Shrimps : Their Diversity, Intriguing Adaptations and Varied Lifestyles

This book explores the biology of decapod shrimps, a group of animals known to most people as a nutritious and tasty food item. Shrimps are amazingly diverse in size, shape, coloration, behavior and natural history. Shrimp fisheries and aquaculture are a vital part of the USA and world economies. These crustaceans are key ecological and food-web components of marine and freshwater habitats. The book synthesizes information on the taxonomic and ecological diversity of shrimps, the structure and function of shrimp anatomy, antifouling adaptations, coloration and camouflage, reproductive biology, sexual systems, mating systems and behavior, life history strategies, symbioses between shrimps and other organisms, shrimp fisheries and aquaculture, as well as the evolution and phylogeny of shrimps. All chapters are written within an adaptational and evolutionary perspective. Important questions about shrimp biology are asked, and hypotheses for testing in future research are proposed. The book is spiced up with personal anecdotes and observations from the author’s research experiences. This book is intended as a comprehensive reference, a “go to” book about the biology of shrimps. The text is scientifically rigorous but written in a style intended for a varied readership. Thus, the book is a valuable resource for budding or working research scientists such as zoologists, aquatic biologists, fisheries and aquaculture professionals, as well as amateur naturalists, aquarium hobbyists and interested laypersons. As the saying goes, “a picture is worth a thousand words,” so that the book is amply illustrated with figures and diagrams.The numerous color plates, composed of photos contributed by expert colleagues, make the world of shrimps come alive. Preface Contents Chapter 1: The Shrimps: An Overview 1.1 What Are Decapod Shrimps? 1.2 Highly Adapted Swimmers 1.3 Reduction of the Caridoid Facies in the Decapoda 1.4 Systematic Position of the Decapod Shrimps Within the Crustacea 1.5 Biological Success of Shrimp Groups 1.6 Which Group of Shrimps? References Chapter 2: Structure and Function 2.1 From Stem to Stern 2.2 Cephalothorax and Carapace 2.3 Gills 2.4 Eyes 2.5 Appendages 2.6 Antennules 2.7 Second Antennae 2.8 Mouthparts 2.9 Maxillipeds 2.10 Pereopods 2.10.1 Chelae: Shrimp Pincers 2.10.2 Walking Legs 2.11 The Abdomen and Its Appendages References Chapter 3: The Families of Decapod Shrimps 3.1 Taxonomic Considerations 3.2 Suborder Dendrobranchiata 3.3 Superfamily Penaeoidea 3.3.1 Aristeidae 3.3.2 Benthesicymidae 3.3.3 Penaeidae 3.3.4 Sicyoniidae 3.3.5 Solenoceridae 3.4 Superfamily Sergestoidea 3.4.1 Luciferidae 3.4.2 Sergestidae 3.5 Infraorder Caridea 3.6 Superfamily Alpheoidea 3.6.1 Alpheidae 3.6.2 Hippolytidae (s.l.) Families 3.6.3 Barbouriidae 3.6.4 Bythocarididae 3.6.5 Hippolytidae (s.s.) 3.6.6 Lysmatidae 3.6.7 Merguiidae 3.6.8 Thoridae 3.6.9 Ogyrididae 3.7 Superfamily Atyoidea: Atyidae 3.8 Superfamily Bresilioidea 3.8.1 Family Agostocarididae 3.8.2 Family Alvinocarididae 3.8.3 Family Anchialocarididae 3.8.4 Family Bresiliidae (s.s.) 3.8.5 Disciadidae 3.8.6 Pseudochelidae 3.9 Superfamily Campylonotoidea 3.9.1 Bathypalaemonellidae 3.9.2 Family Campylonotidae 3.10 Superfamily Crangonoidea 3.10.1 Crangonidae 3.10.2 Glyphocrangonidae 3.11 Superfamily Nematocarcinoidea 3.11.1 Eugonatonotidae 3.11.2 Lipkiidae 3.11.3 Nematocarcinidae 3.11.4 Rhynchocinetidae 3.11.5 Xiphocarididae 3.12 Superfamily Oplophoroidea 3.12.1 Oplophoridae and Acanthephyridae 3.13 Superfamily Palaemonoidea 3.13.1 Desmocarididae 3.13.2 Euryrhynchidae 3.13.3 Palaemonidae (s.l.) 3.13.4 Typhlocarididae 3.14 Superfamily Pandaloidea 3.14.1 Pandalidae 3.14.2 Chlorotocellidae 3.15 Superfamily Pasiphaeoidea: Pasiphaeidae 3.16 Superfamily Physetocaridoidea: Physetocarididae 3.17 Superfamily Processoidea: Processidae 3.18 Superfamily Psalidopodoidea: Psalidopodidae 3.19 Superfamily Stylodactyloidea: Stylodactylidae 3.20 Amphionides reynaudii: Taxon Inquirendum or Pandalidae? 3.21 Infraorder Procarididea: Procarididae 3.22 Infraorder Stenopodidea 3.22.1 Stenopodidae 3.22.2 Spongicolidae 3.22.3 Macromaxillocarididae 3.23 Last Word References Chapter 4: Shrimp Hygiene: Antifouling Adaptations 4.1 Fouling and Antifouling Countermeasures 4.2 Grooming of Chemosensory Appendages 4.2.1 Antennular Grooming 4.2.2 Adaptive Value of Antennular Grooming 4.2.3 Cleaning of the Antennal Flagellum: The Pereopod 1 Carpal-Propodal Brush 4.3 Gill-Cleaning Mechanisms 4.3.1 Reversal and Filtering of the Respiratory Current 4.3.2 Setiferous Epipods 4.3.3 Epipod-Setobranch Complexes 4.3.4 Scaphognathite Gill Cleaning 4.3.5 Gill Cleaning by Chelipeds 4.3.6 Passive Versus Active Gill Cleaning 4.4 General Body Grooming 4.4.1 Morphology and Behavior 4.4.2 Adaptive Value of General Body Grooming 4.5 Embryo Care 4.6 Behavioral Aspects of Grooming 4.7 Other Antifouling Adaptations References Chapter 5: Coloration, Color Change, and Camouflage 5.1 Adaptive Value of Coloration 5.2 Types and Sources of Color 5.3 Color and Camouflage 5.4 Mechanisms of Color Change 5.5 Rapid Color Change 5.6 Day-Night Color Change 5.7 Morphological Color Change 5.8 Applied Value of Color Change Studies 5.9 Color Polymorphism and Camouflage 5.10 Warning Coloration 5.11 Color and Depth 5.12 Bioluminescence and Photophores 5.13 Defensive Bioluminescence References Chapter 6: Reproduction 6.1 Complexity of Shrimp Reproductive Morphology 6.1.1 Female Gonads: Ovaries 6.1.2 Male Gonads and Spermatophores 6.1.3 Male Gonadal System: Caridea, Stenopodidea, and Procarididea 6.1.4 Male Gonadal System: Penaeoidea and Sergestoidea 6.2 Secondary Sexual Characters: Genitalia 6.2.1 Genitalia of the Procarididea, Stenopodidea, and Caridea 6.2.2 Genitalia of Dendrobranchiata: Thelyca, Petasmata, and Appendices Masculinae 6.3 Mechanics of Copulation and Insemination 6.4 Eggs, Sperm, and Fertilization 6.4.1 Eggs 6.4.2 Sperm 6.4.3 Sperm Meets Egg: Fertilization 6.5 Temporal Cycles of Molting, Mating, Spawning, and Incubation 6.6 Spawning, Parental Care, and Breeding Dress References Chapter 7: Sexual Systems 7.1 Sexual Systems and Hermaphroditism 7.2 The Direction of Sex Change and the Size Advantage Model 7.3 Types and Variations in Sexual Systems 7.3.1 Pure Protandry 7.3.2 Partial Protandry with Primary and/or Early-Maturing Females 7.3.3 Partial Protandry with Primary Males, Primary Females 7.3.4 Partial Protandry with Primary Males 7.3.5 Protandric Simultaneous Hermaphroditism 7.3.6 Pure Simultaneous Hermaphroditism? 7.4 Identification of Protandrous Sexual Systems 7.4.1 Size-Frequency Distributions 7.4.2 Transitional Characters 7.4.3 Direct Observation of Sex Change 7.5 Taxonomic Distribution of Hermaphroditism in Shrimps 7.5.1 Dendrobranchiata (Sergestoidea and Penaeoidea) 7.5.2 Alpheidae 7.5.3 Atyidae 7.5.4 Campylonotidae 7.5.5 Crangonidae 7.5.6 Hippolytidae 7.5.7 Merguiidae 7.5.8 Pandalidae 7.5.9 Processidae 7.5.10 Rhynchocinetidae 7.5.11 Thoridae 7.5.12 Lysmatidae/Barbouriidae: Protandric Simultaneous Hermaphroditism 7.6 Role of the Androgenic Gland in Protandrous Sexual Systems 7.7 Environmentally Determined Sex Change (ESD) 7.8 Evolution of Protandrous Sexual Systems 7.8.1 Female-First or Male-First Origin of Protandrous Sexual Systems? 7.8.2 The Dendrobranchiate/Palaemonid Paradox 7.8.3 Evolution of Protandric Simultaneous Hermaphroditism (PSH) 7.8.4 Final Thoughts References Chapter 8: Mating Systems 8.1 Sexual Tactics and Strategies 8.2 Mate Attraction and Recognition 8.2.1 Visual Cues 8.2.2 Tactile Cues 8.2.3 Sex Pheromones 8.2.4 Olfactory Pheromones 8.2.5 Contact Sex Pheromones 8.2.6 Chemical Composition of Sex Pheromones 8.3 Shrimp Mating Systems 8.3.1 Pure Searching 8.3.2 Temporary Mate (Female) Guarding Systems 8.3.3 Search and Attend 8.3.4 Neighborhoods of Dominance (NOD) 8.3.5 Monogamy 8.4 Female Choice, Multiple Paternity and Sperm Competition References Chapter 9: Life Histories 9.1 What Is a Life History? 9.2 Embryonic Development and Hatching 9.3 Larval Development: Costs and Benefits 9.4 Extended Larval Development 9.4.1 Dendrobranchiata: Penaeoids and Sergestoids 9.4.2 Carideans 9.4.3 Stenopodideans 9.5 Abbreviated and Direct Larval Development 9.6 Developmental Strategies of Freshwater Species 9.6.1 Freshwater Abbreviated Larval Development and Direct Development Species 9.6.2 Amphidromous Species 9.6.3 Freshwater Extended Larval Development Species 9.7 Abbreviated and Direct Development in Marine Species 9.7.1 High-Latitude Species 9.7.2 Deep-Sea Species 9.7.3 ALD/DD in Shallow-Water Marine Species 9.8 Reproductive Investment, Output, and Fecundity 9.9 Variation of Fecundity with Female and Egg (Embryo) Size 9.10 Fecundity (Brood) Loss 9.11 Parasitic Castration by Bopyrid Isopods 9.12 Life Cycles, Seasonality, and Latitudinal Trends 9.12.1 Tropical and Warm-Temperate Shallow-Water Species: Dendrobranchiates 9.12.2 Tropical and Warm-Temperate Shallow-Water Species: Pleocyemates 9.12.3 Freshwater Carideans 9.12.4 Cool-Temperate and Boreal/Austral Species 9.12.5 Polar Species 9.13 Within Taxon Latitudinal Variation in Life-History Traits in Neritic Marine Species 9.14 Deep-Sea Species 9.15 The Life-History Continuum of Shrimps 9.16 Global Warming and Climatic Effects on Life-History Patterns References Chapter 10: Symbioses 10.1 Symbioses in Shrimps 10.2 Taxonomic and Geographic Distribution of Shrimp Symbioses 10.3 Common Adaptations of Symbiotic Shrimps 10.4 Sponge Dwellers 10.4.1 Caridean Sponge Associates 10.4.2 Spongicolid Sponge Associates 10.4.3 Social Organization in Synalpheus Sponge Inhabitants 10.5 Shrimps Associated with Cnidarians 10.5.1 Shrimp Associates of Corals and Related Colonial Anthozoans 10.5.2 Anemone Shrimps 10.5.3 Jellyfish Shrimps 10.6 Shrimp Residents of Echinodermata 10.6.1 Echinoidea (Sea Urchin) Associates 10.6.2 Crinoid Shrimps 10.6.3 Other Echinoderm-Shrimp Associations 10.6.4 Host Recognition and Fidelity 10.7 Molluscan Hostels 10.8 Ascidian Homes 10.9 Shrimp Associations with Burrow Dwellers and Other Crustacea 10.10 Shrimps as Hosts: Goby Fish-Alpheid Shrimp Symbioses 10.11 Shrimps That Clean Fishes: Cleaning Symbiosis 10.12 Evolution of Symbioses in Shrimps References Chapter 11: Fisheries and Aquaculture 11.1 A Desirable Food Product 11.2 Facts and Figures 11.3 Shrimp Fisheries and Aquaculture Production: Economic Value 11.4 Composition of the World ́s Shrimp Fisheries 11.5 Life-History Traits and Shrimp Productivity 11.6 Globally Important Fishery Species 11.6.1 Sergestidae 11.6.2 Penaeoidea 11.6.3 Caridea 11.7 Top Shrimp Fishing Countries 11.8 Fishing Techniques 11.8.1 Catchability and Behavior 11.8.2 Otter and Beam Trawls 11.8.3 Skimmer and Butterfly Nets 11.8.4 Fyke Nets and Traps (Pots) 11.9 Management of Shrimp Fisheries 11.9.1 Overfishing 11.9.2 Shrimp Fishery Forecasting and Regulation 11.10 Environmental Impacts of the Shrimp Fishing Industry 11.10.1 Bycatch and Discards 11.10.2 Trawling Disturbance of the Sea Bottom and Its Communities 11.10.3 Loss of Coastal Wetlands and Shrimp Fisheries 11.11 Scale and Growth of Shrimp Aquaculture 11.12 Penaeid Aquaculture 11.12.1 Some History 11.12.2 Principal Species 11.12.3 Principal Countries 11.12.4 Techniques 11.13 Freshwater (Macrobrachium) Shrimp Aquaculture 11.13.1 Principal Species 11.13.2 Principal Countries and Some History 11.13.3 Techniques 11.14 Shrimp Diseases 11.14.1 Penaeids 11.14.2 Macrobrachium Species 11.15 Biosecurity 11.16 Monosex Populations in Shrimp Aquaculture 11.17 Environmental Impacts of Shrimp Aquaculture 11.18 Ornamental Shrimps 11.19 Climate Change: Impacts on Fisheries and Aquaculture 11.20 Past, Present, and Future State of Shrimp Fisheries and Aquaculture References Chapter 12: Evolutionary History and Relationships of Shrimps 12.1 Shrimp Chronicles 12.2 The Fossil Record 12.3 Classification and Systematics 12.4 Morphological Characters and Their Evolutionary Polarities 12.5 Origin of Shrimps from a Decapod Ancestor 12.6 Relationships Among Major Shrimp Groups and Other Decapods 12.7 Chronological Estimates of Taxa Origins 12.8 Relationships Within the Major Shrimp Taxa 12.8.1 Dendrobranchiata 12.8.2 Sergestoidea 12.8.3 Penaeoidea 12.8.4 Caridea and Procarididea 12.8.5 Traditional Views of Caridean Relationships 12.8.6 Christoffersen ́s Cladistic Phylogeny 12.8.7 Molecular Tests of Caridean Phylogeny 12.8.8 Stenopodidea References Subject Index Taxonomic Index