Donald L.J. Quicke retired in 2013 to live in Thailand where he is a Visiting Professor at Chulalongkorn University. Hestudied zoology at Oxford University where he became especially interested in mimicry. In 1976 he travelled to Kenya to experience tropical biodiversity and more of the diversity of life and his work there on insect coloration fertilised his interests as well as on parasitoid wasps, another of his many passions. From then on he kept abreast of the increasingly experimental and theoretical developments in the field even though his academic research took him in diverse other directions. Having now retired he has been able devote his time, in addition to bird watching and butterfly photography, to synthesising and extending his interest in this topic. Mimicry, Crypsis, Masquerade and other Adaptive Resemblances is the result of this work.

Preface, xiii

A comment on statistics, xv

A comment on scientific names, xvi

Acknowledgements, xvii

1 INTRODUCTION AND CLASSIFICATION OF MIMICRY SYSTEMS, 1

A brief history, 2

On definitions of 'mimicry' and adaptive resemblance, 3

The concept of 'adaptive resemblance', 8

The classification of mimicry systems, 9

Wickler's system, 9

Vane¿Wright's system, 10

Georges Pasteur (1930-2015), 11

Other approaches, 13

Endler, 13

Zabka & Tembrock, 13

Maran, 14

Mimicry as demonstration of evolution, 14

2 CAMOUFLAGE: CRYPSIS AND DISRUPTIVE COLOURATION IN ANIMALS, 19

Introduction, 20

Distinguishing crypsis from masquerade, 20

Crypsis examples, 24

Countershading, 24

Experimental tests of concealment by countershading, 27

Bioluminescent counter¿illumination, 28

Background matching, 29

Visual sensitivity of predators, 30

To make a perfect match or compromise, 31

Colour polymorphism, 32

Seasonal colour polymorphism, 32

Butterfly pupal colour polymorphism, 32

Winter pelage: pelts and plumage, 35

Melanism, 37

Industrial melanism, 37

Fire melanism, 40

Background selection, 41

Orientation and positioning, 43

Transparency, 45

Reflectance and silvering, 47

Adaptive colour change, 49

Caterpillars and food plant colouration, 50

Daily and medium¿paced changes, 54

Rapid colour change, 56

Chameleons, 56

Cephalopod chromatophores and dermal papillae, 57

Bird eggs and their backgrounds, 58

Disguising your eyes, 61

Disruptive and distractive markings, 61

Edge¿intercepting patches, 61

Distractive markings, 63

Zebra stripes and tsetse flies, 66

Stripes and motion dazzle - more zebras, kraits and tigers, 69

Computer graphics experiments with human subjects, 69

Observations on real animals, 69

Comparative analysis, 71

Dual signals, 72

Protective crypsis in non¿visual modalities, 73

Apostatic and antiapostatic selection, 73

Search images, 74

Experimental tests of search image, 76

Gestalt perception, 76

Effect of cryptic prey variability, 77

Reflexive selection and aspect diversity, 77

Searching for cryptic prey - mathematical models, 80

Ontogenetic changes and crypsis, 81

Hiding the evidence, 82

Petiole clipping by caterpillars, 82

Exogenous crypsis, 82

Military camouflage and masquerade, 85

3 CAMOUFLAGE: MASQUERADE, 87

Introduction, 88

Classic examples, 88

Twigs as models, 88

Leaves (alive or dead) as models, 88

Bird dropping resemblances, 89

Spider web stabilimenta, 93

Tubeworms, etc., 94

Experimental tests of survival value of masquerade, 94

Ontogenetic changes and masquerade, 97

Thanatosis (death feigning), 97

Feign or flee? The trade¿offs of thanatosis, 100

Other aspects of death mimicry, 100

Seedless seeds and seedless fruit, 100

4 APOSEMATISM AND ITS EVOLUTION, 103

Introduction, 104

Initial evolution of aposematism, 108

Associations of unpalatable experience with place, 109

Mathematical models and ideas of warning colouration evolution, 112

Kin selection models, 112

Green beard selection, 112

Family selection models, 113

Individual selection models, 113

Spatial models and metapopulations, 116

Handicap and signal honesty, 117

Early warnings - reflex bleeding, vomiting and other noxious secretions, 120

Longevity of aposematic protected taxa, 121

Macroevolutionary consequences, 121

Experimental studies, 121

Tough aposematic prey and individual selection, 121

Pyrazine and other early warnings, 123

Learning and memorability, 124

Strength of obnoxiousness, 126

Is the nature of the protective compound important?, 126

Neophobia and the role of novelty, 127

Innate responses of predators, 130

Aposematism and gregariousness, 132

Phylogenetic analysis of aposematism and gregariousness, 134

Behaviour of protected aposematic animals, 135

Of birds and butterflies, 135

Evolution of sluggishness, 139

Origins of protective compounds, 140

Plant¿derived toxins, 140

Cardiac glycosides, 141

Pyrrolizidine alkaloids, 144

De novo synthesis of protective compounds, 145

Obtaining toxins from animal sources, 147

Costs of chemical defence, 149

Aposematism with non¿chemical defence, 150

Escape speed and low profitability, 150

Parasitoids and aposematic insects, 152

Diversity of aposematic forms, 152

Egg load assessment, 154

Proof of aposematism, 154

Bioluminescence as a warning signal, 155

Warning sounds, 155

Warning colouration in mammals, 157

Weapon advertisement, 158

Mutualistic aposematism, 160

Aposematism induced by a parasite, 161

Aposematic commensalism, 161

Polymorphism and geographic variation in aposematic species, 161

Aposematism in plants, 163

Synergistic selection of unpalatability in plants, 165

Aposematism in fungi, 166

Why are some unpalatable organisms aposematic and others not?, 167

5 ANTI¿PREDATOR MIMICRY. I. MATHEMATICAL MODELS, 171

Introduction, 172

Properties of models, rewards, learning rates and numerical relationships, 172

Simple models and their limitations, 173

Muller's original model, 173

Simple models of Batesian and Mullerian mimicry, 173

Are Batesian and Mullerian mimicry different?, 174

An information theory model, 176

Monte¿Carlo simulations, 177

More refined models - time, learning, forgetting and sampling, 180

Importance of alternative prey, 181

Signal detection theory, 181

Genetic and evolutionary models, 182

Coevolutionary chases, 185

Models involving population dynamics, 185

Neural networks and evolution of Batesian mimicry, 188

Automimicry in Batesian/Mullerian mimicry, 188

Predator's dilemma with potentially harmful prey, 190

6 ANTI¿PREDATOR MIMICRY. II. EXPERIMENTAL TESTS, 191

Introduction, 192

Experimental tests of mimetic advantage, 192

How similar do mimics need to be?, 194

Is a twöstep process necessary?, 198

Relative abundances of models and mimics in nature, 198

Sex¿limited mimicries and mimetic load, 198

Mimetic load, 203

Apostatic selection and Batesian mimicry, 204

Mullerian mimicry and unequal defence, 204

Imperfect (satyric) mimicry, 206

7 ANTI¿PREDATOR MIMICRY. III. BATESIAN AND MULLERIAN EXAMPLES, 213

Introduction, 214

Types of model, 214

Mimicry of slow flight in butterflies, 214

The Batesian/Mullerian spectrum, 215

Famous butterflies: ecology, genetics and supergenes, 216

Heliconius, 216

Hybrid zones, 217

Wing pattern genetics, 219

Modelling polymorphism, 220

Danaus and Hypolimnas, 220

Papilio dardanus, 221

Papilio glaucus, 223

Papilio memnon, 223

Supergenes and their origins, 223

Mimicry between caterpillars, 224

Some specific types of model among insects, 225

Wasp (and bee) mimicry, 225

How to look like a wasp, 228

Time of appearance of aculeate mimics, 228

Pseudostings and pseudostinging behaviour, 230

Wasmannian (or ant) mimicry, 231

Ant mimicry as defence against predation, 231

Ant mimicry by spiders, 234

Spiders that feed on ants, 236

How to look like an ant or an ant carrying something?, 236

Myrmecomorphy by caterpillars, 237

Ant chemical mimicry by parasitoid wasps, 237

Protective mimicries among vertebrates, 239

Fish, 239

Batesian mimicry among fish, 239

Mullerian mimicry among fish, 239

Batesian and Mullerian mimicry among terrestrial vertebrates, 239

The coral snake problem - Emsleyan (or Mertensian) mimicry, 240

Other snakes, zig¿zag markings and head shape, 244

Mimicry of invertebrates by terrestrial vertebrates, 246

Inaccurate (satyric) mimics, 248

Mimicry of model behaviour, 249

Aide mémoire mimicry, 250

Batesian-Poultonian (predator) mimicry, 251

Mimicry within predator-prey and host-parasite systems, 253

Bluff and appearing larger than you are, 253

Collective mimicry including an aggressive mimicry, 255

Jamming, 255

Man as model - the case of the samurai crab, 258

8 ANTI¿PREDATOR MIMICRY. ATTACK DEFLECTION, SCHOOLING, ETC., 259

Introduction, 260

Attack deflection devices, 260

Eyespots, 260

Experimental tests of importance of eyespot features, 262

Eyespots in butterflies, 266

Wing marginal eyespots, 267

Eyes with sparkles, 267

Eyespots on caterpillars, 269

Importance of eyespot conspicuousness, 269

Eyespots and fish, 269

Not just an eyespot but a whole head, winking and other enhancements, 271

Reverse mimicry, 271

Insects, 271

Reverse mimicry in flight, 275

Reverse mimicry in terrestrial vertebrates, 275

Other deflectors, 277

Injury feigning in nesting birds, 277

Tail¿shedding (urotomy) in lizards and snakes, 277

Flash and startle colouration, 280

Intimidating displays and bizarre mimicries, 283

Schooling,...