAPTERFOUR.ATTACKoFHOSTSTAGESBY NAM FEMALES OF Hemiptarsenus varicornis AND Opius cinerariae 67 4.L. Abstract 67 4.2. Introduction 68 4.3. Materials and Methods 70 4.3.1. No-choice ExPeriments 70 4.3.2. Choice ExPeriments 7l 4.3.3. Statistical AnalYsis 72 4.4. Results 72 4.4.1. No-choice ExPeriments 72 4.4.1.1. Parasitism 72 4.4.1.2. Host MortalitY 81 4.4.2. Choice ExPeriments 86 4.4.2.1. Parasitism 86 4.4.2.2. Host MortalitY 91 4.5. Discussion 96 CHAPTER FIVF.. ATTACK OF HOST STAGES BY EXPERIENCED FEMALES OF Hemiptarsenus varicornis AND Opius cinerariae 103 5.1. Abstract 103 5.2. Introduction 104 5.3. Materials and Methods 105 5.3.1. No-choice ExPeriments 105 5.3.2. Choice ExPeriments 105 5.3.3. Statistical AnalYsis 105 5.4. Results 106 5.4.1. No-choice ExPeriments 106 5.4.2. Choice ExPeriments 109 5.5. Discussion 109 v CHAPTER SIX. SEX ALLOCATION BY Hemiptarsenus varicornis AND OPius cinerariae rt7 6.1. Abstract tr7 6.2. Innoduction 118 6.3. Materials and Methods 120 6.4. Results t22 6.4.1. No-choice ExPeriments r22 6.4.2. Choice ExPeriments 135 6.4.3. Comparisoi Between no-choice and choice exPeriments t42 6.5. Discussion r42 CHAPTER SEVEN. RELATIVE FITNESS OF PARASITOIDS DEVELOPING FROM DIFFERENT HOST INSTARS 1s0 7.1. Abstract 150 7.2. Introduction 151 7.3. Materials and Methods t54 7.3.1. DeveloPment t54 7.3.2. LongevitY t54 7.3.3. Egg Load 156 7.3.4. Mating Success 156 7.3.5. Size 157 7.4. Results r57 7.4.1. DeveloPment r57 7.4.2. Size 158 7.4.3. l-ongevitY r65 7.4.4. Egg Load 165 7.4.5. Mating Success r67 7.4.5.1. O. cinerariae t67 7 .4.5.2. H. varicornis r70 7.5. Discussion 175 CHAPTER EIGHT. HOST UTILIZATION BY Hemiptarsenus varicornis AND Opius cinerariae IN RELATIOI{ TO RECENT THEORY 181 8.1. Introduction 181 183 8.2. Summary-Which of Relevant Models 8.2.1. host tYPes to attack? 183 8.2.2. Whether to oviposit or host feed? 184 8.2.3. Which sex to allocate? 184 8.3. Summary of Factors Influencing Host Utilization by H. varicornis and O. cinerariac 186 8.3.1. H. varicornis 187 8.3.2. O. cinerariac 188 8.4. Test of Predictions of Models 189 vl 8.4.1. ProgenY Allocation 189 8.4.2. Host Feeding 190 8.4.3. Sex Allocation 191 S.5.otherApproachestoUnderstandingHostUtilization of H.varicornis andO. cincrariae 194 8.6. General Conclusions 19s APPENDIX ONE. METHODS FOR REARING Liriomvza brassicae IN THE LABORATORY lee 41.1. Method One r99 41.1.1. Rearing r99 Al.I.2. Additional Activities 201 A1.2. Method Two 202 APPENDIX TTryO. LOWER DEVELOPMENTAL THRESHOLDS CALCULATED FROM DEVELOPMENTAL TIMES OF Liriomyza brassicae PUBLISHED BY BERI AND CHANDRA (1983) 206 APPENDIX THREE. ABUNDANCE OF PARASITOIDS ATTACKING Liriomyza brassicae IN THE ADELAIDE REGION AS SHOWN BY FIELD SURVEYS FROM 1989 TO 1991 208 APPENDIX FOUR. CORRELATION OF HIND TIBIA LENGTH WITH HEAD CAPSULE \ryIDTH FOR Opius cínerariae AND Hemiptarsenus varicornis 2t7 APPENDIX FIVE. RELATIONSHIPS BETWEEN RELATIVE FITNESS COMPONENTS OF Hemiptarsenus varicornis AND Opius cinerariae FROM DIFFERENT HOST INSTARS 2r9 vll REFERENCES 228 ADDENDUM TO THESIS Introduction 258 Re-analysis of Sex Allocation Data in Chapter Six 258 Discussion of Examiner's Comments 260 Minor Corrections throughout Thesis 264 vu.l LIST OF FIGURES CHAPTER ONE CHAPTER TWO Fieure 2.1. Location of collection sites for field surveys of L.Vrassicae populations within the Adelaide region' t4 Figure 2.2. Relationship between.temperature .and,.^ ã.irfop.n"ntal fate (mean'+ standard enór) of va¡ious life stages of L. brassicae. 2l Figure 2.3. Relationship between lifespan and the number of viibte eggs laid by female L. brassicae at three constant temperatures (15, 20, 25'c). 24 Figure 2.4. Relationship between-temperature and lifesp.an (mËan * standard error) fôr male and female adult L. brassicae. 26 Figure 2.5. Levels of acrive, dead and^empty L. brassica¿ mines coilected during the first yeaf survey (1988-1989) from various host plants around Adelaide. 27 Figure 2.6. Levels of active, dead and eTPty 1ules of L. brassicae oíT. maius, C. maritima, R. rugosum and S' fficinale during the surveys of years 2-3 (1989-91). 32 Figure 2.7. Population strucrure of active L. brassicae mines collected früir f. majus, C. maritima, R. rugosum and S. officinale during the surveys of years 2-3 (1989-91). 36 CHAPTER THREE Figure 3.1. Phenology of the parasitoids of L. brassicae in the Adelaide region. 54 Figure 3.2. Number of emergences- of each sex of H. varicornis itoï Zn¿ and 3rd instar L. brassicae lanae pooled for all sites during the 2nd and 3rd year surveys (1989-91). 59 lx 74 79 88 Figure 4.4. Fate of L. brassicae exposed to naive, mated and urimated female O. cinerariae and those in control cages for the choice experiment (dau pooled for each type of cage)' 92 t24 r27 131 133 Figure 6.5. Total production of female and male prog.eny by O.'cinerariae ondifferent L. brassicae instars between days I and 10 (pooled) for the choice experiments' r37 Figure 6.6. Proportion males (mean * standard enor) allocated by O.-cinerariae dunng the choice experiments. 140 x CHAPTER SEVEN Figure 7.1. Developmental rates (mean t standard deviation) of O.'A*ror¡ae atth¡eé constant temperaffies for males and females *fti.tt began development in lst inlta¡, 2nd instar and 3rd instar L. brassicae larvae. 160 Figure 7.2. Developmental rate (mean t standard deviation) of r62 H. varicornis at three constant temperatures. Fieure 7.3. Hind tibia lengths (range and mean * standard error) of O.'lni*i¡ae, female H. varícornis añd male H. varicornis in relation to 163 the host insta¡ on which development bgan. Figure 7.4. Longevity (mean 1 standard enor) fot o. cinerariae -ã A. varicornis-in ráaiion to the insta¡ of L. brassicae on which development began. r66 Figure 7.5. Number of mature eggs in relation to body size (hind tibia length) of O. cinerariae. 169 172 CHAPTER EIGHT APPENDIX ONE Figure Al.l. caging sysrem used for ovipositing L. brassicae during rearing method 1. 200 Fieure AI.2. Number of feeding and egg punctures (mãan * standard error) produced by L. brassicae between òg00-t00, 1100-1400, 1400-17ü) and 1700-0800 hours for the 204 first 5 days of oviPosition. APPENDIX TWO APPENDIX THREE xl APPENDIX FOUR Figure 44.1. Conelation between hind tibia length and head capsule 218 wiät¡ for O. cineraríae and H ' varicornis. APPENDIX FIVE Figure 45.1. Relationships between relative fitness components t standa¡d enor) for female progeny of H.varicornis from2nd i"i.- 220 and 3rd insta¡ L. brassicae larvae. Figure A5.2. Relationships between relative fitness components. * standa¡d error) for'male progeny of H. varicornis fromZnd i-?ãn 222 and 3rd instar L. brassicac lan¡ae. Figure 45.3. Relationships between relative fitness components i1¡?- t standard error) for iemale progeny of O. cineraríac ftomeach L. brassicae instar. 224 Figure 45.4. Relation-ships between relative fitness components. (m?an + standa¡d error) for male progeny of O. cinerariac îromeach L. brassica¿ insta¡. 226 xll LIST OF TABLES CHAPTER ONE CHAPTER TWO Table 2.1. Plant species and sitesincluded in the first year survey of L. brassicae pofulations (1983-89). 16 Table 2.2. Length and width of leaf mine, Iength and width of tu*." and mouth--hook length of different life stages of L. brassicae. 19 Table 2.3. Lower temperature thresholds and mean thermal .-*tt-tt for ãevelopment of various life stages of L' brassicac (sexes pooled)."alculated 23 Table 2.4. Fecundity (no. of viable eggs) and lifespan (days) of L. brassicae when kept with plants at three constant temperatures.