DIPTERA: CALLIPHORIDAE), and the PHYSIOLOGICAL and BEHAVIOURAL CHANGES in ITS HOST, Macrotermes Carbonarius (BLATTODEA: TERMITIDAE)
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PREVALENCE, DISTRIBUTION AND THE BIOLOGY OF AN ENDOPARASITOID FLY, Verticia fasciventris (DIPTERA: CALLIPHORIDAE), AND THE PHYSIOLOGICAL AND BEHAVIOURAL CHANGES IN ITS HOST, Macrotermes carbonarius (BLATTODEA: TERMITIDAE) TO THI MAI DUYEN UNIVERSITI SAINS MALAYSIA 2018 PREVALENCE, DISTRIBUTION AND THE BIOLOGY OF AN ENDOPARASITOID FLY, Verticia fasciventris (DIPTERA: CALLIPHORIDAE), AND THE PHYSIOLOGICAL AND BEHAVIOURAL CHANGES IN ITS HOST, Macrotermes carbonarius (BLATTODEA: TERMITIDAE) by TO THI MAI DUYEN Thesis submitted in fulfilment of the requirements for the Degree of Master of Science April 2018 ACKNOWLEDGEMENT I would like to express my heartfelt thanks to my supervisor Prof. Dr. Lee Chow-Yang for his unwaverly support that has enabled me to complete my thesis. His mentorship, wisdom, enthusiasm, and kindness have inspired, motivated and guided me during my M.Sc. research in the world renowned Urban Entomology Lab (affectionately known as the Urban Lab). I also would like to thank all the past and present Urban Lab labmates, Dr. Lee Ching Chen, Dr. Tee Hui Siang, Dr. Foo Foong Kuan, and Dr. Neoh Kok-Boon for their valuable discussions on my research, and assistance in data collection in the field. I would like to thank my bench mate, Sophia Liew for her support and care during my stay in Penang. In addition, I also am indebted to other labmates namely Chris Kim Daeyun, Mark Ooi Chuan Sen, Jong Zheng Wei, Gan Li Yan, Leong Xin Yeng, Edmund Hang, Low Yue Jua, and Liu Kun Peng for their friendship, assistance and support. Lastly, my special thanks to my parents, sister, cousin and friends. This thesis would not have been possible without their endless support. ii TABLE OF CONTENTS ACKNOWLEDGEMENT ii TABLE OF CONTENTS iii LIST OF TABLES vi LIST OF FIGURES vii LIST OF PLATES viii ABSTRAK ix ABSTRACT xi CHAPTER ONE: GENERAL INTRODUCTION 1 CHAPTER TWO: LITERATURE REVIEW 4 2.1 Parasitism 4 2.2 Macrotermes carbonarius (Hagen) (Isoptera : Termitidae) 5 2.3 Family Calliphoridae 10 2.3.1 Eggs 10 2.3.2 Larvae 10 2.3.3 Pupae 11 2.3.4 Adults 11 2.3.5 Genus Verticia 11 2.4 Host selection 12 2.4.1 Environmental factors 13 2.4.2 Host factors 14 2.5 Parasitoid – induced physiological changes in the host 15 2.6 Parasitoid – induced behaviour changes in the host 16 CHAPTER THREE: PREVALENCE OF THE ENDOPARASITOID Verticia fasciventris IN COLONIES OF THE FUNGUS-GROWING TERMITE Macrotermes carbonarius ON PENANG ISLAND, MALAYSIA 19 3.1 Introduction 19 3.2 Materials and Methods 20 3.2.1 Survey sites 20 iii 3.2.2 Termite sampling 20 3.2.3 Degree of parasitism and status of colony health 22 3.2.4 Mound size and light intensity measurement 22 3.3.5. Observation of infected soldiers with no morphological changes 23 3.2.6 Statistical analysis 23 3.3 Results 23 3.3.1 Frequency of parasitism 23 3.3.2 Degree of parasitism and status of colony health 25 3.3.3 Mound size and light intensity of parasitized and unparasitized mounds 25 3.3.4 Observation of infected soldiers with no morphological changes 26 3.4 Discussion 26 CHAPTER FOUR: MORPHOLOGY AND DEVELOPMENT OF THE ENDOPARASITOID Verticia fasciventris IN ITS TERMITES HOST 30 4.1 Introduction 30 4.2 Material and Methods 31 4.2.1 Specimen collection and maintenance 31 4.2.2 Morphology of different stages of V. fasciventris 32 4.2.3 Development of different stages of V. fasciventris 33 4.2.4 Statistical analysis 35 4.3 Results 35 4.3.1 Morphology of different stages of V. fasciventris 35 4.3.2 Development of different stages of V. fasciventris 41 4.4 Discussion 44 CHAPTER FIVE: WATER BALANCE AND SURVIVAL OF PARASITIZED AND UNPARASITIZED SOLDIERS IN A Macrotermes carbonarius COLONY 47 5.1 Introduction 47 5.2 Material and Methods 48 5.2.1 Termite collection 48 iv 5.2.2 Water balance 48 5.2.3 Statistical analysis 50 5.3 Results 51 5.3.1 Water balance profile of parasitized and unparasitized major soldiers 51 5.3.2 Survival of parasitized and unparasitized major soldiers under 4% RH 54 5.4. Discussion 55 CHAPTER SIX: BEHAVIOURAL CHANGES IN Macrotermes carbonarius IN COLONIES PARASITIZED BY THE ENDOPARASITOID 58 6.1 Introduction 58 6.2 Materials and Methods 58 6.2.1 Termite collection 59 6.2.2 Agonistic behaviours of parasitized soldiers during foraging activities 60 6.2.3 Trophallaxis and allogrooming activities between workers and soldiers 61 6.2.4 Statistical analysis 62 6.3 Results 63 6.3.1 Agonistic behaviour of parasitized soldiers during foraging activity 63 6.3.2 Trophallaxis and allogrooming frequencies between workers and parasitized or unparasitized soldier 65 6.3.3 Behaviour of workers in groups with or without parasitized or unparasitized individuals 65 6.4 Discussion 69 SUMMARY AND CONCLUSION 74 REFERENCES 78 LIST OF PRESENTATIONS v LIST OF TABLES Page Table 3.1 Frequencies of parasitism at each surveillance site on Penang Island 24 Table 4.1 Measurements (mean ± SD, mm) and comparisons of morphological characteristics of three larval instars of V. fasciventris Table 4.2 Development duration (mean, days) of the stages of 43 V. fasciventris under laboratory conditions Table 5.1 Initial body weight, percentage of total body water, 53 cuticular permebility, and surface area to volume ratio (mean ± SD) of parasitized and unparasitized major soldiers of M. carbonarius Table 6.1 Results of observations of larvae from non-aggressive and 64 aggressive parasitized soldiers collected from foraging trails Table 6.2 Frequencies of trophallaxis and allogrooming between 66 workers and parasitized or unparasitized soldiers Table 6.3 Frequencies of trophallaxis and allogrooming between 67 workers and unparasitized soldiers in the presence or absence of parasitized soldiers Table 6.4 Frequencies of trophallaxis and allogrooming between 68 workers and parasitized soldiers in the presence or absence of unparasitized soldiers vi LIST OF FIGURES Page Figure 3.1 Surveyed sites on Penang Island, Malaysia 21 Figure 4.1 Canonical discriminant functions of larval instars of 39 V. fasciventris Figure 4.2 The life cycle of V. fasciventris 42 Figure 5.1 Cumulative % total body water loss of parasitized and 52 unparasitized major soldiers of M. carbonarius Figure 5.2 Survival rate of parasitized and unparasitized major 54 soldiers vii LIST OF PLATES Page Plate 2.1 Major soldiers of M. carbonarius 7 Plate 2.2 The newly formed parasitized soldier derived from a pre- soldier of M. carbonarius 8 Plate 2.3 A parasitized pre-soldier of M. carbonarius 9 Plate 4.1 Photos of larva stages of V. fasciventris 36 Plate 4.2 V. fasciventris pupa 39 Plate 4.3. V. fasciventris adult 40 viii PREVALENS, TABURAN, DAN BIOLOGI SUATU LALAT ENDOPARASITOID, Verticia fasciventris (DIPTERA: CALLIPHORIDAE), PERUBAHAN FISIOLOGI DAN KELAKUAN PADA PERUMAHNYA Macrotermes carbonarius (BLATTODEA: TERMITIDAE) ABSTRAK Tesis ini menumpu ke atas prevalans, distribusi dan biologi lalat endoparasitoid, Verticia fasciventris Malloch (Diptera: Calliphoridae), perubahan fisiologi serta tingkah laku dalam rumahnya Macrotermes carbonarius (Hagen) (Blattodae: Termitidae). Perkembangan larva parasitoid V. fasciventris boleh dibahagikan kepada tiga peringkat instar. Larva menamatkan instar pertama mereka dalam kapsul kepala pra-askar, manakala larva instar kedua berkembang dalam kapsul kepala askar. Larva instar ketiga pula berkembang dalam kapsul kepala askar dan kemudiannya hidup bebas setelah keluar dari badan perumah anai-anai. Purata jangka masa perkembangan untuk peringkat pupa adalah 13 hari. Saiz parasitoid betina adalah lebih kecil (P < 0.05) berbanding dengan saiz parasitoid jantan. Kemandirian parasitoid betina adalah lebih tinggi (P < 0.05) berbanding dengan kemandirian parasitoid jantan. Tinjauan kadar parasitisme oleh endoparasitoid V. fasciventris pada koloni M. carbonarius di Pulau Pinang, Malaysia dijalankan dari April 2015 hingga April 2016. Daripada 313 koloni M. carbonarius yang telah ditinjau, 105 (33.5%) telah diparasitkan oleh V. fastrivencis. Di kesemua tempat kajian, koloni yang diparasit masih lagi sihat. Kadar parasitisme adalah rendah dengan purata 42.7 ± 12.9 askar M. carbonarius. Saiz busut koloni yang diparasit adalah jauh lebih besar daripada yang tidak berparasit. Keamatan cahaya persekitaran busut koloni yang diparasit adalah jauh lebih rendah (P < 0.05) daripada busut koloni yang tidak diparasit. Kehadiran V. ix fasciventris akan menyebabkan perubahan fisiologi dalam M. carbonarius. Profil kandungan air untuk askar major yang diparasit dan tidak berparasit telah diselidik dalam kajian ini. Askar major yang diparasit menunjukkan jisim badan awal, peratusan jumlah kandungan air badan serta kebolehtelapan kutikel yang jauh lebih rendah (P < 0.05), tetapi mempunyai nisbah permukaan kepada isipadu badan yang lebih tinggi berbanding dengan askar major yang tidak diparasit. Askar major yang diparasit mempunyai kadar hidup yang jauh lebih tinggi (P < 0.05) berbanding dengan askar major tidak diparasit pada kelembapan relatif 4%. Askar M. carbonarius major yang diparasit masih mengekalkan peranan melindungi anai-anai pekerja yang mencari makanan. Askar major yang diparasit boleh dikelaskan kepada kumpulan yang agresif dan yang kurang agresif. Purata masa larva keluar dari perumah yang kurang agresif adalah lebih singkat berbanding dengan purata masa larva untuk keluar dari perumah agresif di lapangan. Kekerapan trofalaksis dan pembersihan alo (‘allogrooming’) pekerja terhadap askar major yang diparasit adalah lebih tinggi berbanding dengan kekerapan kedua faktor tersebut untuk mereka yang tidak diparasit. Kehadiran askar major yang diparasit mempengaruhi kekerapan trofalaksis dan pembersihan alo oleh anai-anai pekerja, manakala kehadiran askar major yang tidak berparasit tidak mempengaruhi kekerapan aktiviti tersebut (P > 0.05). x PREVALENCE, DISTRIBUTION AND THE BIOLOGY OF AN ENDOPARASITOID FLY, Verticia fasciventris (DIPTERA: CALLIPHORIDAE), AND THE PHYSIOLOGICAL AND BEHAVIOURAL CHANGES IN ITS HOST, Macrotermes carbonarius (BLATTODEA: TERMITIDAE) ABSTRACT This thesis focuses on prevalence, distribution and the biology of an endoparasitoid fly, Verticia fasciventris Malloch (Diptera: Calliphoridae) and the physiological and behavioural changes in its host Macrotermes carbonarius (Hagen) (Blattodae: Termitidae).