Mimicry, Crypsis, Masquerade and other Adaptive Resemblances

Title Page
Copyright Page
Contents
Preface
Acknowledgements
Chapter 1 Introduction and Classification of Mimicry Systems
	A brief history
	On definitions of ‘mimicry’ and adaptive resemblance
		The concept of ‘adaptive resemblance’
	The classification of mimicry systems,
		Wickler’s system
		Vane-Wright’s system
		Georges Pasteur (1930–2015)
		Other approaches
			Endler
			Zabka & Tembrock
			Maran
	Mimicry as demonstration of evolution
Chapter 2 Camouflage: Crypsis and Disruptive Colouration in Animals
	Introduction
	Distinguishing crypsis from masquerade
	Crypsis examples
	Countershading
		Experimental tests of concealment by countershading
		Bioluminescent counter-illumination
	Background matching
		Visual sensitivity of predators
		To make a perfect match or compromise
	Colour polymorphism
		Seasonal colour polymorphism
		Butterfly pupal colour polymorphism
		Winter pelage: pelts and plumage
	Melanism
		Industrial melanism
		Fire melanism
	Background selection
		Orientation and positioning
	Transparency
	Reflectance and silvering
	Adaptive colour change
		Caterpillars and food plant colouration
		Daily and medium‐paced changes
		Rapid colour change
			Chameleons
			Cephalopod chromatophores and dermal papillae
	Bird eggs and their backgrounds
	Disguising your eyes
	Disruptive and distractive markings
		Edge-intercepting patches
		Distractive markings
		Zebra stripes and tsetse flies
	Stripes and motion dazzle – more zebras, kraits and tigers
		Computer graphics experiments with human subjects
		Observations on real animals
		Comparative analysis
	Dual signals
	Protective crypsis in non-visual modalities
	Apostatic and antiapostatic selection
		Search images
		Experimental tests of search image
		Gestalt perception
	Effect of cryptic prey variability
		Reflexive selection and aspect diversity
		Searching for cryptic prey – mathematical models
	Ontogenetic changes and crypsis
	Hiding the evidence
		Petiole clipping by caterpillars
		Exogenous crypsis
	Military camouflage and masquerade
Chapter 3 Camouflage: Masquerade
	Introduction
	Classic examples
		Twigs as models
		Leaves (alive or dead) as models
		Bird dropping resemblances
		Spider web stabilimenta
		Tubeworms, etc.
	Experimental tests of survival value of masquerade
	Ontogenetic changes and masquerade
	Thanatosis (death feigning)
		Feign or flee? The trade-offs of thanatosis
		Other aspects of death mimicry
	Seedless seeds and seedless fruit
Chapter 4 Aposematism and its Evolution
	Introduction
	Initial evolution of aposematism
		Associations of unpalatable experience with place
	Mathematical models and ideas of warning colouration evolution
		Kin selection models
		Green beard selection
		Family selection models
		Individual selection models
		Spatial models and metapopulations
	Handicap and signal honesty
		Early warnings – reflex bleeding, vomiting and other noxious secretions
	Longevity of aposematic protected taxa
	Macroevolutionary consequences
	Experimental studies
		Tough aposematic prey and individual selection
		Pyrazine and other early warnings
	Learning and memorability
		Strength of obnoxiousness
		Is the nature of the protective compound important?
		Neophobia and the role of novelty
	Innate responses of predators
	Aposematism and gregariousness
		Phylogenetic analysis of aposematism and gregariousness
	Behaviour of protected aposematic animals
		Of birds and butterflies
		Evolution of sluggishness
	Origins of protective compounds
		Plant-derived toxins
		Cardiac glycosides
		Pyrrolizidine alkaloids
		De novo synthesis of protective compounds
		Obtaining toxins from animal sources
		Costs of chemical defence
	Aposematism with non-chemical defence
		Escape speed and low profitability
	Parasitoids and aposematic insects
	Diversity of aposematic forms
		Egg load assessment
	Proof of aposematism
		Bioluminescence as a warning signal
		Warning sounds
		Warning colouration in mammals
		Weapon advertisement
		Mutualistic aposematism
		Aposematism induced by a parasite
		Aposematic commensalism
	Polymorphism and geographic variation in aposematic species
	Aposematism in plants
		Synergistic selection of unpalatability in plants
	Aposematism in fungi
	Why are some unpalatable organisms aposematic and others not?
Chapter 5 Anti-Predator Mimicry. I. Mathematical Models
	Introduction
	Properties of models, rewards, learning rates and numerical relationships
	Simple models and their limitations
		Müller’s original model
		Simple models of Batesian and  Müllerian mimicry
		Are Batesian and Müllerian mimicry different?
		An information theory model
		Monte-Carlo simulations
	More refined models – time, learning, forgetting and sampling
		Importance of alternative prey
		Signal detection theory
		Genetic and evolutionary models
		Coevolutionary chases
		Models involving population dynamics
		Neural networks and evolution of Batesian mimicry
	Automimicry in Batesian/Müllerian mimicry
	Predator’s dilemma with potentially harmful prey
Chapter 6 Anti-predator mimicry. II. Experimental tests
	Introduction
	Experimental tests of mimetic advantage
		How similar do mimics need to be?
		Is a two-step process necessary?
	Relative abundances of models and mimics in nature
	Sex-limited mimicries and mimetic load
		Mimetic load
	Apostatic selection and Batesian mimicry
		Müllerian mimicry and unequal defence
	Imperfect (satyric) mimicry
Chapter 7 Anti-predator Mimicry. III. Batesian and Müllerian Examples
	Introduction
	Types of model
		Mimicry of slow flight in butterflies
	The Batesian/Müllerian spectrum
	Famous butterflies: ecology, genetics and supergenes
		Heliconius
			Hybrid zones
			Wing pattern genetics
			Modelling polymorphism
		Danaus and Hypolimnas
		Papilio dardanus
		Papilio glaucus
		Papilio memnon
		Supergenes and their origins
	Mimicry between caterpillars
	Some specific types of model among insects
		Wasp (and bee) mimicry
			How to look like a wasp
			Time of appearance of aculeate mimics
			Pseudostings and pseudostinging behaviour
		Wasmannian (or ant) mimicry
			Ant mimicry as defence against predation
			Ant mimicry by spiders
			Spiders that feed on ants
			How to look like an ant or an ant carrying something?
			Myrmecomorphy by caterpillars
			Ant chemical mimicry by parasitoid wasps
	Protective mimicries among vertebrates
		Fish
			Batesian mimicry among fish
			Müllerian mimicry among fish
			Batesian and Müllerian mimicry among terrestrial vertebrates
			The coral snake problem – Emsleyan (or Mertensian) mimicry
			Other snakes, zig-zag markings and head shape
			Mimicry of invertebrates by terrestrial vertebrates
	Inaccurate (satyric) mimics
	Mimicry of model behaviour
		Aide mémoire mimicry
	Batesian–Poultonian (predator) mimicry
	Mimicry within predator–prey and host–parasite systems
	Bluff and appearing larger than you are
	Collective mimicry including an aggressive mimicry
	Jamming
	Man as model – the case of the samurai crab
Chapter 8 Anti-predator Mimicry. Attack Deflection, Schooling, etc.
	Introduction
	Attack deflection devices
		Eyespots
			Experimental tests of importance of eyespot features
			Eyespots in butterflies
			Wing marginal eyespots
			Eyes with sparkles
			Eyespots on caterpillars
			Importance of eyespot conspicuousness
			Eyespots and fish
			Not just an eyespot but a whole head, winking and other enhancements
		Reverse mimicry
			Insects
			Reverse mimicry in flight
			Reverse mimicry in terrestrial vertebrates
	Other deflectors
		Injury feigning in nesting birds
		Tail-shedding (urotomy) in lizards and snakes
	Flash and startle colouration
		Intimidating displays and bizarre mimicries
	Schooling, flocking and predator confusion
		‘Social’ mimicry in birds and fish
		Alarm call mimicry for protection
Chapter 9 Anti-herbivory deceptions
	Introduction
	Crypsis as protection in plants
		Leaf mottling and variegation for crypsis
		Mistletoes and lianas
		Fruit masquerade by leaves
	Protective Batesian and Müllerian mimicry in plants
		False indicators of damage or likely future damage
		Conspicuousness of leafmines
		Dark central florets in some Apiaceae
		Mimicry of silk or fungal hyphae
		Insect egg mimics
		Defensive aphid and caterpillar mimicry in plants
		Aphid deterrence by alarm pheromone mimicry
		Ant mimicry in plants
		Of orchids and bees
		Carrion mimicry as defence
		Algae and corals
		Plant galls
		Experimental evidence for plant aposematism and Batesian mimetic potential in plants
Chapter 10 Aggressive deceptions
	Introduction
		Cryptic versus alluring features
	Crypsis and masquerade by predators
		Stealth
		Shadowing
		Seasonal polymorphisms in predators
		Why seabirds are black and white (and grey)
		Chemical crypsis by a predatory fish
	Alluring mimicries
		Flower mimicry
		Rain mimicry
		Physical lures
		Angling fish
		Caudal (and tongue) lures in reptiles
		Caudal lure in a dragonfly
		Death feigning as a lure
		Other prey and food mimicry
		The case of the German cockroach
	Wolves in sheeps’ clothing
		Vulture-like hawks
		Cleaner fish and their mimics
		Mingling with an innocuous crowd
	Duping by mimicry of competitors
	Seeming to be conspecific
		Getting close
		Appearing to be a potential mate
		Pheromone lures
	Mimicking danger as a flushing device
		Human use of aggressive mimicry
	Cuckoldry, inquilines and brood parasitism
		Cuckoldry in birds
		Gentes and ‘cuckoo’ eggs
			Cues for egg rejection
		Mimicry by chicks – genetic and substantive differences
			Cuckoo chick appearance
			Begging calls
		Cuckoo and host coevolution
		Mimicry between adult cuckoos and their hosts
		Hawk mimicry by adult cuckoos
		Mimicry of harmless birds by adult cuckoos
		Brood parasitism and inquilinism in social insects
		Cuckoo bees and cuckoo wasps
		Kleptoparasites of bees
		Myrmecophily
		Acquired chemical mimicry in social parasites and inquilines
		Brood-parasitic and slave-making ants
		Chemical mimicry and ant and termite inquilines
			A brood-parasitic aphid
		Ants and aphid trophallaxis
		Aphidiine parasitoids of ant‐attended aphids
	Does aggressive mimicry occur in plants?
Chapter 11 Sexual Mimicries in Animals (Including Humans)
	Introduction
	Mimicking the opposite sex
		Female mimicry by males
		Avoiding aggression from competing males
		Mate guarding through distracting other males
		Androchromatism and male mimicry by females
		Egg dummies on fish
		Food dummies and sex
		Mimicry by sperm-dependent all-female lineages
		Female genital mimicry in a female
		Energy-saving cheating for sex
	Behavioural deceptions in higher vertebrates
		Polygynous birds
		Deceptive use of alarm calls and paternity protection
		Female–female mounting behaviour in mammals and birds
		Mimicry in humans
		Make-up, clothes and silicone
		Cryptic oestrus in humans
		Flirting in humans
Chapter 12 Reproductive mimicries in plants
	Introduction
	Pollinator deception
		Pollinator sex pheromone mimicry
		Food deception
		Specific floral mimicry
		Generalised floral mimicry
		Mimicry of a fungus-infected plant
		Brood-site/oviposition-site deception
		Shelter mimicry
	Flower similarity over time
	Flower automimicry – intraspecific food deception (bakerian mimicry)
	Mathematical modelling of sexual deception by plants
	Pollinator guild syndromes
		Bird-pollinated systems
Chapter 13 Intra- and Interspecific Cooperation, Competition and Hierarchies
	Introduction
	Remaining looking young
		Delayed plumage maturation
		Interspecific social dominance mimicry
		Bird song and alarm call mimicry – deceptive acquisition of resources
	Wicklerian mimicry – mimicry of opposite sex to reduce aggression
		Female resemblance in male primates
		Social appeasement by female mimicry in an insect
	Hyperfemininity in prereproductive adolescent primates
	Mimicry of male genitalia by females
		The case of the spotted hyaena
		Mimicry of male genitalia in other mammals
		Phallic mimicry by males
	Appetitive (foraging) mimicry
		Appetitive mimicry and deceptive use of alarm calls
		Beau Geste and seeming to be more than you are
		Appearing older than you are
	Weapon automimicry
Chapter 14 Adaptive Resemblances and Dispersal: Seeds, Spores and Eggs
	Introduction
	Fruit and seed dispersal by birds
		Warningly coloured fruit
		Fruit mimicry by seeds
		Seed dispersal by humans, arable weeds and Vavilovian mimicry
		Seed elaiosomes and their insect mimics
	Mimicry by parasites to facilitate host finding
		The trematode and the snail
		The trematode and the fish
		Pocketbook clams and fish
		‘Termite balls’
		Pseudoflowers, pseudo-anthers and pseudo-pollen
		Truffles
		Mimicry of dead flesh by fungi and mosses
		Deception of dung beetles by fruit
Chapter 15 Molecular Mimicry: Parasites, Pathogens and Plants
	Introduction
	Macro-animal systems
		Anemone fish
		Parasitic helminthes
		Platyhelminthes (Trematoda)
		Tapeworms (Platyhelminthes: Cestoda)
		Parasitic nematodes
		Parasitoid wasp eggs
	Pathogenic fungi
	Protista
		Chagas’ disease
	Microbial systems
		Bacterial chemical mimicry and  autoimmune responses
		Helicobacter pylori
		Campylobacter jejuni
		Mimicry by plant-pathogenic bacteria
	Viruses
	Plants
		Sugar, toxin and satiation mimicry
		Phytoecdysteroids – plant chemicals that mimic insect moulting hormone
		Plant oestrogens – phyto-contraceptives
Extended glossary
References
Author index
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