Biology of Sharks and Their Relatives (CRC Marine Biology Series)

Winner of Choice Magazine’s Outstanding Academic Title award, January 2005! Sharks and their relatives are the subjects of tremendous interest. The public’s fascination is influenced by their roles in movies and popular literature, while the media races to cover stories of predators endangering helpless humans. The alarming threat to shark populations is also garnering significant publicity and leading to a worldwide increase in conservation initiatives. Finally, technological advances are impacting every area of shark research and revealing incredible secrets about these mysterious animals. These major factors indicate the need for a timely synthesis of the biology of sharks and their relatives. Biology of Sharks and Their Relatives brings together the latest information on the phylogeny, physiology, behavior, and ecology of sharks and their relatives, the skates, rays, and chimaeras. Written by a “Who’s Who” lineup in North American elasmobranch research, this single-source review of elasmobranch fishes presents cohesive and integrated coverage of key topics and discusses technological advances used in modern shark research. The text establishes relationships among the sharks and their relatives that dominate the Chondrichthyes, describes their functions and physiological processes, and examines issues relevant to managing depleted and threatened fisheries. Each of the 19 chapters includes a comprehensive review of the subject with extensive up-to-date citations. This authoritative book provides a synopsis of the current understanding of elasmobranch fishes while identifying gaps in our knowledge to stimulate further study. Its broad coverage and inclusive nature make this an important resource for marine and conservation biologists, fishery scientists, biological oceanographers, zoologists, ecologists, environmental planners, and students. Cover Page......Page 1 Title: BIOLOGY OF SHARKS AND THEIR RELATIVES......Page 4 ISBN 084931514X......Page 5 Dedication......Page 6 Preface......Page 7 The Editors......Page 9 Contributors......Page 10 Contents......Page 13 Color Figures......Page 15 Part I: Phylogeny and Zoogeography......Page 17 1.1 Introduction......Page 19 1.2.1 Tesserate Mineralization......Page 20 1.3 Historic Evidence of Early Chondrichthyans......Page 21 1.3.1.2 Bear Gulch Limestone......Page 23 1.3.1.2.2 The Bear Gulch and Devonian-Permian Chondrichthyan Groups......Page 24 1.3.1.2.3 Community Structure and Population Dynamics......Page 25 1.3.1.2.4 Segregation According to Age, Sex, and Reproductive Stage......Page 27 1.3.1.2.5 Reproductive Strategies......Page 31 1.3.2 Upper Carboniferous and Permian Record......Page 32 1.4.2 On Elasmobranch Origins......Page 33 1.5.1 Trends in the Basal Diversification of the Chondrichthyes (Figure 1.6, Appendix 1.1)......Page 35 1.5.2 Other Concluding Remarks on the Origins of Chondrichthyans, Trends in Chondrichthyan Evolution, and on Characters of the Class......Page 37 Acknowledgments......Page 38 Appendix 1.1: Characters and States for the Cladogram of Figure 1.6......Page 39 References......Page 43 2.1.1 Zoogeographic Patterns......Page 49 2.1.3 Zoogeographic Methods; Brief History......Page 52 2.2.1 Vagility......Page 53 2.3.1.2 Species Diversity......Page 54 2.3.2.1.1 Heterodontiformes......Page 58 2.3.2.1.2 Orectolobiformes......Page 59 2.3.2.1.3 Lamniformes......Page 62 2.3.2.1.4 Carcharhiniformes......Page 65 2.3.2.2.2 Echinorhiniformes......Page 71 2.3.2.2.5 Squaliformes......Page 72 2.4 Summary......Page 76 Acknowledgments......Page 78 Appendix 2.1......Page 79 References......Page 91 3.1 Introduction......Page 95 3.2 Analyses......Page 96 3.3.1 Claspers......Page 97 3.3.2 Phylogenetic Analyses......Page 99 3.4.1 Phylogenetic Implications......Page 102 3.4.2 Evolutionary Implications......Page 108 3.4.4 Classification of Batoids......Page 109 3.4.5 Biogeography of Batoids......Page 112 Acknowledgments......Page 116 Appendix 3.1: Specimens Examined......Page 117 Appendix 3.2: Character Matrix......Page 118 Tooth Root Vascularization and Structure......Page 119 Skeletal Structures......Page 120 Cephalic and Branchial Musculature......Page 124 References......Page 125 4.1 Overview of Living Holocephali......Page 131 4.2 General Ecology and Behavior......Page 133 4.3.1.1 Lateral Line Canals......Page 134 4.3.1.2 Fins and Fin Spines......Page 135 4.3.2 Skeleton......Page 136 4.3.3 Secondary Sexual Characteristics......Page 137 4.5.1 Callorhinchidae (Figure 4.1A)......Page 139 4.5.2 Rhinochimaeridae (Figure 4.1B)......Page 140 Acknowledgments......Page 142 Appendix 4.1: Provisional Key to Species......Page 144 References......Page 147 Part II: Form, Function, and Physiological Processes......Page 153 5.1.1 Approaches to Studying Locomotion in Chondrichthyans......Page 155 5.1.3 Body Form and Fin Shapes......Page 157 5.2.1 Function of the Body during Steady Locomotion and Maneuvering......Page 158 5.2.2 Function of the Caudal Fin during Steady Locomotion and Maneuvering......Page 160 5.2.3.1 Anatomy of the Pectoral Fins......Page 164 5.2.3.2 Role of the Pectoral Fins during Steady Swimming......Page 165 5.2.3.3 Role of the Pectoral Fins during Vertical Maneuvering......Page 168 5.2.3.4 Function of the Pectoral Fins during Benthic Station-Holding......Page 170 5.2.3.5 Motor Activity in the Pectoral Fins......Page 171 5.2.4 Synthesis......Page 172 5.3 Locomotion in Skates and Rays......Page 173 5.4 Locomotion in Holocephalans......Page 175 5.5 Future Directions......Page 176 Acknowledgments......Page 177 References......Page 178 6.1 Introduction......Page 181 6.2.1 Predatory Behaviors......Page 182 6.2.2 Feeding Location and Prey Capture......Page 184 6.3.1 Mechanics of Prey Capture......Page 187 6.3.2 Evolution of the Feeding Mechanism......Page 189 6.3.3.1 Sharks......Page 190 6.3.3.2 Batoids......Page 199 6.4.1 Arrangement and Terminology......Page 202 6.4.2 Evolutionary and Functional Patterns......Page 203 6.5 The Enigma of Jaw Protrusion......Page 208 6.6 Future Directions......Page 209 References......Page 210 7.1 Introduction......Page 219 7.2.1.1 Annular/Circular Respirometers......Page 220 7.2.1.2 Swim Tunnel Respirometers......Page 221 7.2.2.1 Muscle Temperature Telemetry......Page 222 7.2.2.2 Heart Rate Telemetry......Page 223 7.2.2.3 Swimming Speed Telemetry......Page 224 7.3.1 Standard Metabolic Rate......Page 225 7.3.2 Maximum Metabolic Rate......Page 226 7.4.1 Swimming Efficiency......Page 227 7.4.3 Cost of Transport......Page 229 7.5.1 Background......Page 230 7.5.2 Indirect Calorimetry: Endotherms vs. Ectotherms......Page 231 7.6.1 Temperature......Page 232 7.6.3 Dissolved Oxygen......Page 233 7.7 Conclusions and Future Directions......Page 234 References......Page 235 8.1 Introduction......Page 241 8.2.1 Quantification of Diet......Page 242 8.2.2 Broad Dietary Groups......Page 243 8.2.3 Diet Shifts......Page 244 8.2.4 Feeding Relationships......Page 245 8.2.5 Feeding Patterns......Page 246 8.3.1 Daily Ration......Page 247 8.3.2 Gastric Evacuation......Page 248 8.4.2 Egestion......Page 251 8.5 Production......Page 252 References......Page 253 9.1 Introduction......Page 263 9.2.1 Rectal Gland......Page 264 9.2.2 Kidney......Page 266 9.2.3 Gill......Page 269 9.3.1 Rectal Gland......Page 271 9.3.2 Kidney......Page 273 9.3.3 Gills......Page 274 9.4.2 Kidney......Page 276 9.4.3 Gill......Page 277 References......Page 279 10.1 Introduction......Page 285 10.2.1 Oviparous Forms......Page 286 10.2.2.3 Oophagy......Page 287 10.2.2.5 Placental Viviparity......Page 288 10.3 Male Reproductive System......Page 289 10.4.1 Anatomy......Page 294 10.4.3 Sperm Storage......Page 295 10.6 Mating and Reproductive Behaviors......Page 296 10.7 Conclusion......Page 299 References......Page 300 CONTENTS......Page 303 11.2.2 Digestive Hormones......Page 304 11.2.4 Interactions between Feeding and Ion Homeostasis......Page 306 11.2.5 Hormones Involved in Energy Metabolism......Page 307 11.4.1 Chromaffin Tissue and Catecholamines......Page 308 11.4.2 Hypothalamo—Pituitary—Interrenal Axis......Page 310 11.5.2 Interrenal Corticosteroids......Page 311 11.5.5 C-Type Natriuretic Peptide......Page 313 11.6.2 Melanocyte-Stimulating Hormone......Page 314 11.7.2 Gonadotropin-Releasing Hormone......Page 316 11.7.4 Gonadal Steroid Hormones in the Female......Page 319 11.7.6 Gonadal Steroids in the Male......Page 322 11.7.8 Other Hormones Involved in Reproduction: Thyroid Hormones......Page 325 11.7.9 Other Hormones Involved in Reproduction: Calcitonin......Page 326 References......Page 327 CONTENTS......Page 341 12.2 Vision......Page 342 12.2.1 Ocular Anatomy and Optics......Page 343 12.2.2 Retina and CNS......Page 345 12.2.3 Visual Performance......Page 347 12.3.1.2 Macula Neglecta......Page 348 12.3.2.1 Audiograms......Page 349 12.4 Mechanosenses......Page 351 12.4.1 Peripheral Organization......Page 352 12.4.2 Adequate Stimulus and Processing......Page 355 12.4.3 Behavior......Page 356 12.5 Electrosenses......Page 357 12.5.1.1 Ampullae of Lorenzini......Page 358 12.5.2.1 Peripheral Physiology......Page 359 12.5.2.2 Central Physiology......Page 360 12.5.3.1 Prey and Predator Detection......Page 361 12.5.3.2 Orientation and Navigation......Page 362 12.5.3.3 Conspecific Detection......Page 364 12.6.1.1 Peripheral Organ and Epithelium......Page 366 12.6.1.2 Olfactory Bulb......Page 367 12.6.1.3 Higher Level Systems......Page 368 12.6.2.1 Studies of Sharks in Large Enclosures and Open Water......Page 369 12.6.2.2 Laboratory Studies......Page 371 12.6.5 Gustation......Page 372 12.7 Conclusions......Page 373 References......Page 374 13.1 Introduction......Page 385 13.2.1 Lymphocytes......Page 386 13.2.2 Granulocytes......Page 387 13.2.3 Monocytes and Macrophages......Page 388 13.3 Lymphomyeloid Tissues......Page 389 13.3.2 Spleen......Page 390 13.3.3 Epigonal and Leydig Organs......Page 391 13.4.1 Nitric Oxide......Page 393 13.4.3 Phagocytosis and Pinocytosis......Page 394 13.4.4 Nonspecific Cytotoxic Cells......Page 395 13.5.2 Cell-Mediated Immunity......Page 396 13.5.3 Immune System Genes......Page 397 13.6 Ontogeny of the Elasmobranch Immune System......Page 398 13.6.1 Organogenesis of Lymphomyeloid Tissues......Page 399 13.6.2 Embryonic Expression of Immune Regulatory Genes......Page 400 13.7.2 Experimental Induction of Apoptosis......Page 401 13.7.4 Transplantation Studies......Page 402 13.7.5 Effects of Ionizing Radiation on Elasmobranch Immune System......Page 403 13.8 Future Directions......Page 404 References......Page 405 Part III: Ecology and Life History......Page 413 14.1 Introduction......Page 415 14.2.1.1 Vertebrae......Page 416 14.2.1.3 Neural Arches......Page 417 14.2.2.1 Taking Samples......Page 418 14.2.2.2 Centrum Cleaning and Sample Preparation......Page 419 14.3 Age Determination......Page 420 14.3.1 Centrum Staining......Page 423 14.3.3 Back-Calculation......Page 424 14.4 Verification and Validation......Page 425 14.4.2 Tag−Recapture......Page 426 14.4.3 Marking, Field Tag−Recapture, and Laboratory Studies......Page 427 14.4.4 Centrum Edge and Relative Marginal Increment Analysis......Page 428 14.4.5 Captive Rearing......Page 429 14.5 Review and Status of Chondrichthyan Age, Growth, Verification, and Validation Studies......Page 430 14.6 Growth Models......Page 440 14.8.1 Age of Maturity Estimates......Page 441 14.9 Implication of Growth, Longevity, and Demography to Fisheries Management......Page 454 References......Page 455 15.1 Introduction......Page 465 15.2.1 Comparative Life History Patterns......Page 466 15.2.2 Life History Relationships......Page 467 15.3.1 Methodological Background......Page 469 15.3.1.2 Population Sampling Design......Page 473 15.3.2 Biomass Dynamic Models......Page 474 15.3.3.1.1 Age-Structured Models......Page 475 15.3.3.1.3 Yield-per-Recruit Models......Page 476 15.3.4 Models Incorporating Uncertainty and Stochasticity......Page 477 References......Page 480 16.1.1 Genome Sizes......Page 487 16.1.3 Sex-Determination......Page 488 16.2.1.1 Isozymes and Allozymes......Page 489 16.2.1.3 Nuclear DNA......Page 490 16.2.2 Measuring Stock Structure with Molecules......Page 492 16.2.3 Forensic Identification and Cryptic Species......Page 494 16.3.1 Philopatry and Sex-Biased Dispersal......Page 495 16.3.2 Parentage and Multiple Paternity......Page 496 References......Page 497 CONTENTS......Page 503 17.2.2 Avoiding Predators......Page 504 17.2.2.1 Habitat Use......Page 505 17.2.2.4 Group Formation......Page 506 17.2.2.6 Deterrence and Defense......Page 507 17.3 Elasmobranchs as Predators......Page 508 17.3.1 Diets of Elasmobranchs and Optimal Diet Theory......Page 509 17.3.2.1 Habitat Use......Page 510 17.3.2.3 Activity Levels and Patterns......Page 513 17.3.3.1 Stalking and Ambush......Page 514 17.3.3.3 Prey Debilitation......Page 515 17.3.3.7 Scavenging......Page 516 17.3.4 Competition......Page 517 17.3.5.1 Foraging—Safety Trade-Offs......Page 518 17.3.5.3 Intraguild Predation......Page 520 17.4 Regulation of Elasmobranch Populations......Page 521 17.5 Role of Elasmobranchs in Marine Ecosystems......Page 522 17.5.1 Density-Mediated Effects......Page 523 17.5.2 Behavior-Mediated Effects......Page 525 17.5.3 Community Consequences of Elasmobranch Predation......Page 526 References......Page 528 CONTENTS......Page 539 18.1 Introduction......Page 540 18.2.1 Skin......Page 545 18.2.2.1 Eyes......Page 550 18.2.2.3 Acousticolateralis System......Page 551 18.2.3.2 Gills and Branchial Chamber......Page 552 18.2.4.1 Buccal Cavity and Esophagus......Page 553 18.2.4.2 Stomach......Page 554 18.2.4.3 Spiral Intestine......Page 555 18.2.4.4 Rectum......Page 556 18.2.5.1 Heart and Vasculature......Page 557 18.2.6.3 Uterus......Page 558 18.2.8.2 Peritoneal Cavity......Page 559 18.3 General Observations......Page 560 References......Page 561 19.1 Introduction......Page 569 19.2.2 Habitat Preference......Page 571 19.3.3 Acoustic Tracking......Page 572 19.3.4 Acoustic Monitoring......Page 575 19.3.5 Satellite Telemetry......Page 577 19.3.6 Archival Tags......Page 578 19.4 The Importance of Scale in Habitat Use Studies......Page 581 19.4.1.1 Diel Effects......Page 582 19.4.1.2 Longer-Term Effects......Page 583 19.4.2 Spatial Factors......Page 584 19.5.2 Biotic Factors......Page 585 References......Page 586 B......Page 589 D......Page 590 F......Page 591 G......Page 592 H......Page 593 L......Page 594 O......Page 595 P......Page 596 R......Page 597 S......Page 598 W......Page 599 Z......Page 600 A......Page 601 C......Page 602 D......Page 603 G......Page 604 H......Page 605 L......Page 606 O......Page 607 P......Page 608 S......Page 609 X......Page 611 Z......Page 612 Back Page......Page 613 Synopsis: Sharks And Their Relatives Are The Subjects Of Tremendous Interest. The Public's Fascination Is Influenced By Their Roles In Movies And Popular Literature, While The Media Races To Cover Stories Of Predators Endangering Helpless Humans. The Alarming Threat To Shark Populations Is Also Garnering Significant Publicity And Leading To A Worldwide Increase In Conservation Initiatives. Finally, Technological Advances Are Impacting Every Area Of Shark Research And Revealing Incredible Secrets About These Mysterious Animals. These Major Factors Indicate The Need For A Timely Synthesis Of The Biology Of Sharks And Their Relatives. Biology Of Sharks And Their Relatives Brings Together The Latest Information On The Phylogeny, Physiology, Behavior, And Ecology Of Sharks And Their Relatives, The Skates, Rays, And Chimaeras. Written By A Who's Who Lineup In North American Elasmobranch Research, This Single-source Review Of Elasmobranch Fishes Presents Cohesive And Integrated Coverage Of Key Topics And Discusses Technological Advances Used In Modern Shark Research. The Text Establishes Relationships Among The Sharks And Their Relatives That Dominate The Chondrichthyes, Describes Their Functions And Physiological Processes, And Examines Issues Relevant To Managing Depleted And Threatened Fisheries. Each Of The 19 Chapters Includes A Comprehensive Review Of The Subject With Extensive Up-to-date Citations. This Authoritative Book Provides A Synopsis Of The Current Understanding Of Elasmobranch Fishes While Identifying Gaps In Our Knowledge To Stimulate Further Study. Its Broad Coverage And Inclusive Nature Make This An Important Resource For Marine And Conservation Biologists, Fishery Scientists, Biological Oceanographers, Zoologists, Ecologists, Environmental Planners, And Students. The Origin And Relationships Of Early Chondrichthyes / Eileen D. Grogan, Richard Lund -- Historical Zoogeography Of The Selachii / John A. Musick, Melanie M. Harbin, Leonard J.v. Compagno -- Phylogeny Of Batoidea / John D. Mceachran, Neil Aschliman -- Phylogeny And Classification Of Extant Holocephali / Dominique A. Didier -- Biomechanics Of Locomotion In Sharks, Rays, And Chimeras / Cheryl A.d. Wilga, George V. Lauder -- Prey Capture Behavior And Feeding Mechanics Of Elasmobranchs / Philip J. Motta -- Metabolism, Energetic Demand, And Endothermy / John K. Carlson, Kenneth J. Goldman, Christopher G. Lowe -- Food Consumption And Feeding Habits / Bradley M. Wetherbee, Enric Cortés -- Homeostasis : Osmoregulation, Ph Regulation, And Nitrogen Excretion / David H. Evans, Peter M. Piermarini, Keith P. Choe -- Reproductive Biology Of Elasmobranchs / Jeffrey C. Carrier, Harold L. Pratt, Jr., José I. Castro -- Hormonal Regulation Of Elasmobranch Physiology / James Gelsleichter -- Sensory Biology Of Elasmobranchs / Robert E. Hueter ... [and Others] -- The Immune System Of Sharks, Skates, And Rays / Carl A. Luer, Catherine J. Walsh, Ashby B. Bodine -- Age Determination And Validation In Chondrichthyan Fishes / Gregor M. Cailliet, Kenneth J. Goldman -- Life History Patterns, Demography, And Population Dynamics / Enric Cortés -- Genetics Of Sharks, Skates, And Rays / Edward J. Heist -- Predator-prey Interactions / Michael R. Heithaus -- Elasmobranchs As Hosts Of Metazoan Parasites / Janine N. Caira, Claire J. Healy -- Assessing Habitat Use And Movement / Colin A. Simpfendorfer, Michelle R. Heupel. [edited By] Jeffrey C. Carrier, John A. Musick, And Michael R. Heithaus. Includes Bibliographical References And Index. To address subject areas and subdisciplines where coverage was absent or superficial in volume one, Jeffrey Carrier and associates have assembled in the current volume a collection of works that reveal patterns of biodiversity, the physiological attributes that contribute to elasmobranchs' successful exploitation of oceanic and freshwater realms, and the unique issues associated with the interaction between elasmobranchs and humans, all of this with overarching attention to issues of conservation. "We begin with chapters examining biodiversity. We have chosen to approach this discussion by presenting elasmobranchs as inhabitants of the range of zoogeographic provinces, realizing that significant overlap may occur for more pelagic species. This realization was reflected in the dialogue that occurred during preparation of the book between our chapter authors, and the recognition that many species simply cannot be confined to a specific habitat or range of habitats. We then continue by examining some of the unique physiological adaptations that allow these animals to exploit the range of habitats where they are found, from unique sensory modalities to compensatory mechanisms for physiological and environmental stress. "Our concluding section presents some of the challenges faced by members of these groups. We have asked our authors to consider human interactions and anthropogenic effects on worldwide populations and the potential extinction risks posed from survival under increasing threats from changes in habitat, changes in water chemistry, and increasing commercial exploitation. Conservation of species under threat remains a theme throughout the book Since the award-winning first volume, The Biology of Sharks and Their Relatives, published in 2004, the field has witnessed tremendous developments in research, rapid advances in technology, and the emergence of new investigators beginning to explore issues of biodiversity, distribution, physiology, and ecology in ways that eluded more traditional studies. As an entirely new companion volume, Sharks and Their Relatives II: Biodiversity, Adaptive Physiology, and Conservation brings you up to speed on these significant changes, specifically examining how elasmobranch fishes - the sharks, skates, rays, and chimaeras - successfully survive in a wide range of habitats. Emphasizes Conservation of Threatened Species This multidisciplinary volume begins by examining elasmobranch biodiversity patterns and their integrated sensory systems. It then explores the physiological adaptations - from unique sensory modalities to compensatory mechanisms for physiological and environmental stress - that make these animals particularly well-suited for the range of habitats where they are found, in both oceanic and freshwater realms. Features Established Researchers and Introduces New Pioneers in the Field The book then considers the human interactions and anthropogenic effects on worldwide elasmobranch populations and the potential extinction risks posed by increasing threats from changes in habitat, changes in water chemistry, and growing commercial exploitation. This text truly is unrivaled in terms of coverage and readability, and it is a must-have reference for marine biologists, fishery scientists, oceanographers, and also marine, zoo, and aquarium veterinarians