PRINCIPLES OF INSECT BEHAVIOUR

2nd Revised and Enlarged Edition

Glass-wing butterfly Greta sp. is overlooked and invisible against flowers and other vegetation due to its transparent wings with very few scales.

PRINCIPLES OF INSECT BEHAVIOUR

2nd Revised and Enlarged Edition

V.B. Awasthi M.Sc. Ph.D.

Former Associate Professor Department of Zoology

J.N.V. University, Jodhpur (India)

Published by:

Scientific Publishers (India) 5 A, New Pali Road, P.O. Box 91 Jodhpur 342 001 (India) E-mail: info@scientificpub.com Website: www.scientificpub.com

Branch Office Scientific Publishers (India) 4806/24, Ansari Road, Daryaganj New Delhi - 110 002 (India)

First Edition, 2001 Second Edition, 2013 © 2013, Awasthi, V.B. All rights reserved. No part of this publication or the information contained herein may be reproduced, adapted, abridged, translated, stored in a retrieval system, computer system, photographic or other systems or transmitted in any form or by any means, electronic, mechanical, by photocopying, recording or otherwise, without written prior permission from the authors and the publishers. Disclaimer: Whereas every effort has been made to avoid errors and omissions, this publication is being sold on the understanding that neither the author nor the publishers nor the printers would be liable in any manner to any person either for an error or for an omission in this publication, or for any action to be taken on the basis of this work. Any inadvertent discrepancy noted may be brought to the attention of the publishers, for rectifying it in future editions, if published. ISBN: 978-81-7233-825-1 eISBN: 978-93-86237-86-6 Printed in India

Preface (1st Edition)

Although there are several books on animal behaviour which mainly deals with vertebrate fauna but there are only a few books on insect behaviour. The information on insect behaviour is very vast, but is scattered in research papers, review articles and monographs. Hence, information from all the available sources has been collected and compiled. The principal aim of writing this book has been to provide in short, the relevant information on the basic principles of insect behaviour together with modern trends on this fascinating and advancing subject.

This book is useful to the students of Entomology and to the insect Ethologists engaged in research work because it provides basic as well as advanced information with an exhaustive but selected bibliography which deals with most of the major aspects of the subject. This book is equally useful to the students of Zoology who study insect behaviour along with animal behaviour as a part of their syllabus in most of the Indian Universities.

This book is also useful to the general readers who have some basic knowledge of Biology and have curiosity to know why the insects are vividly coloured, have peculiar shapes, produce sound and light, and how they communicate.

It may be pointed out here that in each act of insect behaviour numerous well known external, internal as well as some little known factors are involved at times, because nature visualizes each pattern of behaviour in totality and not in individuality, hence it was very difficult to single out which one factor is more important for a particular behaviour and one is less. Some factor(s) may be significant or more pronounced in a particular pattern of behaviour, while they may also be of equal importance for controlling other pattern(s) of behaviour. Hence, the readers may find some sort of overlapping here and there in the text. It may be added that for a proper understanding of a particular behaviour pattern which is

influenced by more than one factor such sort of overlapping was unavoidable.

It is hoped that this book will be useful not only to the students of Entomology-Zoology and research scholars of insect behaviour but also to naturalists who wish to know about some of the mysteries of these small creatures and their life style. For the sake of convenience the book has been divided into 20 chapters viz —

(1) Kinds of insect behaviour (2) Camouflage, warning colouration and defensive behaviour (3) Social behaviour and parental care (4) The language of bees and their behaviour (5) Interactions between insects and plants as related to pollination (6) Food and feeding behaviour (7) Intraspecific and interspecific interactions (8) Adaptations in insects (9) Thermoregulation in desert insects (10) Photoperiodism and insect behaviour (11) Photoperiodism and ecological adaptations (12) Hormonal control of insect behaviour (13) Pheromonal control of insect behaviour (14) Genetic control of behaviour and evolution of behaviour (15) Communication in insects (16) Insect pest management by behavioural means (17) Migratory and dispersal behaviour of some insects (18) Influence of host plants on sex pheromone behaviour of phytophagous insects (19) The role of sense organs in mating behaviour of cockroaches and (20) Reproductive behaviour.

I would like to mention that a large number of insect Etholo-gists have helped me directly or indirectly in this endeavour but it is not possible to acknowledge them individually. I express my most humble and profound regards and sincere gratitude to all of them.

Thanks are also due to the authorities of the Scientific Publishers, Jodhpur, Shri Prahlad Rai Sharma and Shri Pawan Kumar Sharma for their encouragement, and keen interest in my work and bringing out the book in an excellent form.

I am also grateful to my friend Shri A.P. Jain, Botanical Survey of India, Jodhpur, for carefully going through the proof and offering valuable suggestions.

Finally, I also wish to thank my family members : Wife Sneh Lata, Son Santosh and daughter Shubha for their whole hearted co-operation without which this work was not possible.

V.B. Awasthi

30th May, 2000 Jodhpur

Preface to the Second Edition

With great pleasure I am presenting the Second edition of my book "Principles of Insect Behaviour". In this edition, a lot of new information has been added in the text at appropriate places alongwith a good number of illustrations. Two new chapters viz. 1. Ecological, behavioural and other adaptations of desert insects and 2. Thermoregulation by behavioural means have been included. Further, three appendixes have also been given dealing with 1. Amazing Insects, 2. Bee facts and 3. A copy of letter of Prof. V.B. Wigglesworth - The Father of Insect Physiology, in response to one of my critical letter which shows his magnanimity. I take this opportunity to express my gratitude to Prof. Wigglesworth for his useful suggestions received from time to time.

I hope the readers will find the book interesting as well as useful and welcome this edition with a great zeal.

V.B. Awasthi

28th November, 2012 Jodhpur

Contents

1 Kinds of Insect Behaviour 1

2 Camouflage, Warning Colouration and Defensive Behaviour 11

3 Social Behaviour and Parental Care 31

4 The Language of Bees and Their Behaviour 43

5 Interactions between Insects and Plants as Related to Pollination

48

6 Food and Feeding Behaviour 61

7 Intraspecific and Interspecific Interactions 76

8 Adaptations in Insects 86

9 Thermoregulation in Desert Insects 98

10 Ecological, Behavioural and other Adaptations of Desert Insects

102

11 Thermoregulation by Behavioural Means 115

12 Photoperiodism and Insect Behaviour 127

13 Photoperiodism and Ecological Adaptations 135

14 Hormonal Control of Insect Behaviour 139

15 Pheromonal Control of Insect Behaviour 147

16 Genetic Control of Behaviour and Evolution of Behaviour 163

17 Communication in Insects 171

18 Insect Pest Management by Behavioural Means 182

19 Migratory and Dispersal Behaviour of Some Insects 190

20 Influence of Host Plants on Sex Pheromone Behaviour of Phytophagous Insects

206

21 The Role of the Sense Organs in Mating Behaviour of Cockroaches

215

22 Reproductive Behaviour 223

Selected Bibliography 245

Glossary of Selected Technical Terms 268

Appendix - 1. Amazing Insects 279

Appendix - 2. Bee Facts 280

Appendix - 3. Letter of Prof. V.B. Wigglesworth 281

Subject Index 282

Acknowledgement and Figure Sources 300