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Studies on the Biology and Ecology STUDIES ON THE BIOLOGY AND ECOLOGY OF THE CABBAGE MOTH, MAMESTRA BRASSICAE L. (LEPIDOPTERA : NOCTUIDAE) by AQUILES MONTAGNE Ingeniero Agronomo (Venezuela) A thesis submitted for the degree of Doctor of Philosophy of the University of London and the Diploma of Imperial College Department of Zoology and Applied Entomology, Imperial College Field Station, Silwood Park, Ascot Berkshire September 1977 TABLE OF CONTENT Page ABSTRACT ' GENERAL INTRODUCTION 2 SECTION 1 THE BIOLOGY OF M. brassicae 5 1.1 Introduction 5 1.2 General Description of the Stages 6 1.3 Life History and Habits 8 1.4 Laboratory Studies on the Effect of Temperature on the Development and Survival of the Immature Stages 25 1.4.1 Materials and methods 25 1.4.2 Results and discussion 26 1.5 Laboratory Studies on Longevity and Fecundity 36 1.5.1 Materials and methods 36 1.5.2 Results and discussion 37 1.5.2.1 Longevity 37 1.5.2.2 Fecundity and Fertility 38 ' 1.6 Section General Discussion 47 SECTION 2 STUDIES ON THE EFFECTS OF LARVAL DENSITY ON M. 50 brassicae 2.1 Introduction 50 2.2 Review of Literature 50 2.3 Material and Methods 59 ii Table of Contents (Continued) Page 2.4 Results and Discussion 60 2.4.1 Colour variations in larval stage 60 2.4.1.1 Larval colour types 60 2.4.1.2 The colour of larvae reared at various densities 62 2.4.2 The pattern of larval and pupal development 66 2.4.2.1 The duration of larval development 66 2.4.2.2 The pattern of larval growth 71 2.4.2.3 The duration of prepupal and pupal periods 72 2.4.2.4 Larval and pupal mortality 75 2.4.2.5 Sex ratio 75 2.4.2.6 Diapause incidence in the pupae 75 2.4.3 The behaviour pattern of larvae 75 2.4.3.1 Introduction 75 2.4.3.2 General activity level 78 a)Materials and methods 78 b)Results and discussion 78 2.4.3.3 Response to physical disturbance 80 a)Materials and methods 80 b)Results and discussion 81 2.4.4 The resistance of the final instar larvae to stress factors 83 2.4.4.1 Introduction 83 lil Table of Contents (Continued) Page 2.4.4.2 Resistance to starvation 84 a) Materials and methods 84 b) Results and discussion 84 2.4.4.3 Resistance to desiccation 86 a) Materials and methods 86 b) Results and discussion 86 2.4.4.4 Resistance to DDT 89 a) Materials and methods 89 b) Results and discussion 89 2.4.5 The effect of density during hatching and first instar larvae 91 2.4.5.1 Introduction 91 2.4.5.2 The synchronization of larval hatching 92 a) Materials and methods 92 b) Results and discussion 92 2.4.5.3 The pattern of larval growth during the first instar 96 a) Materials and methods 96 b) Results and discussion 98 2.4.5.4 Influence of leaf toughness on the initiation of feeding by newly-hatched larvae 99 a) Materials and methods 99 b) Results and discussion 102 2.4.5.5 Larval mortality 104 2.5 Section General Discussion 111 iv Page SECTION 3 FIELD STUDIES ON THE MORTALITY OF EGG AND LARVAL STAGES AND DISPERSAL OF FULLY—GROWN LARVAE 116 3.1 Introduction 116 3.2 Studies on Egg Mortality 117 3.2.1 Materials and methods 117 3.2.2 Results and discussion 117 3.2.2.1 Characteristics of the egg mass infestations 117 3.2.2.2 Frequency of egg mass losses in relation to egg mass size 119 3.2.2.3 The effect of date of infestation on rate of loss in egg masses 120 3.2.2.4 The frequency of egg mass losses in relation to their position on the leaf 120 3.2.2.5 Egg mortality factors 121 a)Parasitism 125 b)Predation 127 c)Dislodgement 130 d)Disappearance 131 e)Other causes of mortality 132 3.3 Studies on Larval Mortality 132 3.3.1 Materials and methods 132 3.3.1.1 The plot 132 3.3.1.2 The exclusion devices 133 3.3.1.3 The setting up of the field experiments 134 3.3.1.4 Associated laboratory studies 138 a)Precipitin test 138 b)Feeding tests 138 V Page c) Rearing of field collected material 139 3.3.1.5 Details of the 1975 and 1976 field experiments 140 3.3.1.6 Meteorological data 140 3.3.2 Results and discussion 141 3.3.2.1 Survivorship curves and life tables 141 3.3.2.2 Mortality factors 157 a)Predators 157 b)Parasites 172 c)Diseases 174 d)Weather 175 3.4 Studies on the Dispersal of Fully—grown Larvae 176 3.4.1 Introduction 176 3.4.2 Materials and methods 177 3.4.2.1 Tagging release and recovery 177 3.4.2.2 Effect of tagging method on the insect 178 3.4.3 Results and discussion 179 3.4.3.1 Effect of tagging on the insect 179 3.4.3.2 Tag recovery 181 3.4.3.3 Larval dispersal and mortality 182 3.5 Section General Discussion 186 GENERAL DISCUSSION 192 SUMMARY 194 ACKNOWLEDGEMENTS 198 REFERENCES 200 APPENDICES 228 ABSTRACT Mdmestra brassicae L. (Lepidoptera: Noctuidae) is mainly a pest of brassicas. Experiments with this species were conducted both in constant temperature rooms and in the field. In the laboratory, the effects of different constant temperatures on the rate of development and mortalities of eggs, larvae and pupae were investigated;, at 20°C, fecundity, fertility and longevity of adults were also measured. The effects of larval rearing density on the behaviour of larvae and morphological and morphometrics characteristics of larvae and pupae were examined. A comparison was made of the abilities of individually-reared larvae and larvae reared in groups to survive starvation and dessication, and treatment with the insecticide DDT. Both the synchrony of eggs within an egg mass and the effects of grouping during the first instar larvae were studied. Using artificial infestations, the mortality factors acting on the egg and larval stages were studied in the field. The study of larval mortality factors was aided by caging to exclude ground predator, parasite, birds and rainfall; additional observations were made using the natural uncaged population. Observations were made on the distribution of egg masses and different larval instars on the host plant, and the egg masses distribution over the leaf surface. The dispersal and mortality of the sixth instar larvae and their pupation sites were studied using radioactive tags (1r192). 2. GENERAL INTRODUCTION The cabbage moth, Mamestra brassicae L. is mainly a pest on cabbages and although it has been recorded from a number of different plant species belonging to several families, it prefers cabbage and it is consistently reported as a serious pest on it more than on any other. M. brassicae is distributed almost completely throughout the Paleartic Region, except in the far north, from Western Europe to the coasts of the Pacific ocean reaching Japan. In the south it reaches the coasts and islands of the Mediterranean sea, the subtropical zone of Asia Minor, and the mountain districts of north-western India and southern China. The adaptability of M. brassicae to different climatic conditions and its recognised status as a major pest in many temperate and subtropical regions makes the study of M. brassicae important from the economic as well as from the biological point of view. Therefore, it is not surprising to find that in a survey of the literature many papers deal with this insect. However, most of them are concerned with brief accounts of obser- vations on the bionomics including biology, life history, seasonal variation, geographical distribution, host-plants or just its occurrence. A few deal with some aspects of its ecology and population dynamics, and in this respect Japanese workers have contributed the most. Many of the published papers on Mamestra are total or partially devouted to its control, either chemical or by other means e.g., parasite, disease, cultural measures. This seems to be justified because of the relatively high value of the crops which it attacks. 3. Along with a number of other species, the larvae of M. brassicae progressively change colour from pale to dark with increasing population density, and at the same time, other changes occur in its behaviour and physiology. This type of continuous polymorphism has been called "Phase variation" because it is very similar in many respects to that in locusts (Iwao, 1968). This and the frequent reports of mass outbreaks cited in the literature make "Phase variation" an important aspect in the study of the biology of this insect. Although M. brassicae is considered a major pest in eastern Europe, U.S.S.R., Japan and other places, in England it is a minor pest. Attacks are most common in gardens and allotments occurring occasionally on field scale; which perhaps accounts for the relatively little attention given to it by British entomologists. In this respect, it is then, inter- esting to obtain information on the mortality factors that operate under field conditions in this country as the findings may be relevant to other situations. The aims of the present study were to investigate some aspects of the biology and ecology of the cabbage moth and can be summarised as follows: a) To give a general description of the life history and habits of M. brassicae; to study in some detail the effect of tem- perature on the pre-imaginal stages; and the fecundity and longevity of the adults.
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