Description

Book Synopsis

Symbiosis in Fishes provides comprehensive coverage of the biology of partnerships between fishes and invertebrates, ascending the phylogenetic scale, from luminescent bacteria, sponges and coelenterates to molluscs, crustaceans and echinoderms. Both facultative and obligatory partnerships are reviewed with emphasis on the behavioral, ecological and evolutionary aspects of fish symbiosis. Each of the eight chapters of this book focuses on a different group of partners. The structure, physiology and anti-predatory strategies of each group are described to provide the necessary background for the understanding of their partnerships with fishes. The formation of the associations, the degree of partner specificity and its regulation, as well as the benefits and costs for the fishes and their associates, communication between partners and their possible co-evolution are discussed in each chapter.

This is the first attempt to critically review in a single volume all associat

Trade Review

�Overall, Symbiosis in Fishesis a valuable scientific contribution because although symbiotic interactions interactions involving fish are common worldwide and can affect the dynamics of many ecosystems, this is the first comprehensive review of the topic.� (The Quarterly Review of Biology, 1 June 2015)

"Overall, the book reflects an exceptional breadth of knowledge and depth of integrative thinking, together with a deep appreciation for the subject of interspecific partnerships in fishes. As such, the book provides a superb resource and foundation for learning, teaching and further scientific inquiry." ( Journal of Fish Biology, 2015)



Table of Contents

Preface x

Introduction 1

1 The Associations between Fishes and Luminescent Bacteria 6

Luminescent Bacteria 6

Symbiotic Luminescent Bacteria in Fish Light Organs 8

Flashlight Fishes 11

Taxonomy and Distribution 11

The Light Organs 13

The Eye and the Light Organ 17

Reproduction, Larval and Light Organ Development 18

The Photophobic Response 20

The Use of Light by Flashlight Fishes 21

School Formation 22

Territorial Defense 22

Sexual Signaling 22

Deep Sea Ceratioid Anglerfishes 24

Structure, Diversity and Distribution 24

Reproductive Strategies 25

Obligatory Sexual Parasitism 26

Temporary Associations 28

Facultative Sexual Parasitism 29

Light Organ Structure and Development: Light and the Mechanisms Controlling its Emission 29

The Use of Lures by Anglerfishes 34

Ponyfishes 37

Structure, Distribution and Taxonomy 37

The Light Organ System (LOS) and Diversity of the Generated Light Patterns 38

Disruptive Illumination 40

Discrete Projected Luminescence (DPL) 41

Ventral Body Flash 41

Opercular Flash 42

Buccal Luminescence 42

Sex-Specific Signaling 43

Inception of the Association between Luminescent Bacteria and Ponyfishes 43

Sexual Dimorphism of the LOS, Sex-Specific Signaling and the Role of Sexual

Selection in the Evolution of Leiognathid Fishes 44

Specificity of the Partnerships between Luminescent Bacteria and Fishes 47

Optimization of the Benefits to Fishes from their Association with Bacteria 48

The Evolution of the Partnerships between Fishes and Luminescent Bacteria 49

References 52

2 The Associations between Fishes and Sponges 58

Sponges 58

Predator Deterrence by Sponges 59

Multiple Species Assemblages in Sponges 61

Obligatory Fish Symbionts and Adaptations for Living in Association with Sponges 62

Nutrition, Reproduction and Sponge Occupation by Obligatory Symbiotic Fishes 68

Partner Specificity and Sponge Sharing by Obligatory Symbiotic Fishes 69

Evolution of the Partnership Between Obligatory Fish Symbionts and Sponges 70

Sponges as Living Incubators of Fish Eggs 72

Facultative Partnerships Between Fishes and Sponges 74

References 75

3 The Associations between Fishes and Anthozoans 79

Sea Anemones 79

The Stinging Cells and their Release Mechanism 80

Obligatory Associations with Sea Anemones of Fishes of the Genera Amphiprion and Premnas 81

The Taxonomy, Distribution and Ecology of Host Sea Anemones and their Associated Fishes 81

The Protection of Anemone Fishes from Sea Anemones 86

Recognition, Attraction to and Selection of Sea Anemones by Anemone Fishes 93

Partner Specificity 104

Host Preference 106

Competitive Interactions 106

Stochastic Processes 109

Habitat Preference 109

Geographical Overlap 109

Protection from Sea Anemones 109

Species Coexistence 110

Adaptations of Anemone Fishes for Living with Sea Anemones 111

Protandric Sex Reversal 111

Monogamy and Mate Recognition 114

Step-fathering 118

Social Control of Growth and the Tolerance of Nonbreeders by the Breeders 118

Fish Territoriality, Aggression and the Sea Anemone 121

Limited Larval Dispersal and Natal Recruitment 124

Benefits and Costs to Anemone Fishes and Sea Anemones from being Associated and their Short-term Mutual Impacts 128

The Evolution of the Anemone Fish–Sea Anemone Partnership 134

The Facultative Associations Between Fishes and Sea Anemones 135

Protection from Sea Anemones 140

Partner Specificity 141

Settlement and Recruitment of D. trimaculatus to Sea Anemones 142

The Sharing of Sea Anemones with Anemone Fishes 143

Benefits and Costs to Facultative Fish Partners and Sea Anemones 144

The Associations Between Fishes and Scleractinian Corals 145

Scleractinian Corals 145

Microhabitat Selection by Coral Dwelling Fishes 146

Attraction of Pomacentrid Fishes to Corals 146

Attraction of Pomacentrids to Corals Inhabited by Conspecifics 150

Coral Occupation, Competiton and Coexistence of Coral dwelling Gobies 153

Adaptations to Habitat by Coral Dwelling Gobies 157

Small Size and Morphology 157

Noxious Skin 158

Hypoxia Tolerance and Air Breathing 158

Bidirectional Sex Reversal 159

Monogamy 161

Social Control of Growth 162

Multiple Species Assemblages Involving Coral Dwelling Gobies and Crustaceans 164

Benefits and Costs to Fishes and Corals for being Associated 169

Benefits to Fishes 169

Costs to Fishes 173

Benefits to Corals 177

Costs to Corals 180

Social Structure and Mating System Evolution in Coral Dwelling Damselfishes of the genus Dascyllus 181

References 186

4 The Associations between Fishes and Siphonophores 202

Siphonophores 202

Physalia physalis −the Portuguese Man-of-War 203

Fishes Associated with Siphonophores other than Physalia physalis 204

Fishes Associated with Physalia physalis 207

References 209

5 The Associations between Fishes and Scyphozoan Medusae 212

Scyphozoan medusae 212

Predation on Scyphozoan Medusae and their Structural and Behavioral Antipredator Defenses 214

Fishes Associated with Scyphozoan Medusae 215

The Protection of Fishes from Scyphozoan Medusae 217

Recognition and Attraction to Scyphozoan Medusae by Associated Fishes 217

Partner Specificity, Duration of the Medusa–Fish Bond and the Effects of the Medusae Size on the Associated Fishes 219

Benefits and Costs to Fishes and Medusae from being Associated 221

The Effects of Medusae on Fish Recruitment 225

The Association of Fishes with Floating Objects and the Fish–Medusa Partnership 226

References 227

6 The Associations between Fishes and Molluscs 230

The Association between Fishes and Cephalopods 230

Cephalopods 230

Octopus Dens, Foraging and Antipredatory Behavior 231

Scavenging Fishes Associated with Octopus Dens 233

Fishes Associated with Foraging Octopuses 234

Octopuses and Cleaning Symbiosis 238

Transport Associations between Octopuses and Fishes 239

Fishes Associated with Squid Schools 239

The Association between Fishes and Gastropods 241

Gastropods 241

Predation on Conchs, Antipredatory Strategies and Foraging in Conchs 241

The Association between Cardinal Fishes and Conchs 242

The Association between Nudibranchs and Gobiid Fishes 245

The Association between a Pearlfish and an Opisthobranch Gastropod 246

The Association between Fishes and Bivalves 246

Bivalves 246

The Glochidia Larvae of Freshwater Mussels and their Host Fishes 247

Bitterlings and their Freshwater Mussel Hosts 248

Attraction of the European Bitterling to Mussels and Choice of Oviposition Sites 250

Adaptations of Bitterling for Development Inside Freshwater Mussels 252

Male Reproductive Behavior and the Mussel 254

Female Reproductive Behavior and the Mussel 257

Host Utilization by Sympatric Bitterling Species 260

Costs and Benefits for the Mussel and Possible Coevolution of the

Bitterling–Mussel Partnership 263

Pearl Fishes Associated with Bivalves 265

The Association of Snailfish and Red Hake with Sea Scallops 265

References 269

7 The Associations between Fishes and Crustaceans 276

The Associations between Fishes and Cleaner Shrimps 276

Cleaning Symbiosis and Shrimp 276

Taxonomy, Morphology, Coloration and Distribution of Cleaner Shrimp 276

Cleaner Shrimp Activity 287

Associations between Cleaner Shrimp and Sea Anemones 288

Communication between Fishes and Cleaner Shrimp 292

Removal of Parasites versus Mucus by Cleaner Shrimp 294

Costs and Benefits for Cleaner Shrimp and Fish Clients and the Proximate Mechanisms for Cleaning 296

The Evolution of the Cleaner Shrimp–Fish Partnership 298

Feeding associations between fishes and crustaceans 299

Mixed Species Schools of Fishes and Crustaceans 300

Liparid Fishes Associated with Lithodid Crabs 301

The Associations between Fishes and Burrowing Brachyuran Crabs 303

Gobiid Fishes Associated with Burrowing Thalassinid Shrimp 305

Thalassinid Shrimp and their Burrows 305

The Facultative Association of Clevelandia ios with Callianassa californiensis and Upogebia pugettensis 307

The Obligatory Association of the Blind Goby Typhlogobius californiensis with Callianassa affinis 311

The Obligatory Association of Austrolethops wardi with Neaxius acanthus 313

The Obligatory Association of Didogobius amicuscardis with Axiopsis serratifrons 314

Gobiid Fishes Associated with Burrowing Alpheid Shrimps 316

Systematics of Gobies and Shrimps 316

Biogeography 318

Diet and Feeding Behavior 319

Habitat Specificity 322

Population Structure and Dynamics 324

Burrow Structure, Construction and Dynamics 326

Activity Rhythms 330

Aggressive Behavior and Territoriality of Goby and Shrimp 334

Reproduction of Goby and Shrimp 336

Interspecific Communication 338

Communication under Natural Conditions in Indo-Pacific Partnerships 338

Warning Signal Generation by Indo-Pacific Gobies in Response to

Predators and Models of Predators 340

Sequence and Information Analyses in Indo-Pacific Partnerships 342

Film Analysis of the Communication between the Goby Amblyeleotris steinitzi and the Shrimp Alpheus purpurilenticularis 344

Communication between Gobies and Shrimp in the Western Atlantic 347

Partner Specificity 349

Field Observations 349

Laboratory Experiments 350

The Mechanism Regulating Specificity 352

Goby–Shrimp Phylogeography 353

Costs and Benefits for Goby and Shrimp 357

Evolution 358

References 360

8 The Associations between Fishes and Echinoderms 371

The Association between Fishes and Sea Urchins 371

Sea Urchins 371

Sea Urchin Structural Defenses, Predation by Fishes and Antipredatory Strategies 371

Associated Fishes, their Size, Coloration and Sea Urchin Hosts 373

The Attraction of Associated Fishes to Sea Urchins 386

Benefits and Costs of the Fish–Sea Urchin Partnership 388

Partner Specificity in the Fish–Sea Urchin Association 390

The Evolution of the Fish–Sea Urchin Partnership 390

Mimicry of Sea Urchins by Fishes 391

The Association between Fishes and Crinoids 392

Crinoids 392

Predation on Crinoids by Fishes and Antipredatory Strategies of Feather Stars and Sea Lilies 393

Multiple Species Assemblages in Crinoids 394

Associated Fishes and Adaptations for Living with Crinoids 395

Attraction of Associated Fishes to Crinoids and Partner Specificity 398

Benefits and Costs of the Fish–Crinoid Partnership 398

Scarcity of Knowledge 398

The Association between Fishes and Sea Cucumbers 399

Sea Cucumbers 399

Predation on Sea Cucumbers by Fishes and their Structural and

Behavioral Antipredatory Defenses 400

Fishes Associated with Sea Cucumbers and their Life Cycles 401

Host Location, Penetration and Occupation by Pearlfishes 405

Pearlfish Nutrition 406

Pearlfish Reproductive Biology 408

Ecology and Partner Specificity of Pearlfish–Holothurian Associations 410

Acoustic Communication in Pearlfishes 414

Morphological and Physiological Adaptations to Inquilism 415

Benefits and Costs of the Pearlfish–Sea Cucumber Partnership 417

The Evolution of the Partnership between Pearlfishes and their Hosts 417

The Association between Fishes and Sea Stars 418

Sea Stars 418

Sea Star Structural and Behavioral Antipredatory Defenses 419

Feeding Associations between Sea Stars and Fishes 420

Cardinal Fishes Sheltering among Sea Star Spines 421

Pearlfishes Associated with Sea Stars 421

References 423

Species Index 431

Subject Index 443

Symbiosis in Fishes

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      Publisher: John Wiley and Sons Ltd
      Publication Date: Publication Date: 16/05/2014
      ISBN13: 9781405185899, 978-1405185899
      ISBN10: 1405185899

      Description

      Book Synopsis

      Symbiosis in Fishes provides comprehensive coverage of the biology of partnerships between fishes and invertebrates, ascending the phylogenetic scale, from luminescent bacteria, sponges and coelenterates to molluscs, crustaceans and echinoderms. Both facultative and obligatory partnerships are reviewed with emphasis on the behavioral, ecological and evolutionary aspects of fish symbiosis. Each of the eight chapters of this book focuses on a different group of partners. The structure, physiology and anti-predatory strategies of each group are described to provide the necessary background for the understanding of their partnerships with fishes. The formation of the associations, the degree of partner specificity and its regulation, as well as the benefits and costs for the fishes and their associates, communication between partners and their possible co-evolution are discussed in each chapter.

      This is the first attempt to critically review in a single volume all associat

      Trade Review

      �Overall, Symbiosis in Fishesis a valuable scientific contribution because although symbiotic interactions interactions involving fish are common worldwide and can affect the dynamics of many ecosystems, this is the first comprehensive review of the topic.� (The Quarterly Review of Biology, 1 June 2015)

      "Overall, the book reflects an exceptional breadth of knowledge and depth of integrative thinking, together with a deep appreciation for the subject of interspecific partnerships in fishes. As such, the book provides a superb resource and foundation for learning, teaching and further scientific inquiry." ( Journal of Fish Biology, 2015)



      Table of Contents

      Preface x

      Introduction 1

      1 The Associations between Fishes and Luminescent Bacteria 6

      Luminescent Bacteria 6

      Symbiotic Luminescent Bacteria in Fish Light Organs 8

      Flashlight Fishes 11

      Taxonomy and Distribution 11

      The Light Organs 13

      The Eye and the Light Organ 17

      Reproduction, Larval and Light Organ Development 18

      The Photophobic Response 20

      The Use of Light by Flashlight Fishes 21

      School Formation 22

      Territorial Defense 22

      Sexual Signaling 22

      Deep Sea Ceratioid Anglerfishes 24

      Structure, Diversity and Distribution 24

      Reproductive Strategies 25

      Obligatory Sexual Parasitism 26

      Temporary Associations 28

      Facultative Sexual Parasitism 29

      Light Organ Structure and Development: Light and the Mechanisms Controlling its Emission 29

      The Use of Lures by Anglerfishes 34

      Ponyfishes 37

      Structure, Distribution and Taxonomy 37

      The Light Organ System (LOS) and Diversity of the Generated Light Patterns 38

      Disruptive Illumination 40

      Discrete Projected Luminescence (DPL) 41

      Ventral Body Flash 41

      Opercular Flash 42

      Buccal Luminescence 42

      Sex-Specific Signaling 43

      Inception of the Association between Luminescent Bacteria and Ponyfishes 43

      Sexual Dimorphism of the LOS, Sex-Specific Signaling and the Role of Sexual

      Selection in the Evolution of Leiognathid Fishes 44

      Specificity of the Partnerships between Luminescent Bacteria and Fishes 47

      Optimization of the Benefits to Fishes from their Association with Bacteria 48

      The Evolution of the Partnerships between Fishes and Luminescent Bacteria 49

      References 52

      2 The Associations between Fishes and Sponges 58

      Sponges 58

      Predator Deterrence by Sponges 59

      Multiple Species Assemblages in Sponges 61

      Obligatory Fish Symbionts and Adaptations for Living in Association with Sponges 62

      Nutrition, Reproduction and Sponge Occupation by Obligatory Symbiotic Fishes 68

      Partner Specificity and Sponge Sharing by Obligatory Symbiotic Fishes 69

      Evolution of the Partnership Between Obligatory Fish Symbionts and Sponges 70

      Sponges as Living Incubators of Fish Eggs 72

      Facultative Partnerships Between Fishes and Sponges 74

      References 75

      3 The Associations between Fishes and Anthozoans 79

      Sea Anemones 79

      The Stinging Cells and their Release Mechanism 80

      Obligatory Associations with Sea Anemones of Fishes of the Genera Amphiprion and Premnas 81

      The Taxonomy, Distribution and Ecology of Host Sea Anemones and their Associated Fishes 81

      The Protection of Anemone Fishes from Sea Anemones 86

      Recognition, Attraction to and Selection of Sea Anemones by Anemone Fishes 93

      Partner Specificity 104

      Host Preference 106

      Competitive Interactions 106

      Stochastic Processes 109

      Habitat Preference 109

      Geographical Overlap 109

      Protection from Sea Anemones 109

      Species Coexistence 110

      Adaptations of Anemone Fishes for Living with Sea Anemones 111

      Protandric Sex Reversal 111

      Monogamy and Mate Recognition 114

      Step-fathering 118

      Social Control of Growth and the Tolerance of Nonbreeders by the Breeders 118

      Fish Territoriality, Aggression and the Sea Anemone 121

      Limited Larval Dispersal and Natal Recruitment 124

      Benefits and Costs to Anemone Fishes and Sea Anemones from being Associated and their Short-term Mutual Impacts 128

      The Evolution of the Anemone Fish–Sea Anemone Partnership 134

      The Facultative Associations Between Fishes and Sea Anemones 135

      Protection from Sea Anemones 140

      Partner Specificity 141

      Settlement and Recruitment of D. trimaculatus to Sea Anemones 142

      The Sharing of Sea Anemones with Anemone Fishes 143

      Benefits and Costs to Facultative Fish Partners and Sea Anemones 144

      The Associations Between Fishes and Scleractinian Corals 145

      Scleractinian Corals 145

      Microhabitat Selection by Coral Dwelling Fishes 146

      Attraction of Pomacentrid Fishes to Corals 146

      Attraction of Pomacentrids to Corals Inhabited by Conspecifics 150

      Coral Occupation, Competiton and Coexistence of Coral dwelling Gobies 153

      Adaptations to Habitat by Coral Dwelling Gobies 157

      Small Size and Morphology 157

      Noxious Skin 158

      Hypoxia Tolerance and Air Breathing 158

      Bidirectional Sex Reversal 159

      Monogamy 161

      Social Control of Growth 162

      Multiple Species Assemblages Involving Coral Dwelling Gobies and Crustaceans 164

      Benefits and Costs to Fishes and Corals for being Associated 169

      Benefits to Fishes 169

      Costs to Fishes 173

      Benefits to Corals 177

      Costs to Corals 180

      Social Structure and Mating System Evolution in Coral Dwelling Damselfishes of the genus Dascyllus 181

      References 186

      4 The Associations between Fishes and Siphonophores 202

      Siphonophores 202

      Physalia physalis −the Portuguese Man-of-War 203

      Fishes Associated with Siphonophores other than Physalia physalis 204

      Fishes Associated with Physalia physalis 207

      References 209

      5 The Associations between Fishes and Scyphozoan Medusae 212

      Scyphozoan medusae 212

      Predation on Scyphozoan Medusae and their Structural and Behavioral Antipredator Defenses 214

      Fishes Associated with Scyphozoan Medusae 215

      The Protection of Fishes from Scyphozoan Medusae 217

      Recognition and Attraction to Scyphozoan Medusae by Associated Fishes 217

      Partner Specificity, Duration of the Medusa–Fish Bond and the Effects of the Medusae Size on the Associated Fishes 219

      Benefits and Costs to Fishes and Medusae from being Associated 221

      The Effects of Medusae on Fish Recruitment 225

      The Association of Fishes with Floating Objects and the Fish–Medusa Partnership 226

      References 227

      6 The Associations between Fishes and Molluscs 230

      The Association between Fishes and Cephalopods 230

      Cephalopods 230

      Octopus Dens, Foraging and Antipredatory Behavior 231

      Scavenging Fishes Associated with Octopus Dens 233

      Fishes Associated with Foraging Octopuses 234

      Octopuses and Cleaning Symbiosis 238

      Transport Associations between Octopuses and Fishes 239

      Fishes Associated with Squid Schools 239

      The Association between Fishes and Gastropods 241

      Gastropods 241

      Predation on Conchs, Antipredatory Strategies and Foraging in Conchs 241

      The Association between Cardinal Fishes and Conchs 242

      The Association between Nudibranchs and Gobiid Fishes 245

      The Association between a Pearlfish and an Opisthobranch Gastropod 246

      The Association between Fishes and Bivalves 246

      Bivalves 246

      The Glochidia Larvae of Freshwater Mussels and their Host Fishes 247

      Bitterlings and their Freshwater Mussel Hosts 248

      Attraction of the European Bitterling to Mussels and Choice of Oviposition Sites 250

      Adaptations of Bitterling for Development Inside Freshwater Mussels 252

      Male Reproductive Behavior and the Mussel 254

      Female Reproductive Behavior and the Mussel 257

      Host Utilization by Sympatric Bitterling Species 260

      Costs and Benefits for the Mussel and Possible Coevolution of the

      Bitterling–Mussel Partnership 263

      Pearl Fishes Associated with Bivalves 265

      The Association of Snailfish and Red Hake with Sea Scallops 265

      References 269

      7 The Associations between Fishes and Crustaceans 276

      The Associations between Fishes and Cleaner Shrimps 276

      Cleaning Symbiosis and Shrimp 276

      Taxonomy, Morphology, Coloration and Distribution of Cleaner Shrimp 276

      Cleaner Shrimp Activity 287

      Associations between Cleaner Shrimp and Sea Anemones 288

      Communication between Fishes and Cleaner Shrimp 292

      Removal of Parasites versus Mucus by Cleaner Shrimp 294

      Costs and Benefits for Cleaner Shrimp and Fish Clients and the Proximate Mechanisms for Cleaning 296

      The Evolution of the Cleaner Shrimp–Fish Partnership 298

      Feeding associations between fishes and crustaceans 299

      Mixed Species Schools of Fishes and Crustaceans 300

      Liparid Fishes Associated with Lithodid Crabs 301

      The Associations between Fishes and Burrowing Brachyuran Crabs 303

      Gobiid Fishes Associated with Burrowing Thalassinid Shrimp 305

      Thalassinid Shrimp and their Burrows 305

      The Facultative Association of Clevelandia ios with Callianassa californiensis and Upogebia pugettensis 307

      The Obligatory Association of the Blind Goby Typhlogobius californiensis with Callianassa affinis 311

      The Obligatory Association of Austrolethops wardi with Neaxius acanthus 313

      The Obligatory Association of Didogobius amicuscardis with Axiopsis serratifrons 314

      Gobiid Fishes Associated with Burrowing Alpheid Shrimps 316

      Systematics of Gobies and Shrimps 316

      Biogeography 318

      Diet and Feeding Behavior 319

      Habitat Specificity 322

      Population Structure and Dynamics 324

      Burrow Structure, Construction and Dynamics 326

      Activity Rhythms 330

      Aggressive Behavior and Territoriality of Goby and Shrimp 334

      Reproduction of Goby and Shrimp 336

      Interspecific Communication 338

      Communication under Natural Conditions in Indo-Pacific Partnerships 338

      Warning Signal Generation by Indo-Pacific Gobies in Response to

      Predators and Models of Predators 340

      Sequence and Information Analyses in Indo-Pacific Partnerships 342

      Film Analysis of the Communication between the Goby Amblyeleotris steinitzi and the Shrimp Alpheus purpurilenticularis 344

      Communication between Gobies and Shrimp in the Western Atlantic 347

      Partner Specificity 349

      Field Observations 349

      Laboratory Experiments 350

      The Mechanism Regulating Specificity 352

      Goby–Shrimp Phylogeography 353

      Costs and Benefits for Goby and Shrimp 357

      Evolution 358

      References 360

      8 The Associations between Fishes and Echinoderms 371

      The Association between Fishes and Sea Urchins 371

      Sea Urchins 371

      Sea Urchin Structural Defenses, Predation by Fishes and Antipredatory Strategies 371

      Associated Fishes, their Size, Coloration and Sea Urchin Hosts 373

      The Attraction of Associated Fishes to Sea Urchins 386

      Benefits and Costs of the Fish–Sea Urchin Partnership 388

      Partner Specificity in the Fish–Sea Urchin Association 390

      The Evolution of the Fish–Sea Urchin Partnership 390

      Mimicry of Sea Urchins by Fishes 391

      The Association between Fishes and Crinoids 392

      Crinoids 392

      Predation on Crinoids by Fishes and Antipredatory Strategies of Feather Stars and Sea Lilies 393

      Multiple Species Assemblages in Crinoids 394

      Associated Fishes and Adaptations for Living with Crinoids 395

      Attraction of Associated Fishes to Crinoids and Partner Specificity 398

      Benefits and Costs of the Fish–Crinoid Partnership 398

      Scarcity of Knowledge 398

      The Association between Fishes and Sea Cucumbers 399

      Sea Cucumbers 399

      Predation on Sea Cucumbers by Fishes and their Structural and

      Behavioral Antipredatory Defenses 400

      Fishes Associated with Sea Cucumbers and their Life Cycles 401

      Host Location, Penetration and Occupation by Pearlfishes 405

      Pearlfish Nutrition 406

      Pearlfish Reproductive Biology 408

      Ecology and Partner Specificity of Pearlfish–Holothurian Associations 410

      Acoustic Communication in Pearlfishes 414

      Morphological and Physiological Adaptations to Inquilism 415

      Benefits and Costs of the Pearlfish–Sea Cucumber Partnership 417

      The Evolution of the Partnership between Pearlfishes and their Hosts 417

      The Association between Fishes and Sea Stars 418

      Sea Stars 418

      Sea Star Structural and Behavioral Antipredatory Defenses 419

      Feeding Associations between Sea Stars and Fishes 420

      Cardinal Fishes Sheltering among Sea Star Spines 421

      Pearlfishes Associated with Sea Stars 421

      References 423

      Species Index 431

      Subject Index 443

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