The Biology of Sharks and Rays
by A. Peter Klimley, illustrated by Steven Oerding
University of Chicago Press, 2013
Cloth: 978-0-226-44249-5 | Electronic: 978-0-226-92308-6
DOI: 10.7208/chicago/9780226923086.001.0001
ABOUT THIS BOOKAUTHOR BIOGRAPHYREVIEWSTABLE OF CONTENTS

ABOUT THIS BOOK

The Biology of Sharks and Rays is a comprehensive resource on the biological and physiological characteristics of the cartilaginous fishes: sharks, rays, and chimaeras. In sixteen chapters, organized by theme, A. Peter Klimley covers a broad spectrum of topics, including taxonomy, morphology, ecology, and physiology. For example, he explains the body design of sharks and why the ridged, toothlike denticles that cover their entire bodies are present on only part of the rays’ bodies and are absent from those of chimaeras. Another chapter explores the anatomy of the jaws and the role of the muscles and teeth in jaw extension, seizure, and handling of prey. The chapters are richly illustrated with pictures of sharks, diagrams of sensory organs, drawings of the body postures of sharks during threat and reproductive displays, and maps showing the extent of the species’ foraging range and long-distance migrations. Each chapter commences with an anecdote from the author about his own personal experience with the topic, followed by thought-provoking questions and a list of recommended readings in the scientific literature. 

The book will be a useful textbook for advanced ichthyology students as well as an encyclopedic source for those seeking a greater understanding of these fascinating creatures.

AUTHOR BIOGRAPHY

A. Peter Klimley is adjunct professor in the Department of Wildlife, Fish, and Conservation and director of the Biotelemetry Laboratory at the University of California, Davis. He is the author of The Secret Life of Sharks.

REVIEWS

“A. Peter Klimley has been studying the biology of sharks for decades. In this book, he combines this extensive research experience with that of others to produce a solid reference for every shark enthusiast.”

— Gregory Skomal, The Shark Handbook: The Essential Guide for Understanding the Sharks of the World

“This book is a novel publication in its field—such a cohesive compilation of all aspects of Chondrichthyan fish life was surprisingly nonexistent prior to this text. . . . We believe Peter Klimley’s attempt would serve admirably as a textbook.”
— Journal of Fish Biology

“An appealing, comprehensive, and exciting resource on the study of many cartilaginous fishes. . . . What separates this work from other highly useful volumes on elasmobranch fishes is its accessibility and tone. Klimley’s experience as a teacher and mentor really shine in the text; the detailed summaries of the science are nicely paced and easy to read and understand for readers of all backgrounds. Additionally, the book is replete with wonderfully illustrated figures and panels (especially on the anatomy and behavioral ethograms), as well as clear and informative data tables, all of which present actual scientific data from published work. . . . Shark and ray scientists and teachers alike will appreciate the attention to detail and accessibility of this book, and its pages serve as a stepping stone for those interested in deeper investigations into the literature. This resource is highly recommended and should be used by a wide-ranging audience, from high school and college students, to teachers, and even experts in the field. Klimley should be very proud of this career-spanning accomplishment.”
— Austin J. Gallagher, University of Miami, Bulletin of Marine Science

“This is really the essential Shark Week companion for nature TV fans, and the chapter on cartilaginous fishes and humans is an especially rigorous antidote to oversensationalization. More than that though, this book is a comprehensive overview of the state of biological knowledge of these fishes. It is logically laid out, with excellent illustrations and abundant, current citations. These features, plus discussion questions for every chapter, make it a very functional textbook, but the spotlight sections and engaging writing should make it appeal to a much broader audience.”
— Conservation Biology

TABLE OF CONTENTS

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0001
[cartilaginous fishes, benchmark studies, sharks, rays, chimaeras, Chondrichthyes]
There are currently 503 species of sharks, 699 species of rays, and 49 species of chimaeras in the class Chondrichthyes. The name Chondrichthyes comes from the Greek prefix khondros meaning “cartilage” and suffix ikhthus for “fish.” These species are referred to as the cartilaginous fishes. This chapter provides an overview of the subsequent chapters, which describe benchmark studies on the cartilaginous fishes. The results are presented in diagrams and graphs to introduce readers to authentic scientific data. “Spotlights” are included in each chapter to focus attention on the sophisticated methods used by pioneer scientists to make important scientific discoveries. (pages 1 - 10)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0002
[evolution, cartilaginous fishes, sharks, chimaeras, Paleozoic era, Mesozoic era]
This chapter discusses the long evolutionary history of the cartilaginous fishes. Topics covered include the sharks and chimaeras of the Paleozoic era (species diversification, Early Paleozoic sharks, Late Paleozoic sharks, Paleozoic chimaeras); the Paleozoic ecosystem (Permian extinctions, sharks and rays of the Mesozoic Era); the demise of the giant-tooth shark; and orders of extant the cartilaginous fishes. (pages 11 - 50)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0003
[swimming styles, sharks, rays, chimaeras, skeleton, muscular anatomy, dermal denticles, tail fins]
This chapter discusses the swimming styles of sharks, rays, and chimaeras. It covers differences in their skeletal and muscular anatomy; their skin and the dermal denticles that cover their bodies; hypotheses in scientific literature explaining how they use their tail fins to propel themselves forward and the empirical evidence supporting them; and commonly used classification of the diverse modes of swimming exhibited by the cartilaginous fishes. (pages 51 - 82)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0004
[sharks, rays, chimaeras, salt composition, osmoregulation, osmoregulatory organs]
This chapter describes the internal hydro-mineral balance of the sharks, rays, and chimaeras, with special attention given to how species regulate the salt composition of their tissues in order to temporarily inhabit estuarine and freshwater environments or reside wholly in the freshwater environment. The discussion covers the process of osmoregulation and the osmoregulatory organs (rectal gland, kidney, gills). (pages 83 - 100)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0005
[sharks, rays, body temperature, cartilaginous fishes, rete mirabile, metabolic heat]
This chapter discusses the physiological processes and anatomical adaptations that enable some sharks and rays to elevate the temperature of key body parts over the ambient temperature of the ocean. The members of two families of sharks, the Lamnidae and Alopiidae, and one family of rays, the Mobulidae, have evolved anatomical and physiological adaptations that enable them to maintain parts of their body warmer than the external environment. These species possess a rete mirabile that provides a barrier against the loss of metabolic heat. This internal network of closely associated arteries and veins is a countercurrent heat exchanger because the metabolic heat diffuses through the thin walls of veins carrying blood to the exterior surface of the fish and again through the thin walls of the arteries to warm the cool blood flowing toward the interior muscles of the fish. (pages 101 - 124)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0006
[sharks, cartilaginous fishes, olfactory receptor, animon acids, amines]
The cartilaginous fishes, in particular the sharks, can detect their prey at great distances using their senses of smell and hearing. This chapter discusses the anatomy of the olfactory receptor, how it differs among different species, the receptor's sensitivity to a myriad of chemicals, and the ability to localize the source of an odor source. Molecules of various chemicals in continuous water flows create a gradient from a high to a low concentration. Cartilaginous fishes detect these molecules, which are dissolved in water, as they pass through their nasal cavities; this process is called chemoreception. The cartilaginous fishes are sensitive to a wide spectrum of chemicals, those most attractive are amino acids and amines. (pages 125 - 152)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0007
[sharks, rays, chimaeras, low-frequency sounds, mechanoreceptors, pressure oscillation, inner ear, neuromasts, Savi]
This chapter discusses how sharks, rays, and chimaeras are able to locate their prey and avoid predators from large distances. This sense is called mechanoreception because mechanical disturbances of the water are detected by two organ systems. The free neuromasts, canal neuromasts, and vessels of Savi on the external body are sensitive to one-way displacements of particles of water. The inner ear is sensitive not only to these displacements but also to two-way displacements of water particles associated with pressure oscillations. The former receptors are active in the near field, where the one-way oscillations exceed the two-way oscillations; the latter receptor is active in both the near and far field, where the two-way oscillations are dominant. The boundary between these fields for a very low frequency of 10 Hz is 150 m while for 100 Hz it is only 15 m. (pages 153 - 184)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0008
[cartilaginous fishes, vision, eye, underwater photic environment]
This chapter discusses the underwater photic environment, the anatomy of the eye, the spectral sensitivities of visual pigments, and the visual capabilities of the different cartilaginous fishes. All of the sharks, rays, and chimaeras have small pores on the underside of their heads connected to gel-filled tubules that lead to the nervous system, called the ampullae of Lorenzini. These sense minute electrical fields, which are produced by fish, clams, and crabs while out of sight buried in the sand. (pages 185 - 212)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0009
[electromagnetic sense, sharks, rays, chimaeras, magnetic field, ampullae of Lorenzini, prey detection, mate detection]
This chapter discusses the electromagnetic sense which enables sharks, rays, and chimaeras to find their way in the apparently featureless ocean by following the subtle patterns of magnetization on the sea floor or the earth's dipolar magnetic field. It covers the anatomy and distribution of ampullae of Lorenzini; prey detection; mate detection; orientation and navigation; and the hierarchy of senses. (pages 213 - 238)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0010
[cartilaginous fishes, sharks, rays, chimaeras, brain development, learning]
This chapter discusses the anatomy of the brains of the cartilaginous fishes and their learning capabilities. Sharks, rays, and chimaeras can learn as quickly as birds and mammals, and have brains of a comparable size and elaboration. (pages 239 - 264)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0011
[sharks, rays, reproductive biology, sexual maturation, spermatogenesis, embryonic development, copulation]
This chapter discusses the courtship behavior and reproductive biology of sharks and rays. It covers the spermatogenesis and sexual maturation of male sharks and rays; sexual maturation and embryonic development of female sharks and rays; the reproductive cycle; courtship and copulation; and promiscuity and multiple paternity. (pages 265 - 310)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0012
[sharks, rays, chimaeras, cartilaginous fishes, teeth, jaws, predation]
This chapter discusses the anatomy of the jaws and teeth of cartilaginous fishes as well as the role of the muscles and teeth in jaw extension, seizure, and handling of prey. Most sharks, rays, and chimaeras are tertiary consumers, occupying the same trophic level as the prey of the white shark. These species feed upon small fishes, cephalopods, and mollusks that in turn feed on zooplankton. There are three steps in active predation: approach, seizure, and handling of the prey. The extent of attachment of the jaws to the cranium has lessened over evolutionary time in the cartilaginous fishes. The ancient sharks needed to swallow their prey whole due to the rigid attachment of the jaws to the cranium. The jaws of the modern sharks trend toward a less rigid attachment to the cranium. (pages 311 - 338)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0013
[cartilaginous fishes, feeding, digestion, intestines]
This chapter discusses what cartilaginous fishes eat, their frequency of feeding, their rates of digestion and growth, and to what age they live. The composition of the diet of the cartilaginous fishes often changes as they grow larger and migrate from one geographical region to another. The diet can also differ among males and females because in many cartilaginous fishes, the sexes segregate from each other and occupy separate feeding grounds. The cartilaginous fishes are unique in their possession of spiral and scrolled intestines, which confers both advantages and disadvantages. Their compactness affords more space in the peritoneum to accommodate a larger liver and uterus. Yet the economy in mucosal surface results in a slower digestion rate and prolongs the interval between foraging bouts. (pages 339 - 372)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0014
[sharks, rays, ambushers, cartilaginous, migration, oscillatory diving, surface swimming]
This chapter discusses the diverse types of movement patterns exhibited by sharks and rays. Some elasmobranchs are ambushers, and their movements are restricted to a confined area. Many skates, some stingrays, and angelsharks remain in wait on the bottom for long periods of time to ambush crabs and shrimps or fishes that walk or swim close to the bottom. Many cartilaginous fishes exhibit two types of travel modes, performing highly directional and rapid movements as they migrate between resting or foraging locations, and then exhibiting randomly directed slow movements when interacting socially or foraging at their feeding grounds. During their migrations, these species display two types of swimming behaviors, oscillatory diving and surface swimming. (pages 373 - 404)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0015
[shark attacks, stingrays, ecotourism, tourist attractions]
This chapter discusses the interactions between the cartilaginous fishes and humans. It describes the risk of being bitten by a shark or being impaled by the barb of a stingray and the growing interest of ecologically aware tourists in viewing sharks underwater. It recounts a few infamous shark attacks to illustrate the severe economic repercussions of such a highly publicized mishap; explains the motivations behind this behavior; it also places the modest risk of shark attack in the context of many other greater risks in daily life. The chapter also considers the economic value of shark and ray ecotourism, and provides information on what species can now be viewed underwater and where to go to see them. (pages 405 - 436)
This chapter is available at:
    https://academic.oup.com/chica...

- A. Peter Klimley
DOI: 10.7208/chicago/9780226923086.003.0016
[sharks, rays, elasmobranch fisheries, fisheries management, marine reserves]
This chapter focuses on the recent expansion of fisheries for sharks and rays, and the decline of the populations of sharks and rays in the world's oceans. It describes historical examples of the collapse of shark fisheries; the methods of managing elasmobranch fisheries including recent regulations; and the recent establishment of reserves to protect sharks and rays. It suggests that the factors which made the sharks, rays, and chimaeras so successful over time also have made them very vulnerable today in a world dominated by humans. (pages 437 - 465)
This chapter is available at:
    https://academic.oup.com/chica...