Mimicry, Crypsis, Masquerade and other Adaptive Resemblances

578 Pages·2017·86.115 MB·English

by  Donald L. J. Quicke

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
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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
and tigers, 69
2 CAMOUFLAGE: CRYPSIS AND DISRUPTIVE
Computer graphics experiments with
COLOURATION IN ANIMALS, 19
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