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WATER BALANCE, HOST UTILISATION, and MASS REARING IMPROVEMENTS of the COCKROACH OOTHECAL PARASITOIDS, Aprostocetus Hagenowii

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WATER BALANCE, HOST UTILISATION, and MASS REARING IMPROVEMENTS of the COCKROACH OOTHECAL PARASITOIDS, Aprostocetus Hagenowii WATER BALANCE, HOST UTILISATION, AND MASS REARING IMPROVEMENTS OF THE COCKROACH OOTHECAL PARASITOIDS, Aprostocetus hagenowii (HYMENOPTERA: EULOPHIDAE) AND Evania appendigaster (HYMENOPTERA: EVANIIDAE) TEE HUI SIANG UNIVERSITI SAINS MALAYSIA 2016 WATER BALANCE, HOST UTILISATION, AND MASS REARING IMPROVEMENTS OF THE COCKROACH OOTHECAL PARASITOIDS, Aprostocetus hagenowii (HYMENOPTERA: EULOPHIDAE) AND Evania appendigaster (HYMENOPTERA: EVANIIDAE) BY TEE HUI SIANG Thesis submitted in fulfillment of the requirements for the degree of Doctor of Philosophy APRIL 2016 ACKNOWLEDGEMENTS I would like to thank my advisor Prof. Lee Chow Yang for his academic guidance, support and expertise, which are fundamental to the completion of this thesis, fulfilling the academic requirements for graduation. Nevertheless, his mentoring, wisdom, enthusiasm, insightfulness, resourcefulness and kindness have inspired, motivated and guided me while pursuing my doctorate studies at his Urban Entomology Laboratory at USM. I am indebted to him for all these. My sincere thanks to Dr. Yoshitaka Kamimura (Keio University, Japan) for providing valuable discussions on my research and his time reviewing my manuscripts. I am grateful to past and present Urban Entomology Labmates, Dr. Neoh Kok Boon, Dr. Bong Lee Jin, Dr. Lee Ching Chen, Dr. Ang Ling Hui, Dr. Foo Foong Kuan, Dr. Tay Jia Wei, Dr. Veera Singham, Nellie Wong Su Chee, Goh Xin Yu, Ikhsan Guswenrivo, Nadiah Mohamed, Nurainie Ismail, Chris Kim Daeyun, Sophia Liew Chiew Liang, Mai Duyen, Mark Ooi Chuan Sen, Jong Zheng Wei, Amy Low Kar Yin and Kathleen Low for their friendship, assistance and support. I would like to thank to Prof. Zairi Jaal, Dr. Hamdan Ahmad as well as the staffs of Vector Control Research Unit, particularly Mr. Adanan Che Rus, Mr. Abdul Malik Yahaya, Mr. Rohaizat Baba, Mr. Haslan Salim, Mr. Mohd Nasir Hassan, Mr. Mohd Nasir Mohd Noor, Mr. Muhammad Noor Hisham Halim, Eugene Chan, Ms. Ashiah Abu Seman and Ms. Nurul Mazni Yusuf for their assistance and technical support (laboratory facilities, materials, etc.). I am grateful to Universiti Sains Malaysia (USM Fellowship) and Agricultural Crop Trust for their financial support of my PhD studies. ii My special thanks to my wife, Shin Yih, and my parents. This thesis would not have been completed without their endless support. iii TABLE OF CONTENTS Acknowledgements ii Table of contents iv List of Tables viii List of Figures ix List of Plates xiv Abstrak xv Abstract xviii CHAPTER ONE ‒ INTRODUCTION 1 CHAPTER TWO ‒ LITERATURE REVIEW 2.1 Periplaneta americana 5 2.2 Aprostocetus hagenowii 8 2.3 Evania appendigaster 11 2.4 Interspecific competition and coexistence of competing 13 species in parasitoid wasps 2.5 Water balance in insects 15 2.6 Factors influenc ing host selection and foraging in parasitoids 17 2.7 Rearing of A. hagenowii and E. appendigaster for biological 21 control of cockroaches 2.8 Biological control efficacy of A. hagenowii and E. 24 appendigaster CHAPTER THREE ‒ WATER BALANCE PROFILES, HUMIDITY PREFERENCE AND SURVIVAL OF TWO SYMPATHRIC COCKROACH OOTHECA PARASITOIDS Evania appendigaster and Aprostocetus hagenowii 3.1 Introduction 25 iv 3.2 Materials and methods 27 3.2.1 Rearing of E. appendigaster and A. hagenowii 27 3.2.2 Water balance profile 30 3.2.3 Survival of parasitoids at different RHs in the presence 31 and absence of a sugar solution 3.2.4 RH preference 32 3.2.5 Data analysis 34 3.3 Results 36 3.3.1 Water balance of E. appendigaster and A. hagenowii 36 3.3.2 Survival time of E. appendigaster reared at different 40 RHs 3.3.3 Survival time of A. hagenowii reared at different RHs 40 3.3.4 RH preference of E. appendigaster 43 3.3.5 RH preference of A. hagenowii 45 3.4 Discussion 47 CHAPTER FOUR – OOTHECA AGE AND TEMPERATURE EFFECTS ON THE LIFE HISTORY OF COCKROACH OOTHECA PARASITOID Aprostocetus hagenowii 4.1 Introduction 55 4.2 Materials and methods 57 4.2.1 Insect rearing 57 4.2.2 Influence of age of oothecae incubated at different 58 temperatures on fitness traits of A. hagenowii 4.2.3 Influence of temperature on immature development of 59 A. hagenowii 4.2.4 Influence of temperature on survival, parasitism 60 activities and lifetime realized fecundity of A. hagenowii v 4.2.5 Data analysis 62 4.3 Results 64 4.3.1 Influence of age of oothecae incubated at different 64 temperatures on fitness traits of A. hagenowii 4.3.2 Influence of temperature on immature development of 66 A. hagenowii 4.3.3 Influence of temperature on survival and parasitism 70 activities of A. hagenowii 4.4 Discussion 74 CHAPTER FIVE ‒ COCKROACH OOTHECA PARASITOID Evania appendigaster EXHIBITS OVIPOSITION PREFERENCE TOWARDS OOTHECA AGE MOST VULNERABLE TO HOST CANNIBALISM 5.1 Introduction 78 5.2 Materials and methods 81 5.2.1 Cockroach and parasitoid cultures 81 5.2.2 Cannibalism of healthy and parasitoid-parasitized 82 oothecae of different ages by P. americana 5.2.3 Ootheca age preference of E. appendigaster 83 5.2.4 Handling time and fitness traits of E. appendigaster on 85 different-aged oothecae 5.2.5 Data analysis 87 5.3 Results 88 5.3.1 Cannibalism of healthy and parasitoid -parasitized 88 oothecae of different ages by P. americana 5.3.2 Ootheca age preference by E. appendigaster 91 5.3.3 Handling time and fitness traits of E. appendigaster on 91 different-aged oothecae 5.4 Discussion 96 vi CHAPTER SIX ‒ FEASIBILITY OF COLD-STORED Periplaneta americana OOTHECAE FOR REARING THE OOTHECA PARASITOIDS Aprostocetus hagenowii and Evania appendigaster : EFFECT OF OOTHECA AGE AND STORAGE DURATION 6.1 Introduction 101 6.2 Materials and methods 10 3 6.2.1 Rearing of P. americana 103 6.2.2 Rearing of A. hagenowii and E. appendigaster 104 6.2.3 Effect of heating on hatchability of oothecae 105 6.2.4 Rearing of A. hagenowii and E. appendigaster from 10 6 live, heat-killed, and cold-stored oothecae aged 1–4 weeks old 6.2.5 Data analysis 108 6.3 Results 110 6.3.1 Effect of heating on hatchability of oothecae 110 6.3.2 Biological parameters of A. hagenowii reared on live, 110 heat-killed, and cold-stored oothecae 6.3.3 Biological parameters of E. appendigaster reared on 117 live, heat-killed, and cold-stored oothecae 6.4 Discussion 121 SUMMARY AND CONCLUSION 12 8 REFERENCES 13 5 List of Publications and Presentations 155 vii LIST OF TABLES Page Table 3.1 Fresh mass, percentage of to tal body water, cuticular 37 permeability, and surface area to volume ratio (mean ± SE) of males and females of E. appendigaster and A. hagenowii . Table 3.2 Statistics for the linear regression between cumulative 39 %TBW loss and desiccation time (h) for males and females of E. appendigaster and A. hagenowii . Table 4.1 Mean (± SE) percentage emergence, development time 68 (days), number of progeny/ootheca and percentage female progeny of A. hagenowii emerged from P. americana oothecae at 20 ‒35 °C. Table 5.1 Results of three-way ANOVA of the effects of ootheca 89 age, cockroach life stage, and ootheca parasitism status on percentage cannibalism of oothecae in P. americana. Table 6.1 Two -way ANOVA of the % parasitism, % emergence and 11 2 log of the developmental time (days) for A. hagenowii and E. appendigaster . Table 6.2 The effect of progeny sex and cold storage conditions on 122 developmental time of E. appendigaster . viii LIST OF FIGURES Page Figure 3.1 Diagram of the linearly connected chambers. Four 33 different salt solutions (MgCl 2, NaBr, NaCl, and KNO 3) were used to create a gradient of increasing RH among the chambers. Figure 3.2 Cumulative % total body water loss of males and females 38 of E. appendigaster and A. hagenowii with desiccation time (0 ‒24 h) at 24.0 ± 0.1°C and 0 ‒2% RH. Figure 3.3 Survival (ST 50 , d) of E. appendigaster males (A) and 41 females (B) when kept at 38%, 56%, 76%, and 87% RH in the absence and presence of food (10% sugar solution). Different letters and asterisks above bars denote significant differences in survival of parasitoids between different RHs in each food condition and between different food conditions in each RH, respectively. The vertical line on each bar represents the ± 95% fiducial limit. Numbers inside bars indicate sample size. Figure 3.4 Survival (ST 50 , d) of A. hagenowii males (A) and females 43 (B) when kept at 38%, 56%, 76%, and 87% RH in the absence and presence of food (10% sugar solution). Different letters and asterisks above bars denote significant differences in survival of parasitoids between different RHs in each food condition and between different food conditions in each RH, respectively. The vertical line on each bar represents the ± 95% fiducial limit. Numbers inside bars indicate sample size. Figure 3.5 Mean % distribution of E. appendigaster males (A) and 44 females (B) in linearly connected chambers with a gradient of 44%, 56%, 76%, and 87% RH. Different letters above each bar indicate significant differences in the distribution of parasitoids among RH chambers at each time interval (LSD test, P < 0.05). The vertical line on each bar represents standard error of the mean (n = 8). Figure 3.6 Mean % distribution of A. hagenowii males (A) and 46 females (B) in linearly connected chambers with a gradient of 44%, 56%, 76%, and 87% RH. Different letters above each bar indicate significant differences in ix the distribution of parasitoids among RH chambers at each time interval (LSD test, P < 0.05). The vertical line on each bar represents standard error of the mean (n = 10). Figure 4.1 (A) P.
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