Shrimps are animals known to most people as nutritious and tasty food items. Shrimp fisheries and aquaculture are a vital part of US and world economies. These crustaceans are key ecological and food-web components of marine and freshwater habitats. Shrimps occur in all oceanic depths and latitudes. The 4400 species of decapod shrimps are amazingly diverse in size, shape, coloration, and natural history. Perhaps, it is no wonder that I have spent the last 52 years of my life in the study, research, publication, and teaching of marine science and crustacean biology, with a focus on shrimps. It became obvious to me that a book covering all the shrimps was needed and desired by marine biologists, zoologists, and carcinologists (crustacean biologists), as well as by applied scientists and amateur naturalists. Thus, in 2017, I launched into the writing of this book to cover those crustaceans recognized by most persons as “shrimps,” i.e., the groups belonging to the crustacean order Decapoda: Dendrobranchiata (penaeoids and sergestoids), Caridea, Procarididea, and Stenopodidea. In this book, I cover important aspects of the organismal biology and natural history of shrimps: taxonomy, ecological diversity, structure and function of shrimp anatomy, antifouling adaptations, coloration and camouflage, reproductive biology, sexual systems, mating systems and behavior, life-history strategies, shrimp symbioses (parasitism, commensalism, and mutualism), and the evolution of shrimps. The book is based on past and present scientific literature (journal publications and books) as well as my own research and experience. I point out remaining questions about shrimps which need investigation and propose hypotheses for testing them. This book is envisioned as a comprehensive reference and source of information, a “go-to” book about shrimps. I have tried to write in a style appealing to a wide audience: advanced undergraduate and postgraduate students, academic scientists, and fisheries and aquaculture professionals, as well as amateur naturalists, aquarists, and interested laypersons.
I would not have been able to do this book without the help and support of numerous persons and key institutions along the way. I had the good fortune of being accepted in 1969 to the Scripps Institution of Oceanography (University of California, San Diego) where I received a doctorate in Marine Biology. William (Bill) A. Newman mentored my dissertation work. It was at his suggestion that I first investigated and sampled the nearby tidepools in search of possible research ideas. There I encountered fascinating species of shrimps, a group which became my life’s work. A class on crustaceans taught by Bill and Robert Hessler cemented my interest in the group. A year’s postdoctoral fellowship, sponsored by Raymond B. Manning, at the Smithsonian Institution’s National Museum of Natural History (NMNH), was another key influence. There I worked among giants of crustacean biology such as Fenner A. Chace, Jr., Brian Kensley, Austin Williams, Horton H. Hobbs, Jr., Thomas Bowman, and Isabel Pérez Farfante, among others.
My first tenure-track position was at the University of Puerto Rico, Río Piedras, as a marine biologist. The beauty and diversity of this island’s tropical reefs, seagrass meadows, cayos, and freshwater streams were the source of many productive studies. This research could not have been done without the help of many enthusiastic and agreeable UPR students. From there, I moved to the University of Louisiana at Lafayette, which became my permanent academic home. The support I have received from ULL throughout the years is gratefully recognized.
In the course of writing this book, I have received considerable help from students, professional colleagues, journal editors, and laypersons who responded to requests for information of various kinds. There are too many to list, but to all of you, my sincere thanks. I have been a member of the Crustacean Society since its inception in late 1979. Its contribution to my work and carcinology has been indispensable. I am indebted to TCS and its Journal of Crustacean Biology for its support of this book.
With the saying “a picture is worth a thousand words” in mind, I have included in the book 26 plates, each composed of several color photographs and/or grayscale micrographs. These plates were made possible in large part by generous image contributions from the colleagues listed below. Special credit and thanks are due Arthur Anker, a prominent carcinologist whose photography of shrimps is renowned. Many of the plate photos were contributed by Arthur. Marine biologist Zdeněk Ďuriš, professor of biology at the University of Ostrava, Czech Republic, kindly contributed several valuable photographs from his work in tropical habitats. T.Y. Chan, National Taiwan Ocean University, kindly sent several important shrimp photos taken during his extensive marine research, as did Alexander Vereshchaka, Institute of Oceanology of Russian Academy of Sciences. Other colleagues who provided much appreciated photographic contributions are Kevin Bourdon, Florent Charpin, David Darom, Robert Ditter, William Folsom, Christian Gloor, David Herasimtschuk, David Hillegas, Ivan Hinojosa, Watcharapong Hongjamrassilp, Gregory Jensen, Werner Klotz, Leslie Knoesen, David Knott, Richard Larocque, Benjamin Magana, Jeremy Monroe, Jun Ohtomi, João Pantaleão, Jennifer Rasch, Greg W. Rouse, Bernard Sainte-Marie, Tomomi Saito, Troy Sakihara, David Shale, Al Stokes, María Morales Suazo, Martin Thiel, Cris Harris Toro, Eva Tóth, Sebastian Vazquez, Fernando Vega-Villasante, Mary Wicksten, and Yusuke Yamada. The Louisiana Sea Grant Program’s Digital Images Collection was a good source of photographs about shrimp fishery techniques.
The WoRMS (World Registry of Marine Species) website has been invaluable for staying current on the taxonomy and classification of shrimps and other aquatic organisms. Vim De Cock of the WoRMS management team and Sammy De Grave and Charles H.J.M. Fransen (editors, Decapoda) were very helpful with questions and data on species numbers. Many thanks to Rose Gulledge (NMNH) for obtaining copies of hard-to-get journal articles and illustrations. It has been a pleasure working with Springer Nature senior editor Éva Loerinczi and the production teams led by Bibhuti Sharma and Jana Yagnavaragan. Their advice, patience, and help of all kinds have been invaluable—many thanks!!
I will finish the Preface with the most important acknowledgment of all: to my wife Lydia, whose constant encouragement, support, and patience were essential during the production of this book. Lydia, I owe you big time; lots of love, Ray

1 The Shrimps: An Overview 1

1.1 What Are Decapod Shrimps? 1

1.2 Highly Adapted Swimmers 3

1.3 Reduction of the Caridoid Facies in the Decapoda 6

1.4 Systematic Position of the Decapod Shrimps Within the Crustacea 9

1.5 Biological Success of Shrimp Groups 11

1.6 Which Group of Shrimps? 13

References 16

2 Structure and Function 19

2.1 From Stem to Stern 19

2.2 Cephalothorax and Carapace 19

2.3 Gills 23

2.4 Eyes 32

2.5 Appendages 36

2.6 Antennules 38

2.7 Second Antennae 41

2.8 Mouthparts 41

2.9 Maxillipeds 45

2.10 Pereopods 50

      2.10.1 Chelae: Shrimp Pincers 51

      2.10.2 Walking Legs 56

2.11 The Abdomen and Its Appendages 57

References 62

3 The Families of Decapod Shrimps 69

3.1 Taxonomic Considerations 69

3.2 Suborder Dendrobranchiata 70

3.3 Superfamily Penaeoidea 73

      3.3.1 Aristeidae 73

      3.3.2 Benthesicymidae 76

      3.3.3 Penaeidae 78

      3.3.4 Sicyoniidae 81

      3.3.5 Solenoceridae 83

3.4 Superfamily Sergestoidea 86

      3.4.1 Luciferidae 86

      3.4.2 Sergestidae 88

3.5 Infraorder Caridea 90

3.6 Superfamily Alpheoidea 91

      3.6.1 Alpheidae 91

      3.6.2 Hippolytidae (s.l.) Families 93

      3.6.3 Barbouriidae 94

      3.6.4 Bythocarididae 95

      3.6.5 Hippolytidae (s.s.) 96

      3.6.6 Lysmatidae 100

      3.6.7 Merguiidae 101

      3.6.8 Thoridae 102

      3.6.9 Ogyrididae 104

3.7 Superfamily Atyoidea: Atyidae 105

3.8 Superfamily Bresilioidea 108

      3.8.1 Family Agostocarididae 108

      3.8.2 Family Alvinocarididae 108

      3.8.3 Family Anchialocarididae 114

      3.8.4 Family Bresiliidae (s.s.) 114

      3.8.5 Disciadidae 115

      3.8.6 Pseudochelidae 116

3.9 Superfamily Campylonotoidea 118

      3.9.1 Bathypalaemonellidae 118

      3.9.2 Family Campylonotidae 119

3.10 Superfamily Crangonoidea 121

      3.10.1 Crangonidae 121

      3.10.2 Glyphocrangonidae 123

3.11 Superfamily Nematocarcinoidea 126

      3.11.1 Eugonatonotidae 126

      3.11.2 Lipkiidae 127

      3.11.3 Nematocarcinidae 129

      3.11.4 Rhynchocinetidae 131

      3.11.5 Xiphocarididae 133

3.12 Superfamily Oplophoroidea 136

      3.12.1 Oplophoridae and Acanthephyridae 138

3.13 Superfamily Palaemonoidea 139

      3.13.1 Desmocarididae 139

      3.13.2 Euryrhynchidae 143

      3.13.3 Palaemonidae (s.l.) 144

      3.13.4 Typhlocarididae 150

3.14 Superfamily Pandaloidea 151

      3.14.1 Pandalidae 151

      3.14.2 Chlorotocellidae 154

3.15 Superfamily Pasiphaeoidea: Pasiphaeidae 154

3.16 Superfamily Physetocaridoidea: Physetocarididae 156

3.17 Superfamily Processoidea: Processidae 157

3.18 Superfamily Psalidopodoidea: Psalidopodidae 160

3.19 Superfamily Stylodactyloidea: Stylodactylidae 162

3.20 Amphionides reynaudii: Taxon Inquirendum or Pandalidae? 163

3.21 Infraorder Procarididea: Procarididae 165

3.22 Infraorder Stenopodidea 168

      3.22.1 Stenopodidae 169

      3.22.2 Spongicolidae 171

      3.22.3 Macromaxillocarididae 172

3.23 Last Word 174

References 174

4 Shrimp Hygiene: Antifouling Adaptations 195

4.1 Fouling and Antifouling Countermeasures 195

4.2 Grooming of Chemosensory Appendages 198

      4.2.1 Antennular Grooming 198

      4.2.2 Adaptive Value of Antennular Grooming 203

      4.2.3 Cleaning of the Antennal Flagellum: The Pereopod 1 Carpal-Propodal Brush 207

4.3 Gill-Cleaning Mechanisms 210

      4.3.1 Reversal and Filtering of the Respiratory Current 211

      4.3.2 Setiferous Epipods 211

      4.3.3 Epipod-Setobranch Complexes 218

      4.3.4 Scaphognathite Gill Cleaning 220

      4.3.5 Gill Cleaning by Chelipeds 222

      4.3.6 Passive Versus Active Gill Cleaning 226

4.4 General Body Grooming 227

      4.4.1 Morphology and Behavior 227

      4.4.2 Adaptive Value of General Body Grooming 232

4.5 Embryo Care 234

4.6 Behavioral Aspects of Grooming 236

4.7 Other Antifouling Adaptations 238

References 239

5 Coloration, Color Change, and Camouflage 247

5.1 Adaptive Value of Coloration 247

5.2 Types and Sources of Color 248

5.3 Color and Camouflage 252

5.4 Mechanisms of Color Change 254

5.5 Rapid Color Change 255

5.6 Day-Night Color Change 257

5.7 Morphological Color Change 260

5.8 Applied Value of Color Change Studies 261

5.9 Color Polymorphism and Camouflage 264

5.10 Warning Coloration 273

5.11 Color and Depth 274

5.12 Bioluminescence and Photophores 275

5.13 Defensive Bioluminescence 281

References 282

6 Reproduction 287

6.1 Complexity of Shrimp Reproductive Morphology 287

      6.1.1 Female Gonads: Ovaries 287

      6.1.2 Male Gonads and Spermatophores 290

      6.1.3 Male Gonadal System: Caridea, Stenopodidea, and Procarididea 290

      6.1.4 Male Gonadal System: Penaeoidea and Sergestoidea 293

6.2 Secondary Sexual Characters: Genitalia 300

      6.2.1 Genitalia of the Procarididea, Stenopodidea, and Caridea 301

      6.2.2 Genitalia of Dendrobranchiata: Thelyca, Petasmata, and Appendices Masculinae 304

6.3 Mechanics of Copulation and Insemination 308

6.4 Eggs, Sperm, and Fertilization 317

      6.4.1 Eggs 317

      6.4.2 Sperm 319

      6.4.3 Sperm Meets Egg: Fertilization 321

6.5 Temporal Cycles of Molting, Mating, Spawning, and Incubation 323

6.6 Spawning, Parental Care, and Breeding Dress 328

References 333

7 Sexual Systems 339

7.1 Sexual Systems and Hermaphroditism 339

7.2 The Direction of Sex Change and the Size Advantage Model 340

7.3 Types and Variations in Sexual Systems 345

      7.3.1 Pure Protandry 346

      7.3.2 Partial Protandry with Primary and/or Early-Maturing Females 346

      7.3.3 Partial Protandry with Primary Males, Primary Females 348

      7.3.4 Partial Protandry with Primary Males 348

      7.3.5 Protandric Simultaneous Hermaphroditism 348

      7.3.6 Pure Simultaneous Hermaphroditism? 349

7.4 Identification of Protandrous Sexual Systems 349

      7.4.1 Size-Frequency Distributions 349

      7.4.2 Transitional Characters 350

      7.4.3 Direct Observation of Sex Change 354

7.5 Taxonomic Distribution of Hermaphroditism in Shrimps 356

      7.5.1 Dendrobranchiata (Sergestoidea and Penaeoidea) 356

      7.5.2 Alpheidae 358

      7.5.3 Atyidae 361

      7.5.4 Campylonotidae 362

      7.5.5 Crangonidae 362

      7.5.6 Hippolytidae 363

      7.5.7 Merguiidae 365

      7.5.8 Pandalidae 366

      7.5.9 Processidae 366

      7.5.10 Rhynchocinetidae 367

      7.5.11 Thoridae 369

      7.5.12 Lysmatidae/Barbouriidae: Protandric Simultaneous Hermaphroditism 372

7.6 Role of the Androgenic Gland in Protandrous Sexual Systems 377

7.7 Environmentally Determined Sex Change (ESD) 378

7.8 Evolution of Protandrous Sexual Systems 383

      7.8.1 Female-First or Male-First Origin of Protandrous Sexual Systems? 383

      7.8.2 The Dendrobranchiate/Palaemonid Paradox 385

      7.8.3 Evolution of Protandric Simultaneous Hermaphroditism (PSH) 386

      7.8.4 Final Thoughts 390

References 390

8 Mating Systems 399

8.1 Sexual Tactics and Strategies 399

8.2 Mate Attraction and Recognition 401

      8.2.1 Visual Cues 401

      8.2.2 Tactile Cues 402

      8.2.3 Sex Pheromones 402

      8.2.4 Olfactory Pheromones 403

      8.2.5 Contact Sex Pheromones 408

      8.2.6 Chemical Composition of Sex Pheromones 408

8.3 Shrimp Mating Systems 410

      8.3.1 Pure Searching 414

      8.3.2 Temporary Mate (Female) Guarding Systems 418

      8.3.3 Search and Attend 418

      8.3.4 Neighborhoods of Dominance (NOD) 420

      8.3.5 Monogamy 426

8.4 Female Choice, Multiple Paternity and Sperm Competition 432

References 435

9 Life Histories 443

9.1 What Is a Life History? 443

9.2 Embryonic Development and Hatching 443

9.3 Larval Development: Costs and Benefits 447

9.4 Extended Larval Development 448

      9.4.1 Dendrobranchiata: Penaeoids and Sergestoids 448

      9.4.2 Carideans 450

      9.4.3 Stenopodideans 453

9.5 Abbreviated and Direct Larval Development 453

9.6 Developmental Strategies of Freshwater Species 456

      9.6.1 Freshwater Abbreviated Larval Development and Direct Development Species 457

      9.6.2 Amphidromous Species 457

      9.6.3 Freshwater Extended Larval Development Species 464

9.7 Abbreviated and Direct Development in Marine Species 465

      9.7.1 High-Latitude Species 466

      9.7.2 Deep-Sea Species 466

      9.7.3 ALD/DD in Shallow-Water Marine Species 467

9.8 Reproductive Investment, Output, and Fecundity 468

9.9 Variation of Fecundity with Female and Egg (Embryo) Size 470

9.10 Fecundity (Brood) Loss 472

9.11 Parasitic Castration by Bopyrid Isopods 474

9.12 Life Cycles, Seasonality, and Latitudinal Trends 476

      9.12.1 Tropical and Warm-Temperate Shallow-Water Species: Dendrobranchiates 477

      9.12.2 Tropical and Warm-Temperate Shallow-Water Species: Pleocyemates 483

      9.12.3 Freshwater Carideans 488

      9.12.4 Cool-Temperate and Boreal/Austral Species 489

      9.12.5 Polar Species 491

9.13 Within Taxon Latitudinal Variation in Life-History Traits in Neritic Marine Species 492

9.14 Deep-Sea Species 495

9.15 The Life-History Continuum of Shrimps 499

9.16 Global Warming and Climatic Effects on Life-History Patterns 502

References 503

10 Symbioses 515

10.1 Symbioses in Shrimps 515

10.2 Taxonomic and Geographic Distribution of Shrimp Symbioses 517

10.3 Common Adaptations of Symbiotic Shrimps 519

10.4 Sponge Dwellers 525

      10.4.1 Caridean Sponge Associates 525

      10.4.2 Spongicolid Sponge Associates 527

      10.4.3 Social Organization in Synalpheus Sponge Inhabitants 529

10.5 Shrimps Associated with Cnidarians 535

      10.5.1 Shrimp Associates of Corals and Related Colonial Anthozoans 536

      10.5.2 Anemone Shrimps 539

      10.5.3 Jellyfish Shrimps 544

10.6 Shrimp Residents of Echinodermata 545

      10.6.1 Echinoidea (Sea Urchin) Associates 546

      10.6.2 Crinoid Shrimps 549

      10.6.3 Other Echinoderm–Shrimp Associations 550

      10.6.4 Host Recognition and Fidelity 551

10.7 Molluscan Hostels 552

10.8 Ascidian Homes 556

10.9 Shrimp Associations with Burrow Dwellers and Other Crustacea 557

10.10 Shrimps as Hosts: Goby Fish–Alpheid Shrimp Symbioses 559

10.11 Shrimps That Clean Fishes: Cleaning Symbiosis 563

10.12 Evolution of Symbioses in Shrimps 571

References 574

11 Fisheries and Aquaculture 583

11.1 A Desirable Food Product 583

11.2 Facts and Figures 583

11.3 Shrimp Fisheries and Aquaculture Production: Economic Value 585

11.4 Composition of the World’s Shrimp Fisheries 586

11.5 Life-History Traits and Shrimp Productivity 588

11.6 Globally Important Fishery Species 596

      11.6.1 Sergestidae 596

      11.6.2 Penaeoidea 596

      11.6.3 Caridea 597

11.7 Top Shrimp Fishing Countries 598

11.8 Fishing Techniques 600

      11.8.1 Catchability and Behavior 600

      11.8.2 Otter and Beam Trawls 600

      11.8.3 Skimmer and Butterfly Nets 602

      11.8.4 Fyke Nets and Traps (Pots) 605

11.9 Management of Shrimp Fisheries 605

      11.9.1 Overfishing 607

      11.9.2 Shrimp Fishery Forecasting and Regulation 608

11.10 Environmental Impacts of the Shrimp Fishing Industry 611

      11.10.1 Bycatch and Discards 611

      11.10.2 Trawling Disturbance of the Sea Bottom and Its Communities 613

      11.10.3 Loss of Coastal Wetlands and Shrimp Fisheries 615

11.11 Scale and Growth of Shrimp Aquaculture 616

11.12 Penaeid Aquaculture 618

      11.12.1 Some History 618

      11.12.2 Principal Species 618

      11.12.3 Principal Countries 623

      11.12.4 Techniques 625

11.13 Freshwater (Macrobrachium) Shrimp Aquaculture 628

      11.13.1 Principal Species 629

      11.13.2 Principal Countries and Some History 630

      11.13.3 Techniques 630

11.14 Shrimp Diseases 633

      11.14.1 Penaeids 633

      11.14.2 Macrobrachium Species 635

11.15 Biosecurity 636

11.16 Monosex Populations in Shrimp Aquaculture 637

11.17 Environmental Impacts of Shrimp Aquaculture 639

11.18 Ornamental Shrimps 641

11.19 Climate Change: Impacts on Fisheries and Aquaculture 643

11.20 Past, Present, and Future State of Shrimp Fisheries and Aquaculture 645

References 647

12 Evolutionary History and Relationships of Shrimps 657

12.1 Shrimp Chronicles 657

12.2 The Fossil Record 657

12.3 Classification and Systematics 660

12.4 Morphological Characters and Their Evolutionary Polarities 663

12.5 Origin of Shrimps from a Decapod Ancestor 665

12.6 Relationships Among Major Shrimp Groups and Other Decapods 667

12.7 Chronological Estimates of Taxa Origins 669

12.8 Relationships Within the Major Shrimp Taxa 670

      12.8.1 Dendrobranchiata 670

      12.8.2 Sergestoidea 671

      12.8.3 Penaeoidea 673

      12.8.4 Caridea and Procarididea 678

      12.8.5 Traditional Views of Caridean Relationships 678

      12.8.6 Christoffersen’s Cladistic Phylogeny 683

      12.8.7 Molecular Tests of Caridean Phylogeny 688

      12.8.8 Stenopodidea 691

References 694

Subject Index 699

Taxonomic Index 711

中国农科院农业信息研究所