Mimicry, Crypsis, Masquerade and Other Adaptive Resemblances
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MIMICRY, CRYPSIS, MASQUERADE AND OTHER ADAPTIVE RESEMBLANCES MIMICRY, CRYPSIS, MASQUERADE AND OTHER ADAPTIVE RESEMBLANCES Donald L. J. Quicke FRES, PhD Professor, Chulalongkorn University, Bangkok, Thailand This edition first published 2017 © 2017 John Wiley & Sons Ltd All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by law. Advice on how to obtain permission to reuse material from this title is available at http://www.wiley.com/go/permissions. The right of Donald L. J. Quicke to be identified as the author of this work has been asserted in accordance with law. 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Neither the publisher nor authors shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages. A catalogue record for this book is available from the Library of Congress and the British Library. ISBN 9781118931530 Cover images: courtesy of Donald L.J. Quicke (except top left on back cover, courtesy of Linda Pitkin) Set in 9/11pt Photina by SPi Global, Pondicherry, India 10 9 8 7 6 5 4 3 2 1 For all the dogs who live on streets everywhere with no‐one to love them, and to those wonderful organisations such as Soi Dog Foundation in Thailand, that work hard to care for them. This is wonderful Puii. She lives at Saphan Taksin Pier. CONTENTS Preface, xiii Butterfly pupal colour polymorphism, 32 A comment on statistics, xv Winter pelage: pelts and plumage, 35 A comment on scientific names, xvi Melanism, 37 Industrial melanism, 37 Acknowledgements, xvii Fire melanism, 40 Background selection, 41 Orientation and positioning, 43 1 INTRODUCTION AND CLASSIFICATION Transparency, 45 OF MIMICRY SYSTEMS, 1 Reflectance and silvering, 47 A brief history, 2 Adaptive colour change, 49 On definitions of ‘mimicry’ and adaptive resemblance, 3 Caterpillars and food plant colouration, 50 The concept of ‘adaptive resemblance’, 8 Daily and medium‐paced changes, 54 The classification of mimicry systems, 9 Rapid colour change, 56 Wickler’s system, 9 Chameleons, 56 Vane‐Wright’s system, 10 Cephalopod chromatophores Georges Pasteur (1930–2015), 11 and dermal papillae, 57 Other approaches, 13 Bird eggs and their backgrounds, 58 Endler, 13 Disguising your eyes, 61 Zabka & Tembrock, 13 Disruptive and distractive markings, 61 Maran, 14 Edge‐intercepting patches, 61 Mimicry as demonstration of evolution, 14 Distractive markings, 63 Zebra stripes and tsetse flies, 66 Stripes and motion dazzle – more zebras, kraits 2 CAMOUFLAGE: CRYPSIS AND DISRUPTIVE and tigers, 69 COLOURATION IN ANIMALS, 19 Computer graphics experiments with human subjects, 69 Introduction, 20 Observations on real animals, 69 Distinguishing crypsis from masquerade, 20 Comparative analysis, 71 Crypsis examples, 24 Dual signals, 72 Countershading, 24 Protective crypsis in non‐visual modalities, 73 Experimental tests of concealment by Apostatic and antiapostatic selection, 73 countershading, 27 Search images, 74 Bioluminescent counter‐illumination, 28 Experimental tests of search image, 76 Background matching, 29 Gestalt perception, 76 Visual sensitivity of predators, 30 Effect of cryptic prey variability, 77 To make a perfect match or compromise, 31 Reflexive selection and aspect diversity, 77 Colour polymorphism, 32 Searching for cryptic prey – mathematical Seasonal colour polymorphism, 32 models, 80 vii viii Contents Ontogenetic changes and crypsis, 81 Behaviour of protected aposematic animals, 135 Hiding the evidence, 82 Of birds and butterflies, 135 Petiole clipping by caterpillars, 82 Evolution of sluggishness, 139 Exogenous crypsis, 82 Origins of protective compounds, 140 Military camouflage and masquerade, 85 Plant‐derived toxins, 140 Cardiac glycosides, 141 Pyrrolizidine alkaloids, 144 3 CAMOUFLAGE: MASQUERADE, 87 De novo synthesis of protective compounds, 145 Introduction, 88 Obtaining toxins from animal sources, 147 Classic examples, 88 Costs of chemical defence, 149 Twigs as models, 88 Aposematism with non‐chemical defence, 150 Leaves (alive or dead) as models, 88 Escape speed and low profitability, 150 Bird dropping resemblances, 89 Parasitoids and aposematic insects, 152 Spider web stabilimenta, 93 Diversity of aposematic forms, 152 Tubeworms, etc., 94 Egg load assessment, 154 Experimental tests of survival value of masquerade, 94 Proof of aposematism, 154 Ontogenetic changes and masquerade, 97 Bioluminescence as a warning signal, 155 Thanatosis (death feigning), 97 Warning sounds, 155 Feign or flee? The trade‐offs of thanatosis, 100 Warning colouration in mammals, 157 Other aspects of death mimicry, 100 Weapon advertisement, 158 Seedless seeds and seedless fruit, 100 Mutualistic aposematism, 160 Aposematism induced by a parasite, 161 4 APOSEMATISM AND ITS EVOLUTION, 103 Aposematic commensalism, 161 Polymorphism and geographic variation in Introduction, 104 aposematic species, 161 Initial evolution of aposematism, 108 Aposematism in plants, 163 Associations of unpalatable experience Synergistic selection of unpalatability with place, 109 in plants, 165 Mathematical models and ideas of warning colouration Aposematism in fungi, 166 evolution, 112 Why are some unpalatable organisms aposematic Kin selection models, 112 and others not?, 167 Green beard selection, 112 Family selection models, 113 5 ANTI‐PREDATOR MIMICRY. I. Individual selection models, 113 MATHEMATICAL MODELS, 171 Spatial models and metapopulations, 116 Handicap and signal honesty, 117 Introduction, 172 Early warnings – reflex bleeding, vomiting Properties of models, rewards, learning rates and and other noxious secretions, 120 numerical relationships, 172 Longevity of aposematic protected taxa, 121 Simple models and their limitations, 173 Macroevolutionary consequences, 121 Müller’s original model, 173 Experimental studies, 121 Simple models of Batesian and Tough aposematic prey and individual selection, 121 Müllerian mimicry, 173 Pyrazine and other early warnings, 123 Are Batesian and Müllerian mimicry Learning and memorability, 124 different?, 174 Strength of obnoxiousness, 126 An information theory model, 176 Is the nature of the protective compound Monte‐Carlo simulations, 177 important?, 126 More refined models – time, learning, forgetting Neophobia and the role of novelty, 127 and sampling, 180 Innate responses of predators, 130 Importance of alternative prey, 181 Aposematism and gregariousness, 132 Signal detection theory, 181 Phylogenetic analysis of aposematism Genetic and evolutionary models, 182 and gregariousness, 134 Coevolutionary chases, 185 Contents ix Models involving population dynamics, 185 Myrmecomorphy by caterpillars, 237 Neural networks and evolution Ant chemical mimicry by parasitoid of Batesian mimicry, 188 wasps, 237 Automimicry in Batesian/Müllerian mimicry, 188 Protective mimicries among vertebrates, 239 Predator’s dilemma with potentially harmful prey, 190 Fish, 239 Batesian mimicry among fish, 239 Müllerian mimicry among fish, 239 6 ANTI‐PREDATOR MIMICRY. II. Batesian and Müllerian mimicry among EXPERIMENTAL TESTS, 191 terrestrial vertebrates, 239 Introduction, 192 The coral snake problem – Emsleyan Experimental tests of mimetic advantage,