Dr Robert 'Bob' Wootton was Reader in Ecology at the Institute of Biological, Environmental and Rural Sciences, Aberystwyth University, UK. Bob's research addressed the ecology, behaviour and evolutionary biology of fishes.Dr Carl Smith is a Lecturer in the School of Biology at the University of St Andrews, UK and also holds a research position at the Institute of Vertebrate Biology of the Czech Academy of Sciences in Brno, Czech Republic. Carl's research focuses on mating system evolution in fishes.

“I was very impressed by this book, even if it does not delve deeply into the mathematics of evolutionary theory applied to the topics.”  (Environ Biol Fish, 1 September 2015)

• Preface xvii Acknowledgements xixAbbreviations and acronyms xxi1 Introduction 1Introduction 1Reproductive modes of the teleosts 2Gender systems of teleosts 2Spawning dynamics 4Modes of fertilisation 5Mating systems 5Secondary sexual characteristics 6Parental care 6Reproductive guilds 6Reproductive diversity in teleosts: an explanatory framework 6Phylogenetic relationships of the teleosts 8Life-history theory 9Aims of the volume 112 Sex determination 13Introduction 13Genotypic sex determination 14Monofactorial genotypic sex determination 14Multifactorial genotypic sex determination 20Polyfactorial (polygenic) genotypic sex determination 20Environmental sex determination 21Evolution of sex-determining mechanisms 263 Sex differentiation 31Introduction 31Embryology of the gonads 32Origin and migration of primordial germ cells 33Sex differentiation in the gonadal anlagen 35Initial differentiation of the gonads 36Initial differentiation of ovaries 38Initial differentiation of testes 38Genetic control of early gonadal differentiation 40Synthesis 424 Gametogenesis 45Introduction 46Structure of ovaries 48Gross morphology of the ovaries 48Development of female gonoducts 51Oogenesis 52Oogonial proliferation and oogonial nest formation 53Chromatin nucleolus stage 53Primary growth 54Secondary growth – vitellogenesis 55Vitellogenin and the zona pellucida proteins 56Polarity of the oocyte 58Oocyte maturation 58Ovulation 60Atresia 60Fertilisation 62Genetic control of oogenesis 64Dynamics of oocyte development 65Spawning dynamics and fecundity 66Structure of the testes 66Gross morphology of a testis 66Development of male gonoducts 69Accessory structures associated with the testes and gonoducts 70Spermatogenesis 71Spermatocysts 71Sertoli cells 71Stages of spermatogenesis within a spermatogenic spermatocyst 73Differentiated spermatogonia 73Primary spermatocysts 73Secondary spermatocysts 73Spermatids 74Spermiation and capacitation 75Types of spermatozoa 76Structure and phylogenetic distribution 76Spermatozoa viability 77Genetic control of spermatogenesis 79Dynamics of spermatogenesis 79Quantitative analysis of sperm production 805 Endocrinology of reproduction 81Introduction 82Brain–pituitary–gonad reproductive axis in vertebrates 83Levels of analysis 85Gonadal steroids 86Sex steroid synthesis 86Sex steroid receptors 87Endocrine control of oogenesis 88Oogonial proliferation and primary growth 88Secondary growth: vitellogenesis 89Maturation and ovulation 90Endocrine control of oogenesis in species with batch spawning 91Endocrine control of spermatogenesis 93Spermatogonial proliferation 95Initiation of meiosis and formation of spermatocysts 95Spermiation 96Patterns of spawning in relation to spermatogenesis 96The pituitary and the gonadotrophins 98Brain–pituitary relationship 98Gonadotrophs and gonadotrophins 99Gonadotrophin receptors 100Hypothalamic control of the pituitary 101Gonadotrophin-releasing hormone 101Functions of gonadotrophin-releasing hormone 102Gonadotrophin-releasing hormone receptors 102Distribution of gonadotrophin-releasing hormone neurons in the brain 103Role of dopamine as an inhibitor of reproduction 104Role of other brain factors in the control of gonadotrophs 105Feedback control in the brain–pituitary–gonad reproductive axis 106A provisional model for the control of the developmental dynamics of ovaries and testes 109Puberty in teleosts 109Physiological control of puberty 110Interactions between growth and reproductive axes 113Endocrines, behaviour and secondary sexual characteristics 116Four model species 117Role of endocrines in regulation of reproductive behaviour 118Sensory systems 118Central integrative systems 118Motor systems 120Endocrines and secondary sex characteristics 120Effect of social interactions on reproductive endocrines 121Role of pheromones in reproductive interactions 122Endocrine-disrupting chemicals 123Conclusions 1256 Environmental control of reproduction 127Introduction 128Seasonal patterns of reproduction 128Global seasonal patterns 128Ultimate factors 129Proximate factors 130Spawning seasonality in freshwater systems 130Spawning seasonality in marine species 132Lunar-related reproductive cycles 134Intertidal spawners 135Coral reef spawners 135Freshwater spawners 136Diel reproductive cycles 136Multiple environmental cycles 136Endogenous cycles and reproductive activity 137Proximate factors 138Photoperiod and temperature 139Lunar-related and tidal cycles 143Rainfall, flooding and other cues 143Social factors as cues 144Stress and reproduction 144Neuroendocrine mechanisms mediating environmental effects 147Food as a proximate factor 150Feeding and capital breeders 151Feeding and income breeders 154Feeding and skipped spawnings 155Physiological basis for the effect of feeding on reproduction 156Conclusions 1577 Migration, territoriality and spawning site selection 159Introduction 160Migration 160Terms and definitions 161Migratory patterns associated with reproduction 162Adaptive significance of reproductive migrations 164Energetic costs of migration 166Survival costs of migration 167Physiological adaptations associated with migration 167Mechanisms of orientation and homing during spawning migrations 168Control of timing of migration 171Human impacts on migration 172Territoriality 173The concept of territoriality and definitions 174Adaptive significance of territoriality 174Cost–benefit analysis and evolutionary stable strategy 174Aggression and territoriality 175Patterns of occurrence of territoriality 176Establishment and maintenance of territories 180Physiological basis of territoriality 182Energetic costs of territoriality 184Genetic basis of territoriality 184Territoriality and population dynamics 185Spawning site selection 188Sites for reproduction 188Site selection without habitat modification 188Site preparation 189Site excavation 189Site preparation and young brooded elsewhere 190Site preparation and eggs buried 191Excavation of shelters and tunnels 192Nest construction 193Terrestrial spawning sites 195Cues used in spawning site selection 197Spawning symbioses 197Conclusions 2008 Mating systems and sexual selection 201Introduction 202Classification of mating systems 203Sexual selection and theories of mate choice 204Direct intersexual selection 205Indirect intersexual selection 206Good genes 207Compatible genes 207Antagonistic genes 209Arbitrary mate choice 210Fisherian selection 210Sensory bias 211Male mate choice and female ornaments 212Mutual mate choice and the ‘double process’ of sexual selection 214Mate choice copying 216Intrasexual selection 216The interaction of intra- and intersexual selection 218The strength of sexual selection and the operational sex ratio 218Courtship 219Functions of courtship 219Mate identification and attraction 219Mate stimulation, mating synchrony and appeasement 220Maintenance of pair bonds 221Ornaments and signals in courtship 221Visual 222Body size 222Morphological trait size 223Colour 224UV reflectance 224Display behaviour 225Auditory 226Olfaction and gustation 227Tactile, electrical and other cues 228Multiple cues 228Sexual signals and reproductive isolation 229Fertilisation 231Sperm competition 231Cost of sperm production 231Risk and intensity of sperm competition 232Sperm competition avoidance 234Pre-oviposition ejaculation 234Seminal fluid composition 235Sperm capacitation and motility 235Ejaculate size and ejaculation frequency 236External fertilisation 237Internal fertilisation 238Quasi-internal fertilisation 239Male mating polymorphisms 240Causal factors 242Floaters, sneakers, streakers, pirates, parasites, fighters and the bourgeoisie 243Male mating polymorphism and sperm competition 244Guarder male response to non-guarders 244Female response to alternative male mating phenotypes 244Female mating polymorphisms 245Sexual conflict 246Phylogenetic history and mating system evolution 2499 Parental care 251Introduction 252Definition of care 252Distribution of parental care 252Modes of parental care 253Protection from predators and disturbance 253Protection from hypoxia and desiccation 253Protection from pathogens 255Nutrition 256External bearing 257Internal bearing 261Costs of parental care 262Cost to parental survival 262Cost to number of offspring produced 264Sex role reversal and parental care 265Phenotypic plasticity in care 266Number of offspring 266Offspring age 267Offspring quality 267Food availability 268Parental age 268Availability of alternative mates 268Helper care systems 269Alloparental care 270Sneaky mating and egg dumping 270Egg stealing, kidnapping and nest takeovers 271Brood parasitism 271Cooperative care 273Misdirected care 274Filial cannibalism 274Evolution of parental care 27610 Unusual reproductive modes 281Introduction 282Viviparity in teleosts 282Systematic distribution of viviparity in teleosts 283Internal fertilisation 283Lecithotrophy and matrotrophy 286Ovarian modifications in viviparity 287Testis modification in viviparity 289Gestation 289Timing of events 289Embryonic nutrition 290Superfetation 291Endocrine control of viviparity 292Evolution of viviparity and matrotrophy 292Male viviparity in the Syngnathidae 295Unisexual teleosts 298Systematic distribution of unisexual biotypes in teleosts 299Cyprinodontiformes 300Poeciliid unisexuals 300Poeciliid unisexuals: Poecilia formosa 300Poeciliid unisexuals: Poeciliopsis 300Atheriniformes 301Cypriniformes 301Phoxinus eos-neogaeus 302Tropidophoxinellus alburnoides 302Carassius gibelio 302Cobitis 303Problems posed by unisexual teleosts 303Developmental 303Evolutionary implications 304Hermaphroditism in teleosts 305Phylogenetic distribution of hermaphroditism in teleosts 306Gonadal organisation in hermaphroditic fishes 308Endocrinology of reproduction in hermaphroditic species 309Simultaneous hermaphrodites 312Self-fertilisation 312Cross-fertilisation 313Sequential hermaphrodites 313Protogyny 314Protandry 315Serial change 315Adaptive significance of hermaphroditism 316Why are hermaphrodites found in teleosts? 319Deep-sea anglerfish dwarf males: quasi-hermaphroditism 32011 Reproduction and life-history evolution 323Introduction 324Life-history variables 324Size and age at maturity 325Fecundity and clutch size 325Egg and offspring size 327Incubation time 331Reproductive lifespan 333Parental care 333Other life-history traits 333Basic concepts in life-history theory 334Trade-offs 334Costs of reproduction 334Measuring the costs of reproduction 335Evidence of costs of reproduction 336Reproductive effort 338Demographic models of life-history evolution 339Measures of fitness 340Predictions of demographic models 341Empirical studies of life-history evolution 342Habitat invasion 344Environmental gradients 346Habitat templet model 347Dimensionless variables and life-history invariants 350Plasticity of life-history traits 351Constraints on life-history evolution 353Alternative life-history strategies 354Conclusions 35612 Reproduction, fisheries and aquaculture 357Introduction 358Reproduction and fisheries management 358Stock–recruitment relationships and reproduction 358Reproductive traits and population dynamics 362Age at maturity 362Fecundity 363Mode of reproduction and susceptibility to overexploitation 364The impact of fisheries on reproductive traits 365Age and size structure 365Population sex ratio 368Population size and density 370Fisheries-induced evolution 372Control of reproduction in aquaculture 373Facilitating reproduction 374Environmental manipulation 374Endocrine manipulation 375Sexual dysfunction in cultivated fishes 376Sexual dysfunction in females 376Sexual dysfunction in males 378Preservation of gametes 378Suppressing reproduction 379Monosex production 379Endocrine manipulation 380Production of female monosex lines 381Production of male monosex lines 382Gynogenesis 383Androgenesis 385Hybridisation 385Sterilisation 386Mechanical, immunological, chemical and irradiation techniques 386Hybridisation 387Polyploidy 38713 Coda 389References 391Systematic index 451Subject index 463

Dr Robert 'Bob' Wootton was Reader in Ecology at the Institute of Biological, Environmental and Rural Sciences, Aberystwyth University, UK. Bob's research addressed the ecology, behaviour and evolutionary biology of fishes.Dr Carl Smith is a Lecturer in the School of Biology at the University of St Andrews, UK and also holds a research position at the Institute of Vertebrate Biology of the Czech Academy of Sciences in Brno, Czech Republic. Carl's research focuses on mating system evolution in fishes.

“I was very impressed by this book, even if it does not delve deeply into the mathematics of evolutionary theory applied to the topics.”  (Environ Biol Fish, 1 September 2015)

• Preface xvii Acknowledgements xixAbbreviations and acronyms xxi1 Introduction 1Introduction 1Reproductive modes of the teleosts 2Gender systems of teleosts 2Spawning dynamics 4Modes of fertilisation 5Mating systems 5Secondary sexual characteristics 6Parental care 6Reproductive guilds 6Reproductive diversity in teleosts: an explanatory framework 6Phylogenetic relationships of the teleosts 8Life-history theory 9Aims of the volume 112 Sex determination 13Introduction 13Genotypic sex determination 14Monofactorial genotypic sex determination 14Multifactorial genotypic sex determination 20Polyfactorial (polygenic) genotypic sex determination 20Environmental sex determination 21Evolution of sex-determining mechanisms 263 Sex differentiation 31Introduction 31Embryology of the gonads 32Origin and migration of primordial germ cells 33Sex differentiation in the gonadal anlagen 35Initial differentiation of the gonads 36Initial differentiation of ovaries 38Initial differentiation of testes 38Genetic control of early gonadal differentiation 40Synthesis 424 Gametogenesis 45Introduction 46Structure of ovaries 48Gross morphology of the ovaries 48Development of female gonoducts 51Oogenesis 52Oogonial proliferation and oogonial nest formation 53Chromatin nucleolus stage 53Primary growth 54Secondary growth – vitellogenesis 55Vitellogenin and the zona pellucida proteins 56Polarity of the oocyte 58Oocyte maturation 58Ovulation 60Atresia 60Fertilisation 62Genetic control of oogenesis 64Dynamics of oocyte development 65Spawning dynamics and fecundity 66Structure of the testes 66Gross morphology of a testis 66Development of male gonoducts 69Accessory structures associated with the testes and gonoducts 70Spermatogenesis 71Spermatocysts 71Sertoli cells 71Stages of spermatogenesis within a spermatogenic spermatocyst 73Differentiated spermatogonia 73Primary spermatocysts 73Secondary spermatocysts 73Spermatids 74Spermiation and capacitation 75Types of spermatozoa 76Structure and phylogenetic distribution 76Spermatozoa viability 77Genetic control of spermatogenesis 79Dynamics of spermatogenesis 79Quantitative analysis of sperm production 805 Endocrinology of reproduction 81Introduction 82Brain–pituitary–gonad reproductive axis in vertebrates 83Levels of analysis 85Gonadal steroids 86Sex steroid synthesis 86Sex steroid receptors 87Endocrine control of oogenesis 88Oogonial proliferation and primary growth 88Secondary growth: vitellogenesis 89Maturation and ovulation 90Endocrine control of oogenesis in species with batch spawning 91Endocrine control of spermatogenesis 93Spermatogonial proliferation 95Initiation of meiosis and formation of spermatocysts 95Spermiation 96Patterns of spawning in relation to spermatogenesis 96The pituitary and the gonadotrophins 98Brain–pituitary relationship 98Gonadotrophs and gonadotrophins 99Gonadotrophin receptors 100Hypothalamic control of the pituitary 101Gonadotrophin-releasing hormone 101Functions of gonadotrophin-releasing hormone 102Gonadotrophin-releasing hormone receptors 102Distribution of gonadotrophin-releasing hormone neurons in the brain 103Role of dopamine as an inhibitor of reproduction 104Role of other brain factors in the control of gonadotrophs 105Feedback control in the brain–pituitary–gonad reproductive axis 106A provisional model for the control of the developmental dynamics of ovaries and testes 109Puberty in teleosts 109Physiological control of puberty 110Interactions between growth and reproductive axes 113Endocrines, behaviour and secondary sexual characteristics 116Four model species 117Role of endocrines in regulation of reproductive behaviour 118Sensory systems 118Central integrative systems 118Motor systems 120Endocrines and secondary sex characteristics 120Effect of social interactions on reproductive endocrines 121Role of pheromones in reproductive interactions 122Endocrine-disrupting chemicals 123Conclusions 1256 Environmental control of reproduction 127Introduction 128Seasonal patterns of reproduction 128Global seasonal patterns 128Ultimate factors 129Proximate factors 130Spawning seasonality in freshwater systems 130Spawning seasonality in marine species 132Lunar-related reproductive cycles 134Intertidal spawners 135Coral reef spawners 135Freshwater spawners 136Diel reproductive cycles 136Multiple environmental cycles 136Endogenous cycles and reproductive activity 137Proximate factors 138Photoperiod and temperature 139Lunar-related and tidal cycles 143Rainfall, flooding and other cues 143Social factors as cues 144Stress and reproduction 144Neuroendocrine mechanisms mediating environmental effects 147Food as a proximate factor 150Feeding and capital breeders 151Feeding and income breeders 154Feeding and skipped spawnings 155Physiological basis for the effect of feeding on reproduction 156Conclusions 1577 Migration, territoriality and spawning site selection 159Introduction 160Migration 160Terms and definitions 161Migratory patterns associated with reproduction 162Adaptive significance of reproductive migrations 164Energetic costs of migration 166Survival costs of migration 167Physiological adaptations associated with migration 167Mechanisms of orientation and homing during spawning migrations 168Control of timing of migration 171Human impacts on migration 172Territoriality 173The concept of territoriality and definitions 174Adaptive significance of territoriality 174Cost–benefit analysis and evolutionary stable strategy 174Aggression and territoriality 175Patterns of occurrence of territoriality 176Establishment and maintenance of territories 180Physiological basis of territoriality 182Energetic costs of territoriality 184Genetic basis of territoriality 184Territoriality and population dynamics 185Spawning site selection 188Sites for reproduction 188Site selection without habitat modification 188Site preparation 189Site excavation 189Site preparation and young brooded elsewhere 190Site preparation and eggs buried 191Excavation of shelters and tunnels 192Nest construction 193Terrestrial spawning sites 195Cues used in spawning site selection 197Spawning symbioses 197Conclusions 2008 Mating systems and sexual selection 201Introduction 202Classification of mating systems 203Sexual selection and theories of mate choice 204Direct intersexual selection 205Indirect intersexual selection 206Good genes 207Compatible genes 207Antagonistic genes 209Arbitrary mate choice 210Fisherian selection 210Sensory bias 211Male mate choice and female ornaments 212Mutual mate choice and the ‘double process’ of sexual selection 214Mate choice copying 216Intrasexual selection 216The interaction of intra- and intersexual selection 218The strength of sexual selection and the operational sex ratio 218Courtship 219Functions of courtship 219Mate identification and attraction 219Mate stimulation, mating synchrony and appeasement 220Maintenance of pair bonds 221Ornaments and signals in courtship 221Visual 222Body size 222Morphological trait size 223Colour 224UV reflectance 224Display behaviour 225Auditory 226Olfaction and gustation 227Tactile, electrical and other cues 228Multiple cues 228Sexual signals and reproductive isolation 229Fertilisation 231Sperm competition 231Cost of sperm production 231Risk and intensity of sperm competition 232Sperm competition avoidance 234Pre-oviposition ejaculation 234Seminal fluid composition 235Sperm capacitation and motility 235Ejaculate size and ejaculation frequency 236External fertilisation 237Internal fertilisation 238Quasi-internal fertilisation 239Male mating polymorphisms 240Causal factors 242Floaters, sneakers, streakers, pirates, parasites, fighters and the bourgeoisie 243Male mating polymorphism and sperm competition 244Guarder male response to non-guarders 244Female response to alternative male mating phenotypes 244Female mating polymorphisms 245Sexual conflict 246Phylogenetic history and mating system evolution 2499 Parental care 251Introduction 252Definition of care 252Distribution of parental care 252Modes of parental care 253Protection from predators and disturbance 253Protection from hypoxia and desiccation 253Protection from pathogens 255Nutrition 256External bearing 257Internal bearing 261Costs of parental care 262Cost to parental survival 262Cost to number of offspring produced 264Sex role reversal and parental care 265Phenotypic plasticity in care 266Number of offspring 266Offspring age 267Offspring quality 267Food availability 268Parental age 268Availability of alternative mates 268Helper care systems 269Alloparental care 270Sneaky mating and egg dumping 270Egg stealing, kidnapping and nest takeovers 271Brood parasitism 271Cooperative care 273Misdirected care 274Filial cannibalism 274Evolution of parental care 27610 Unusual reproductive modes 281Introduction 282Viviparity in teleosts 282Systematic distribution of viviparity in teleosts 283Internal fertilisation 283Lecithotrophy and matrotrophy 286Ovarian modifications in viviparity 287Testis modification in viviparity 289Gestation 289Timing of events 289Embryonic nutrition 290Superfetation 291Endocrine control of viviparity 292Evolution of viviparity and matrotrophy 292Male viviparity in the Syngnathidae 295Unisexual teleosts 298Systematic distribution of unisexual biotypes in teleosts 299Cyprinodontiformes 300Poeciliid unisexuals 300Poeciliid unisexuals: Poecilia formosa 300Poeciliid unisexuals: Poeciliopsis 300Atheriniformes 301Cypriniformes 301Phoxinus eos-neogaeus 302Tropidophoxinellus alburnoides 302Carassius gibelio 302Cobitis 303Problems posed by unisexual teleosts 303Developmental 303Evolutionary implications 304Hermaphroditism in teleosts 305Phylogenetic distribution of hermaphroditism in teleosts 306Gonadal organisation in hermaphroditic fishes 308Endocrinology of reproduction in hermaphroditic species 309Simultaneous hermaphrodites 312Self-fertilisation 312Cross-fertilisation 313Sequential hermaphrodites 313Protogyny 314Protandry 315Serial change 315Adaptive significance of hermaphroditism 316Why are hermaphrodites found in teleosts? 319Deep-sea anglerfish dwarf males: quasi-hermaphroditism 32011 Reproduction and life-history evolution 323Introduction 324Life-history variables 324Size and age at maturity 325Fecundity and clutch size 325Egg and offspring size 327Incubation time 331Reproductive lifespan 333Parental care 333Other life-history traits 333Basic concepts in life-history theory 334Trade-offs 334Costs of reproduction 334Measuring the costs of reproduction 335Evidence of costs of reproduction 336Reproductive effort 338Demographic models of life-history evolution 339Measures of fitness 340Predictions of demographic models 341Empirical studies of life-history evolution 342Habitat invasion 344Environmental gradients 346Habitat templet model 347Dimensionless variables and life-history invariants 350Plasticity of life-history traits 351Constraints on life-history evolution 353Alternative life-history strategies 354Conclusions 35612 Reproduction, fisheries and aquaculture 357Introduction 358Reproduction and fisheries management 358Stock–recruitment relationships and reproduction 358Reproductive traits and population dynamics 362Age at maturity 362Fecundity 363Mode of reproduction and susceptibility to overexploitation 364The impact of fisheries on reproductive traits 365Age and size structure 365Population sex ratio 368Population size and density 370Fisheries-induced evolution 372Control of reproduction in aquaculture 373Facilitating reproduction 374Environmental manipulation 374Endocrine manipulation 375Sexual dysfunction in cultivated fishes 376Sexual dysfunction in females 376Sexual dysfunction in males 378Preservation of gametes 378Suppressing reproduction 379Monosex production 379Endocrine manipulation 380Production of female monosex lines 381Production of male monosex lines 382Gynogenesis 383Androgenesis 385Hybridisation 385Sterilisation 386Mechanical, immunological, chemical and irradiation techniques 386Hybridisation 387Polyploidy 38713 Coda 389References 391Systematic index 451Subject index 463

Dr Robert 'Bob' Wootton was Reader in Ecology at the Institute of Biological, Environmental and Rural Sciences, Aberystwyth University, UK. Bob's research addressed the ecology, behaviour and evolutionary biology of fishes.Dr Carl Smith is a Lecturer in the School of Biology at the University of St Andrews, UK and also holds a research position at the Institute of Vertebrate Biology of the Czech Academy of Sciences in Brno, Czech Republic. Carl's research focuses on mating system evolution in fishes.

“I was very impressed by this book, even if it does not delve deeply into the mathematics of evolutionary theory applied to the topics.”  (Environ Biol Fish, 1 September 2015)

• Preface xvii Acknowledgements xixAbbreviations and acronyms xxi1 Introduction 1Introduction 1Reproductive modes of the teleosts 2Gender systems of teleosts 2Spawning dynamics 4Modes of fertilisation 5Mating systems 5Secondary sexual characteristics 6Parental care 6Reproductive guilds 6Reproductive diversity in teleosts: an explanatory framework 6Phylogenetic relationships of the teleosts 8Life-history theory 9Aims of the volume 112 Sex determination 13Introduction 13Genotypic sex determination 14Monofactorial genotypic sex determination 14Multifactorial genotypic sex determination 20Polyfactorial (polygenic) genotypic sex determination 20Environmental sex determination 21Evolution of sex-determining mechanisms 263 Sex differentiation 31Introduction 31Embryology of the gonads 32Origin and migration of primordial germ cells 33Sex differentiation in the gonadal anlagen 35Initial differentiation of the gonads 36Initial differentiation of ovaries 38Initial differentiation of testes 38Genetic control of early gonadal differentiation 40Synthesis 424 Gametogenesis 45Introduction 46Structure of ovaries 48Gross morphology of the ovaries 48Development of female gonoducts 51Oogenesis 52Oogonial proliferation and oogonial nest formation 53Chromatin nucleolus stage 53Primary growth 54Secondary growth – vitellogenesis 55Vitellogenin and the zona pellucida proteins 56Polarity of the oocyte 58Oocyte maturation 58Ovulation 60Atresia 60Fertilisation 62Genetic control of oogenesis 64Dynamics of oocyte development 65Spawning dynamics and fecundity 66Structure of the testes 66Gross morphology of a testis 66Development of male gonoducts 69Accessory structures associated with the testes and gonoducts 70Spermatogenesis 71Spermatocysts 71Sertoli cells 71Stages of spermatogenesis within a spermatogenic spermatocyst 73Differentiated spermatogonia 73Primary spermatocysts 73Secondary spermatocysts 73Spermatids 74Spermiation and capacitation 75Types of spermatozoa 76Structure and phylogenetic distribution 76Spermatozoa viability 77Genetic control of spermatogenesis 79Dynamics of spermatogenesis 79Quantitative analysis of sperm production 805 Endocrinology of reproduction 81Introduction 82Brain–pituitary–gonad reproductive axis in vertebrates 83Levels of analysis 85Gonadal steroids 86Sex steroid synthesis 86Sex steroid receptors 87Endocrine control of oogenesis 88Oogonial proliferation and primary growth 88Secondary growth: vitellogenesis 89Maturation and ovulation 90Endocrine control of oogenesis in species with batch spawning 91Endocrine control of spermatogenesis 93Spermatogonial proliferation 95Initiation of meiosis and formation of spermatocysts 95Spermiation 96Patterns of spawning in relation to spermatogenesis 96The pituitary and the gonadotrophins 98Brain–pituitary relationship 98Gonadotrophs and gonadotrophins 99Gonadotrophin receptors 100Hypothalamic control of the pituitary 101Gonadotrophin-releasing hormone 101Functions of gonadotrophin-releasing hormone 102Gonadotrophin-releasing hormone receptors 102Distribution of gonadotrophin-releasing hormone neurons in the brain 103Role of dopamine as an inhibitor of reproduction 104Role of other brain factors in the control of gonadotrophs 105Feedback control in the brain–pituitary–gonad reproductive axis 106A provisional model for the control of the developmental dynamics of ovaries and testes 109Puberty in teleosts 109Physiological control of puberty 110Interactions between growth and reproductive axes 113Endocrines, behaviour and secondary sexual characteristics 116Four model species 117Role of endocrines in regulation of reproductive behaviour 118Sensory systems 118Central integrative systems 118Motor systems 120Endocrines and secondary sex characteristics 120Effect of social interactions on reproductive endocrines 121Role of pheromones in reproductive interactions 122Endocrine-disrupting chemicals 123Conclusions 1256 Environmental control of reproduction 127Introduction 128Seasonal patterns of reproduction 128Global seasonal patterns 128Ultimate factors 129Proximate factors 130Spawning seasonality in freshwater systems 130Spawning seasonality in marine species 132Lunar-related reproductive cycles 134Intertidal spawners 135Coral reef spawners 135Freshwater spawners 136Diel reproductive cycles 136Multiple environmental cycles 136Endogenous cycles and reproductive activity 137Proximate factors 138Photoperiod and temperature 139Lunar-related and tidal cycles 143Rainfall, flooding and other cues 143Social factors as cues 144Stress and reproduction 144Neuroendocrine mechanisms mediating environmental effects 147Food as a proximate factor 150Feeding and capital breeders 151Feeding and income breeders 154Feeding and skipped spawnings 155Physiological basis for the effect of feeding on reproduction 156Conclusions 1577 Migration, territoriality and spawning site selection 159Introduction 160Migration 160Terms and definitions 161Migratory patterns associated with reproduction 162Adaptive significance of reproductive migrations 164Energetic costs of migration 166Survival costs of migration 167Physiological adaptations associated with migration 167Mechanisms of orientation and homing during spawning migrations 168Control of timing of migration 171Human impacts on migration 172Territoriality 173The concept of territoriality and definitions 174Adaptive significance of territoriality 174Cost–benefit analysis and evolutionary stable strategy 174Aggression and territoriality 175Patterns of occurrence of territoriality 176Establishment and maintenance of territories 180Physiological basis of territoriality 182Energetic costs of territoriality 184Genetic basis of territoriality 184Territoriality and population dynamics 185Spawning site selection 188Sites for reproduction 188Site selection without habitat modification 188Site preparation 189Site excavation 189Site preparation and young brooded elsewhere 190Site preparation and eggs buried 191Excavation of shelters and tunnels 192Nest construction 193Terrestrial spawning sites 195Cues used in spawning site selection 197Spawning symbioses 197Conclusions 2008 Mating systems and sexual selection 201Introduction 202Classification of mating systems 203Sexual selection and theories of mate choice 204Direct intersexual selection 205Indirect intersexual selection 206Good genes 207Compatible genes 207Antagonistic genes 209Arbitrary mate choice 210Fisherian selection 210Sensory bias 211Male mate choice and female ornaments 212Mutual mate choice and the ‘double process’ of sexual selection 214Mate choice copying 216Intrasexual selection 216The interaction of intra- and intersexual selection 218The strength of sexual selection and the operational sex ratio 218Courtship 219Functions of courtship 219Mate identification and attraction 219Mate stimulation, mating synchrony and appeasement 220Maintenance of pair bonds 221Ornaments and signals in courtship 221Visual 222Body size 222Morphological trait size 223Colour 224UV reflectance 224Display behaviour 225Auditory 226Olfaction and gustation 227Tactile, electrical and other cues 228Multiple cues 228Sexual signals and reproductive isolation 229Fertilisation 231Sperm competition 231Cost of sperm production 231Risk and intensity of sperm competition 232Sperm competition avoidance 234Pre-oviposition ejaculation 234Seminal fluid composition 235Sperm capacitation and motility 235Ejaculate size and ejaculation frequency 236External fertilisation 237Internal fertilisation 238Quasi-internal fertilisation 239Male mating polymorphisms 240Causal factors 242Floaters, sneakers, streakers, pirates, parasites, fighters and the bourgeoisie 243Male mating polymorphism and sperm competition 244Guarder male response to non-guarders 244Female response to alternative male mating phenotypes 244Female mating polymorphisms 245Sexual conflict 246Phylogenetic history and mating system evolution 2499 Parental care 251Introduction 252Definition of care 252Distribution of parental care 252Modes of parental care 253Protection from predators and disturbance 253Protection from hypoxia and desiccation 253Protection from pathogens 255Nutrition 256External bearing 257Internal bearing 261Costs of parental care 262Cost to parental survival 262Cost to number of offspring produced 264Sex role reversal and parental care 265Phenotypic plasticity in care 266Number of offspring 266Offspring age 267Offspring quality 267Food availability 268Parental age 268Availability of alternative mates 268Helper care systems 269Alloparental care 270Sneaky mating and egg dumping 270Egg stealing, kidnapping and nest takeovers 271Brood parasitism 271Cooperative care 273Misdirected care 274Filial cannibalism 274Evolution of parental care 27610 Unusual reproductive modes 281Introduction 282Viviparity in teleosts 282Systematic distribution of viviparity in teleosts 283Internal fertilisation 283Lecithotrophy and matrotrophy 286Ovarian modifications in viviparity 287Testis modification in viviparity 289Gestation 289Timing of events 289Embryonic nutrition 290Superfetation 291Endocrine control of viviparity 292Evolution of viviparity and matrotrophy 292Male viviparity in the Syngnathidae 295Unisexual teleosts 298Systematic distribution of unisexual biotypes in teleosts 299Cyprinodontiformes 300Poeciliid unisexuals 300Poeciliid unisexuals: Poecilia formosa 300Poeciliid unisexuals: Poeciliopsis 300Atheriniformes 301Cypriniformes 301Phoxinus eos-neogaeus 302Tropidophoxinellus alburnoides 302Carassius gibelio 302Cobitis 303Problems posed by unisexual teleosts 303Developmental 303Evolutionary implications 304Hermaphroditism in teleosts 305Phylogenetic distribution of hermaphroditism in teleosts 306Gonadal organisation in hermaphroditic fishes 308Endocrinology of reproduction in hermaphroditic species 309Simultaneous hermaphrodites 312Self-fertilisation 312Cross-fertilisation 313Sequential hermaphrodites 313Protogyny 314Protandry 315Serial change 315Adaptive significance of hermaphroditism 316Why are hermaphrodites found in teleosts? 319Deep-sea anglerfish dwarf males: quasi-hermaphroditism 32011 Reproduction and life-history evolution 323Introduction 324Life-history variables 324Size and age at maturity 325Fecundity and clutch size 325Egg and offspring size 327Incubation time 331Reproductive lifespan 333Parental care 333Other life-history traits 333Basic concepts in life-history theory 334Trade-offs 334Costs of reproduction 334Measuring the costs of reproduction 335Evidence of costs of reproduction 336Reproductive effort 338Demographic models of life-history evolution 339Measures of fitness 340Predictions of demographic models 341Empirical studies of life-history evolution 342Habitat invasion 344Environmental gradients 346Habitat templet model 347Dimensionless variables and life-history invariants 350Plasticity of life-history traits 351Constraints on life-history evolution 353Alternative life-history strategies 354Conclusions 35612 Reproduction, fisheries and aquaculture 357Introduction 358Reproduction and fisheries management 358Stock–recruitment relationships and reproduction 358Reproductive traits and population dynamics 362Age at maturity 362Fecundity 363Mode of reproduction and susceptibility to overexploitation 364The impact of fisheries on reproductive traits 365Age and size structure 365Population sex ratio 368Population size and density 370Fisheries-induced evolution 372Control of reproduction in aquaculture 373Facilitating reproduction 374Environmental manipulation 374Endocrine manipulation 375Sexual dysfunction in cultivated fishes 376Sexual dysfunction in females 376Sexual dysfunction in males 378Preservation of gametes 378Suppressing reproduction 379Monosex production 379Endocrine manipulation 380Production of female monosex lines 381Production of male monosex lines 382Gynogenesis 383Androgenesis 385Hybridisation 385Sterilisation 386Mechanical, immunological, chemical and irradiation techniques 386Hybridisation 387Polyploidy 38713 Coda 389References 391Systematic index 451Subject index 463