Courtship and mating in butterflies

394 Pages·2020·14.183 MB·English

by  Cannon, Raymond Julian Conway

C M B
ourtship and ating in utterflies

C M
ourtship and ating
B
in utterflies
raymond J.C. Cannon
CABI is a trading name of CAB International
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© Raymond J.C. Cannon 2020. All rights reserved. No part of this publication
may be reproduced in any form or by any means, electronically, mechanically, by
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A catalogue record for this book is available from the British Library, London, UK.
Library of Congress Cataloging-in-Publication Data
Names: Cannon, Raymond Julian Conway, author.
Title: Courtship and mating in butterflies : reproduction, mating
behaviour and sexual conflicts / Raymond Julian Conway Cannon.
Description: Boston, MA : CAB International, 2020. | Includes
bibliographical references and index. | Summary: “The aim of this book is to
present a readable account of butterfly behaviour, based on field observations,
great photographs and the latest research. The main focus is on courtship and
mating - including perching, searching and territorial behaviour - but to
understand these subjects it is necessary to explain how mates are chosen
and this requires sections on wing colours and patterns. A chapter on
butterfly vision is also essential in terms of how butterflies see the world and
each other. There have been exciting discoveries in all of these fields in
recent years, including: butterfly vision (butterfly photoreceptors), wing
patterns (molecular biology), wing colouration (structural colours and
nano-architecture), mating strategies and female choice (ecology and
behaviour)”-- Provided by publisher.
Identifiers: LCCN 2019026041 (print) | LCCN 2019026042 (ebook) | ISBN
9781789242638 (hardback) | ISBN 9781789242645 (ebook) | ISBN
9781789242652 (epub)
Subjects: LCSH: Butterflies--Behavior. | Courtship in animals.
Classification: LCC QL562.4 .C36 2020 (print) | LCC QL562.4 (ebook) | DDC
595.78/9156--dc23
LC record available at https://lccn.loc.gov/2019026041
LC ebook record available at https://lccn.loc.gov/2019026042
ISBN-13: 9781789242638 (hardback)
9781789242645 (ePDF)
9781789242652 (ePub)
Commissioning Editor: Ward Cooper
Editorial Assistant: Lauren Davies
Production Editor: Tim Kapp
Typeset by SPi, Pondicherry, India
Printed and bound in the UK by Severn, Gloucester
Contents
Preface xi
Acknowledgements xv
1. Introduction 1
Welcome to the Superfamily! 2
Origins and evolution 2
Butterfly Families 4
Names and numbers 5
Pioneering Investigations of Butterfly Courtship 6
Tinbergen and the grayling 7
Crane and ‘irrelevant actions’ in tropical heliconiids 10
Stride and the danaid eggfly 12
Magnus and the silver-washed fritillary 14
Brower and the queen butterfly 16
2. Sexual Selection and Mate Choice 19
Choosing a Mate – the Female Prerogative? 19
Sexual selection 20
Sexual conflicts 21
Good genes and handicaps 22
Identifying a Potential Mate 22
Visual signals 24
Female Choice 29
Eyespots, ornamentation and ultraviolet light 30
Active and passive choices 34
Iridescent ornaments and a preference for flashy males! 35
Females with a tendency to vary 38
Mr Normal – stabilizing selection 40
v
vi Contents
Male Choice 40
The banded peacock 44
Role Reversals 45
The alba polymorphism in Colias species 46
Sexual Dimorphism 48
Differences in size 48
Differences in shape 49
Differences in colours and patterns 50
Differences in ornamentation 55
Differences in smell and taste 56
3. Strategies for Locating a Mate 60
Perching and Patrolling 60
Optimal search strategies 62
Perching – Staying Put and Waiting for Females 62
Territories 63
Thermoregulation and site selection 64
Patrolling – Moving Around and Searching for Mates 65
Hill-topping – Sexual Rendezvous Points 68
Niche segregation 69
Leks – arenas for contests 70
Pushing and shoving! 71
Landmark mating systems 72
Examples of Perching and Patrolling by Family 72
Hesperiidae 72
Lycaenidae 73
Nymphalidae 73
Riodinidae 75
Papilionidae 75
Pieridae 77
Territoriality 78
Territory formation and turnover 80
Territorial contests 81
Battling swallowtails 83
Crepuscular contests 84
Wars of attrition or mistaken identity? 85
Persistence or endurance? 87
Older males – experienced and more persistent? 88
Case Study: The speckled wood 89
4. Seeing and Being Seen 96
Butterfly Colour Vision 96
The Structure of the Compound Eye 98
Eye Shine and Differences Between the Sexes 100
The Light-Gathering Molecules 101
Gathering the Light in Different Ways 104
Three-Colour and Four-Colour Vision 105
Contents vii
Seeing Ultraviolet and Polarized Light 107
Hindsight: Genital Photoreceptors 108
Pseudopupils 108
Butterflies Using Their Eyes 110
5. Courtship Behaviour 114
What is Courtship? 114
Courtship Patterns and Diversity 116
Tabulating butterfly behaviour 117
Lycaenids – Good Vibrations! 117
The blues 117
Three coppers 119
Hairstreaks 120
The apefly 122
Pierids – Wing Buffeting 123
Three little sulphurs 124
Japanese small white 126
A forest ‘ghost’ 129
Hesperiids 129
Hesperia skippers 129
The fiery skipper 130
Giant skippers 132
The Essex skipper 132
Nymphalids 133
The buckeye 134
Common and danaid eggflies – all of a quiver 135
Banded and scarlet peacocks 136
The squinting bushbrown, Bicyclus anynana 137
Heliconius butterflies 139
Fritillaries and checkerspot butterflies 141
Satyrids 144
Danaiids 145
Monarch butterflies: hell’s angels? 147
Other examples of forced matings and aerial
‘takedowns’ 149
Papilionids 151
Birdwing butterflies 152
Multimodal Displays – the Components
of Courtship 153
Determinants of Success or Failure 155
6. Body Language 160
Refusal and Rejection 160
Playing possum: thanatosis 165
Female Solicitation 167
Clinging on and Standing Guard 168
Pair clinging 168
Mate guarding 168
viii Contents
Antennal Contacts – Courtship Bowing 169
Common graylings 169
Rock and tree graylings 172
Other species 174
Proboscis Waving: Wood Whites 175
Sonic Courtship: Do You Like My Click?! 178
7. The Mating Game 181
Reproduction: A Battle of the Sexes, or a Tug of War? 181
Mating Strategies 183
Copulation 184
Pair carrying in copula 184
Copulation duration 185
Post-nuptial flights and mate-guarding 187
Sperm competition 188
Protandry and female receptivity 190
Sexual Conflicts and Confluences 191
Polyandry – the Risks and Benefits of Multiple Matings 192
Experience counts? Old male mating advantage 197
Cryptic female choice: rewarding generous suitors 198
Costs and benefits 199
Pupal Mating 201
Strange or Enigmatic Liaisons 203
Incest and inbreeding 203
Same-sex interactions 204
Interspecific mating and hybridization 205
8. The Inside Story 211
The Mechanics of Coupling 211
Tickle sticks! 213
Mating Plugs and Sphragides 213
Sperm and Spermatophores 218
The Female Side of the Story 224
9. Scents: Chemical Communication 227
Aphrodisiac Pheromones 228
Detectable odours 229
Studies using Bicyclus anynana 231
Heliconius butterflies 233
Androconia 233
Varieties of androconial structures 234
Sex brands in skippers (Hesperiidae) 236
Pierids 238
The squinting bushbrown 238
Abdominal Organs (‘Hairpencils’) 240
Contact behaviour and perfuming 243
Pyrrolizidine alkaloids (PAs) 244
Antiaphrodisiacs 245
Contents ix
10. Wing Colours and Iridescence 251
The Different Functions of Butterfly Wing Surfaces 251
Structural Colours and Nanoarchitecture 252
Iridescence 253
Black light-traps and solar collectors 254
Hydrophobicity and self-cleaning wings 255
Scales and Ridges 255
Pigments 258
Blue Morpho Wings 259
Lycaenid Wings 263
Partitioning the Function (Different Wings
for Different Things) 268
Pierid Wings 269
Colour Badges as Honest Signals 271
Papilionoid Wings 272
Birdwings 273
Mixing it 274
Nymphalid Wings 277
Clearwings and Nanopillars 279
Concluding Remarks 282
Summing Up 285
Glossary 289
Annex 1 – Internet Links 295
References 297
Index 357