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Pentatomoidea and Their Egg—Parasites

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Pentatomoidea and Their Egg—Parasites THE BIOLOGY OF SOME PENTATOMOIDEA AND THEIR EGG—PARASITES By Mie JAVAHERY A thesis presented for the Degree of Doctor of Philosophy in the Faculty of Science of the University of London. March, 1967. Imperial College Field Station, Silwood Park, Sunninghill, Ascot, Berkshire. 2 ABSTRACT This thesis is divided into two sections. The first concerns the bionomics, habitats, reproductive biology of six Pentatamoidea, Heteroptera. The second section deals with eight species of egg parasites (Scelionidae, Hymenoptera) and includes their taxonomy and biology. The environment in which the six species of Heteroptera, Aelia acuminata (L.), Eurygaster integriceps Put". Neottiglossa pusilla (G.), Palomena prasina (L.), Piezodorus lituratus (F.) and Picromerus htdens (L.) live is described in the first section. Their biology and ecology in southern England and that of E. integriceps in Iran were investigated and compared. The reproductive morphology of all these species is outlined and the effects of environmental factors, especially of the food and temperature on fecundity and longevity of the adults, was investigated. The effects of these factors on the rate of development of the immature stages was also studied. Particular attention was paid to reproduction in Eurygaster integriceps in view of its economical importance and its status as a serious pest of cereal crops in Iran. Changes in the sizes of populations of A. acuminata and P. lituratus were recorded at Silwood Park, Berkshire. The problem of 3 winter diapause, migration and their possible relationship was studied. In the second section, six newly discovered British species of Scelionidae from the genera Asolcus and Telenamus were investigated. The first key to the identification of these wasps and of the closely related non-British species based on "bio-morphological taxonomy" is constructed. A study was made of the immature stages, adults, and the reproductive morphology of all the species in relation to various hosts and temperatures. Aggressive behaviour, ability of the parasites to select suitable host eggs, oviposition, copulation, the rate of development and sex ratio of all species, are described. Life histories, resistance to low temperatures, interspecific interference and host specificity of these minute wasps were observed throughout three seasons. Percentage of parasitism of eggs of several Pentatomoidea by Asolcus and Telenomus species at Silwood and at Yateley, were calculated for 1965 and 1966. New methods of rearing both the shieldbugs and their scelionid egg parasites have been devised. A preliminary distribution maps of hosts and parasites from the study areas are given for Britain and Iran. 4 CONTENTS Page INTRODUCTION ••• ••• ••• ••• ••• ••• 10 DESCRIPTION OF AREA OF STUDY ••• ••• ••• ••41 ••111 17 1. Area in Silwood Park ••• ••• ••• ••• 2. Area near Silwood Park ••• ••• ••• ••• 12Z 3. Other areas of study ••• ••• ••• ••• 22 METHODS 41.0 4100 • Gli • ••• 5 ,•• ••• 26 1. Numerical estimation in the field ••• ••411 26 2. Field rearing of the shieldbugs.. •4.• ••• 29 3. Laboratory rearing of shielJbugs. ••• ••• 35 4. Method of rearing of the larvae.. ••• ••• 46 SECTION 1. BIOLOGY OF SOME PENTATOMOIDEA STUDIES OF THE OVERWINTERING PERIOD IN PENTATOMOIDEA. • ••• 49 1. Biotic factors ... 0041 060 O.. ••• ••• 53 2. Climatic factors... • • • 0.0 O.. ••• ••• 55 3. Other climatic factors • • • 600 S OO •• 0 ••• 62 Aelia acuminata (Linnaeus) 1. The habitat 0•0 ••• • 00 ••• ••• 65 2. The host plants ••• ••• ••• ••• ••• 66 3. Description of stages • • • ••• ••• • • 0 • • • 68 4. Occurrence of Adults ... • • • • • • 68 5. The Number of Adults and Larval Stages of A. acuminata • •• ••• ••• • • • • • • 69 6. Reproductive Biology • 00 ••• ••• • • • • • • T1 7. Sexual Maturation.. ••• ••• ••• • • • • • • 71 8. Fecundity in Captivity and in the Field. • • • • • • 77 9: Rearing of Aelia... 000 • • • • • • • • • • • • 81 10: Aggregation. 000 • • • 41410 • • • • • • 81 11. Migration, Aestivation and Hibernation.. • • 0 • • • 82 12. Sex Ratio... ••• 41•11 6.0 000 • • • • • • 82 Eurygaster integriceps Puton 000 660 ••• 0 • • ••• 83 1. Life History 060 41614 ••• ••• 85 2. Reproductive Biology 0.0 SOO' ••• ••• 86 3. Sexual Maturation.. SOO O.. 000 0 •• ••• 86 3.1. Diapause ..* 000 OW& ••• ••• ••• 86 5 Page 3.2. Conclusion ••• • • • 93 4. Copulation in Eurygaster integriceps. • • • • • • 96 5. FECUNDITY OF E. integriceps IN RELATION TO FOOD AND TEMPERATURE IN THE LABORATORY ••• 101 A) FECUNDITY OF E. inte,,riceps ON WHEAT ... ••• 102 Conclusions 600 *Of IWO* 040 • • • 107 B) FECUNDITY OF E. integriceps ON BARLEY... 107 Conclusions ••• 0444, ••• ••• • • • 112 c) The Effect of a Single Copulation on Fecundity of E. integriceps ••• *Of 112 D) Relationship between Crowding and Fecundity 0 • 0 114 Neottiglossa pusilla (Gmelin) 1. The habitat... C • • •O• ••0 0•• 118 2. The host plants • • • 000 6•0 ••• ••• 113 3. Description of Stages. ••• ••• ••• ••• 119 4. Occurrence of Adults.. ••• ••• 0•• ••• 119 5. ReproductiveBialoGy.. • 4.• ••• ••• ••• 119 6. Sexual Maturation ... 11•0 ••• •O• ••• 119 7. Fecundity in Captivity and in the Field ... ..• 121 8. Rearing of Larval Instars of N. pusilla • .• •• • 122 9. Migration.. ••• • • . • • • •• • 122 Palomena prasina (Linnaeus) 1. The habitat ... •• • • •• •• • ••• o•• 123 2. The host plants ••• ••• ••• ••• ••• 123 3. Description of stages. ... ... *00 OS* 124 4. The Number of Adults and Larval Stages of P. prasina 124 5. Reproductive Biology.. 000 000 *WO 000 125 6. Sexual Maturation ... 00* 060 *06 *60 125 7. Fecundity in Captivity and in the Field 006 600 126 8. Development of the Larval Instars and their Food.. 127 Piezodorus lituratus (Fabricius) 1. The habitat ••• ••• 0 • • GOO 128 2. The host plants • 0 • • • • 128 3. Description of Stages. ••• ••• ••• 129 4. Time of occurrence of Adults. 000 *0* 129 5. Estimation of Numbers of Adults and Larvae of P. lituratus during the three seasons... 131 6. Biometrical Differences in Adults and larval instars of P. lituratus • 0 • ••• 134 7. Reproductive Biology.. ••• ••• ••• 134 8. Sexual Maturation ... 000 000 000 • • • 137 9. Fecundity in the Field and in the Laboratory 139 6 Page Picromerus bidens (Linnaeus) 1. The habitat ... • • • • • • • • • • .• ••• 147 2. Food 0.0 ••• ••• ••• ••• ••• ••• 147 3. Description of Stages.. • • • •• • •• ••• 151 4. The time of Occurrence of Adults and their Larvae... 151 5. The Number of Adults and their Larval Instars *0* 152 6. Reproductive Biology... • • • • • • • • • ••• 152 7. Sexual Maturation ... ... • • • • . • •• • 154 8. Fecundity and Longevity of P. bidens.. *00 0.41 157 Fecundity in the Field ... • • • • • • •• . 157 B1 Fecundity in the Laboratory. SOO • • • 00. 161 9. Rearing of P. bidens... • • • • • • • • . •• • 168 10. Number of Annual Generations... 0.0 • • 0 • • • 169 AGGREGATION IN PENTATOMOIDEA.. • • • •• • • • • .• • 170 MIGRATION BEHAVIOUR IN PENTATOMOIDEA.. 001 0.4 ... 171 Conclusions on migration • • • • . ... •• • 174 EFFECT OF SOME PARASITES AND PREDATORS ON PENTATOMOIDEA IN SOUTHERN ENGLAND •• • ••• • •• • • . • • • ••• 175 SECTION 2. BIOLOGY AND TAXONOMY OF SOME PROOTOTRUPOIDEA INTRODUCTION ... •• • •• • • •• •• • • • • •• • 178 REVIEW OF LITERATURE ... 000 0.0 ... 000 .00 181 MATERIALS, METHODS AND TECHNIQUES • •• •• • ••• ••• 188 A) Material for Investigations 00. 000 000 0.* 188 B) Eggs of Pentatomoidea... ... • • • ... •• • 189 METHODS OF STUDYING THE PARASITES ... • • • 000 0.0 192 Method of Collecting the egg parasites in the field. 192 The Method of placing egg:batches in the field ... 193 Collecting the naturally parasitised egg batches ... 195 Method of finding the adult egg parasites ... 0.0 195 DISSECTION, PREPARATION, PRESERVATION AND DRAWING... ... 1. Dissection ... ... ... • • • • • • ... 198 2. Preparation of specimens • • • • • • ••• •• • 20(' 3. Preserving of the specimens • • • • • • •• • ••• 20',- 4. Drawing... •• • • •• • • • • .. — • •• • 202 TECHNIQUES OF REARING Asolcus AND Telenomus spp. 000 .** REARING OF PARASITES IN THE LABORATORY ... 010 000 204 FIELD REARING CAGES.. 00. • • • • • • 000 *60 214 7 Page THE TAXONOMY OF EGG PARASITES COMPLEX OF BRITISH PENTATOMOIDEA. • 0 • 0 • • • • • •• 04.0 217 I Genus Asolcus NAKAGAWA. 0•• ••• ••• ••• 219 Key to species.. • • .,•• •. • • • • 4o. 219 Asolcus waloffae sp.n.. .• • •• • •• • •,• 225 Asolcus davatchii sp.n. •• . .• • .• • •• • 231 Asolcus silwoodensis sp.n. ... ... •• • ... 238 Asolcus nixo—martini sp.n. •• . •• • •• • •• . 247 II Genus Telenomus Haliday ... ... ... ... 252 Telenomus truncatus Mayr ... 004 000 0*, 254 Telenomus sokolovi Mayr 261 Conclusions ... • • • •• • •• • ••4. ••• 266 THE BIOLOGY AND ECOLOGY OF EGG PARASITES COMPLEX OF BRITISH PENTATOMOIDEA. ... ... ... *66 4410 268 Asolcus waloffae sij,,n. Morphology of the Immature Stages.... ese • • • 268 Reproductive Organs ... ... ... ... ... 270 Mating... • • • • • • .•. • • • •• • •• • 272 Oviposition ... • • • •• • ••• ••• ••• 276 Development ... • • • ••• •• • •• • •• • 285 The Emergence of the Adult Parasites. 000 00* 289 Occurrence in the Field ... 411O4 000 OlDe 291 Number of Annual Generations.. ... •.• 41111 292 •• • ••• Fecundity • .• • • • • • • .•• 293 Parthenogenesis. ... ... •. • 000 • 00 298 Sex Ratio • •• • • . •• • •• • •• • •• • 298 Host Specificity • • • •• • ••• •• • •• • 300 Distribution ... • • • . • • •• • •.. •• . 300 Asolcus davatchii sp.n. Morphology of Immature Stages. ... ... ... 301 Reproductive Organs ... • • • 440 ... ... 304 14641 Mating... ... ... ... 040 .O0 305 Oviposition
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