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Douglas E. Facey

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THE DIVERSITY OF FISHES

The third edition of The Diversity of Fishes is a major revision of the widely adopted ichthyology textbook, incorporating the latest advances in the biology of fishes and covering taxonomy, anatomy, physiology, biogeography, ecology, and behavior. Key information on the evolution of various fishes is also presented, providing expansive and conclusive coverage on all key topics pertaining to the field.

To aid in reader comprehension, each chapter begins with a summary that provides a broad overview of the content of that chapter, which may be particularly useful for those using the text for a course who don’t intend to address every chapter in detail. Detailed color photographs throughout the book demonstrate just some of the diversity and beauty of fishes that attract many to the field. A companion website provides related videos selected by the authors, instructor resources, and additional references and websites for further reading.

Sample topics covered and learning resources included in The Diversity of Fishes are as follows:

  • How molecular genetics has transformed many aspects of ichthyology
  • The close relationship between structure and function, including adaptations to special environments
  • Many physical and behavioral adaptations reflecting the fact that many fishes are both predators and prey
  • Fish interactions with other species within fish assemblages and broader communities, plus their impacts on ecosystems
  • Global maps that more accurately represent the comparative sizes of oceans and land masses than maps used in prior editions

For students, instructors, and individuals with an interest in ichthyology, The Diversity of Fishes is an all-in-one introductory resource to the field, presenting vast opportunities for learning, many additional resources to aid in information retention, and helpful recommendations on where to go to explore specific topics further.

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Veröffentlichungsjahr: 2022

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Table of Contents

Cover

Title Page

Copyright Page

Dedication Page

Preface to the Third Edition

Preface to the Second Edition

Acknowledgments

Preface to the First Edition

Adaptations versus adaptationists

Acknowledgments

Acknowledgments

About the Companion Website

PART I: Introduction

CHAPTER 1: The Science of Ichthyology

Summary

Introduction

What Is a Fish?

Vertebrate Classes

The Diversity of Fishes

Superlative Fishes

A Brief History of Ichthyology

Introduction to Fish Genetics

Additional Sources of Information

Supplementary Reading

CHAPTER 2: Phylogenetic Procedures

Summary

Importance of Classification

Species Concepts

Taxonomy Versus Systematics

Approaches to Classification

Morphological Characters

Molecular Characters

Units of Classification

International Code of Zoological Nomenclature

PhyloCode

Name Changes

Collections

Supplementary Reading

PART II: Form, Function, Ontogeny

CHAPTER 3: Structure and Function of the Head

Summary

Osteology

Cranial Skeleton

Muscles of the Head

Breathing

Functional Morphology of Feeding

Supplementary Reading

CHAPTER 4: Structure and Function of the Trunk and Fins

Summary

Postcranial Skeleton

Appendicular Skeleton

Muscles

Integument

Median Fins

Locomotion: Movement and Shape

Supplementary Reading

CHAPTER 5: Circulation, Gas Transport, Metabolism, Digestion, and Energetics

Summary

Cardiovascular System

Gas Exchange and Transport

Metabolic Rate

Energetics and Buoyancy

Energy Intake and Digestion

Bioenergetics Models

Supplementary Reading

CHAPTER 6: Nervous System and Sensory Organs

Summary

Nervous System

Sensory Organs

Supplementary Reading

CHAPTER 7: Homeostasis

Summary

Coordination and Control of Regulation

Temperature Relationships

Osmoregulation, Excretion, Ion and pH Balance

The Immune System

Stress

Supplementary Reading

CHAPTER 8: Reproduction

Summary

Determination, Differentiation, and Maturation

Gonads and Gametogenesis

Energetic Investment in Reproduction

Sex and Gender Roles in Fishes

The Evolution of Sex Change in Fishes

Sexual Dimorphism

Sexual Selection

Fertilization

Embryo Development

Supplementary Reading

CHAPTER 9: Larvae, Juveniles, Adults, Age, and Growth

Summary

Early Life History

Juveniles

Adults

Age and Growth

The Ontogeny and Evolution of Growth

Supplementary Reading

CHAPTER 10: Special Habitats and Special Adaptations

Summary

The Deep Sea

The Open Sea

Intertidal Fishes

Strong Currents and Turbulent Water

Extreme pH and Salinity

Polar Regions

Deserts and Other Seasonally Arid Habitats

Caves

Supplementary Reading

PART III: Taxonomy, Phylogeny, and Evolution

CHAPTER 11: A History of Fishes

Summary

The First Vertebrates

Extinct Jawless Fishes

Later Evolution of Primitive Agnathous Fishes

Early Gnathostomes: the First Jawed Fishes

Advanced Jawed Fishes I: Teleostomes

Advanced Jawed Fishes II: Chondrichthyes

A History of Fishes: From Bones to Genomes

Supplementary Reading

CHAPTER 12: Chondrichthyes: Sharks, Skates, Rays, and Chimaeras

Summary

Class Chondrichthyes, Subclass Euselachii

Class Chondrichthyes, Subclass Holocephali

Supplementary Reading

CHAPTER 13: Living Representatives of Primitive Fishes

Summary

Lancelets and Jawless Fishes (Hagfishes and Lampreys) – Living Groups

Hagfishes and Lampreys: Evolutionary Relationships

Primitive Bony Fishes – Living Groups

Conclusions

Supplementary Reading

CHAPTER 14: Teleosts I: Elopomorpha Through Paracanthopterygii

Summary

Teleostean Phylogeny

A Survey of Living Teleostean Fishes

Supplementary Reading

CHAPTER 15: Teleosts II: Spiny‐Rayed Fishes

Summary

Superorder Acanthopterygii: Introduction

Series Berycimorphaceae (Berycida)

Series Holocentrimorphaceae

Series Percomorphaceae

Supplementary Reading

PART IV: Fish Behavior

CHAPTER 16: Fishes as Predators andPrey

Summary

Introduction

Search

Pursuit

Attack and Capture

Handling

Scavengers, Detritivores, and Herbivores

Balancing Costs and Benefits

Supplementary Reading

CHAPTER 17: Fishes as SocialAnimals

Summary

Reproduction

Reproductive Patterns Among Fishes

Courtship and Spawning

Parental Care

Alternative Mating Systems and Tactics

Genetic Resolution of Mating Systems

Fish Aggregation, Aggression, and Cooperation

Communication

Agonistic Interactions

Aggregations

Interspecific Relations: Symbioses

Supplementary Reading

CHAPTER 18: Cycles of Activity and Behavior

Summary

Introduction to Cycles of Behavior

Diel Patterns

Semilunar and Lunar Patterns

Seasonal Patterns

Annual and Supra‐Annual Patterns: Migrations

Supplementary Reading

PART V: Fish Distribution, Ecology, and Conservation

CHAPTER 19: Zoogeography and Phylogeography

Summary

Continental Drift, Tectonic Plates, and Fish Distributions

Vicariance and Dispersal

Fish Distributions in Major Zoogeographic Regions

Marine Fishes

Marine Zoogeographic Regions

Marine Fish Phylogeography

Freshwater Fishes

Freshwater Zoogeographic Regions

Archaic Freshwater Fish Distributions

More Recent Distributions

Phylogeography of Freshwater Fishes

Anadromous Fishes

Supplementary Reading

CHAPTER 20: Fish Populations

Summary

Population Ecology

Population Dynamics and Regulation

Population Genetics

Hybridization

Supplementary Reading

CHAPTER 21: The Functional Role of Fishes in Communities and Ecosystems

Summary

Introduction to Assemblages, Communities, and Ecosystems

Assemblages

Interactions within Assemblages and the Broader Ecological Community

The Effects of Fishes on Vegetation

The Effects of Fishes on Invertebrate Activity, Distribution, and Abundance

Fishes and Trophic Structure of Communities

Fishes and Ecosystem Ecology

Influence of Physical Factors and Disturbance

Ecosystem Services Provided by Fishes

Supplementary Reading

CHAPTER 22: Conservation

Summary

Introduction – A Brief History of Conservation Biology

Extinction and Biodiversity Loss

General Causes of Biodiversity Decline

Protecting Fish Biodiversity – What Can Be Done?

Supplementary Reading

Appendix 1 ‐ Glossary of Genetic/Molecular Terminology

References

Index

End User License Agreement

List of Tables

Chapter 1

TABLE 1.1 The diversity of living fishes. Below is a brief listing of highe...

TABLE 1.2 Chromosome number, nuclear genome size, and mitochondrial genome ...

Chapter 2

TABLE 2.1 Classification of Atlantic Herring, Brown Trout, and Chub Mackere...

Chapter 4

TABLE 4.1 Form, function, and locomotion in fishes. About 12 generalized ty...

Chapter 5

TABLE 5.1

Diversity of fishes with air‐breathing capabilities.

Chapter 10

TABLE 10.1 Representative teleostean taxa from the three major deep‐sea hab...

TABLE 10.2 A sampling of freshwater fishes that inhabit torrent and rapid z...

Chapter 11

TABLE 11.1 Repeated trends in fish evolution. Although fishes represent div...

Chapter 12

TABLE 12.1 A summary of embryonic development and nutrition in chondrichthy...

Chapter 13

TABLE 13.1 Characteristics of extant relict fishes. Presence (+) or absence...

Chapter 17

TABLE 17.1 A summary of components of breeding systems in fishes, with repr...

TABLE 17.2 A classification of reproductive guilds in teleost fishes, based...

Chapter 18

TABLE 18.1 Diel activity patterns – defined as when fishes feed – of better...

TABLE 18.2 Families of known diadromous fishes. Modified from McDowall (198...

Chapter 19

TABLE 19.1 Numbers of species in selected genera of inshore fishes from the...

TABLE 19.2 The number of management units and evolutionary significant unit...

Chapter 20

TABLE 20.1 A life table for a cohort of Brook Trout, Salvelinus fontinalis ...

TABLE 20.2 Population genetic diversity averaged across three types of fish...

TABLE 20.3 Comparison of pelagic larval duration and population structure i...

Chapter 21

TABLE 21.1

Top‐down effects of fishes in temperate lakes and streams.

Chapter 22

TABLE 22.1 Ten commonly introduced but controversial fish species. Species ...

TABLE 22.2 Examples from field studies demonstrating reproductive and devel...

List of Illustrations

Chapter 1

FIGURE 1.1 A cladogram showing the relationships of the major groups of chor...

FIGURE 1.2 Fish versus fishes. By convention, “fish” refers to one or more i...

FIGURE 1.3 Some very large fishes. (A) The Ocean Sunfish (

Mola mola

) is esti...

Chapter 2

FIGURE 2.1 Cladogram of hypothesized relationships of the Louvar (

Luvarus

, L...

FIGURE 2.2 A hypothetical phylogeny is showing groups that are monophyletic,...

FIGURE 2.3 Some meristic and morphometric characters are shown on a hypothet...

FIGURE 2.4 The shaded area represents a phylogenetic tree showing the evolut...

FIGURE 2.5 Phylogeny of the most ancient lineages of extant fishes. Previous...

FIGURE 2.6 Phylogenetic relationships among actinopterygian fishes based on ...

FIGURE 2.7 A maximum likelihood phylogeny for the ray‐finned fishes (Actinop...

FIGURE 2.8 Phylogenetic relationships of bonefish species based on maximum l...

Chapter 3

FIGURE 3.1 A cleared and stained specimen of Pacific Cod (

Gadus macrocephalu

...

FIGURE 3.2 Lateral view of the skull of the Dogtooth Tuna (

Gymnosarda unicol

...

FIGURE 3.3 Lateral view of the neurocranium of the Dogtooth Tuna (

Gymnosarda

...

FIGURE 3.4 Dorsal view of the neurocranium of the Dogtooth Tuna (

Gymnosarda

...

FIGURE 3.5 Ventral view of the neurocranium of the Dogtooth Tuna (

Gymnosarda

...

FIGURE 3.6 Left infraorbital bones in lateral view of the Spanish Mackerel (

FIGURE 3.7 Rear view of the skull of a bonito (

Sarda chiliensis

). The crosse...

FIGURE 3.8 Lateral bones of the face and lower jaw suspension of generalized...

FIGURE 3.9 Left hyoid complex in lateral view of a Spanish mackerel (

Scomber

...

FIGURE 3.10 Major types of jaw suspension in fishes.

FIGURE 3.11 Branchial arch of a Spanish mackerel (

Scomberomorus semifasciatu

...

FIGURE 3.12 Cheek muscles of a sculpin,

Jordania zonope

. (A) Superficial mus...

FIGURE 3.13 (A, B) The gill arches of a fish support the gill filaments (als...

FIGURE 3.14 The timing of the expansion and contraction of the buccal (oral)...

FIGURE 3.15 (A) Hagfishes have one or more external gill openings on each si...

FIGURE 3.16 (A) Opening, protrusion, and closing of the jaw in most percoids...

FIGURE 3.17 Extreme jaw protrusion in the Sling‐jaw Wrasse,

Epibulus insidia

...

FIGURE 3.18 Crushing ability of the pharyngeal jaws in three related wrasses...

FIGURE 3.19 The protrusible pharyngeal jaws of moray eels (artwork by Zina D...

FIGURE 3.20 Convergence in dentition among predatory fishes. The triangular,...

FIGURE 3.21 African Tigerfish,

Hydrocynus vittatus

., are predatory alestids....

FIGURE 3.22 Fishes that feed on hard‐bodied prey crush their prey with molar...

FIGURE 3.23 Correspondence among mouth position, feeding habits, and water c...

FIGURE 3.24 The supraterminal mouth of an Atlantic Tarpon (

Megalops atlantic

...

Chapter 4

FIGURE 4.1 Junction of precaudal and caudal vertebrae in a left lateral view...

FIGURE 4.2 Representative precaudal and caudal vertebrae of generalized char...

FIGURE 4.3 Posterior vertebrae and caudal complex of generalized characin (

B

...

FIGURE 4.4  Caudal complex in left lateral view of a Spanish Mackerel (

Scomb

...

FIGURE 4.5 Left pectoral girdle of generalized characin (

Brycon meeki

).

FIGURE 4.6 The anatomy of swimming in teleosts. (A) Lateral view of a Spotte...

FIGURE 4.7 Male Midshipman (

Porichthys

) attract females by rapid contraction...

FIGURE 4.8 The Electric Eel,

Electrophorus electricus

, is closely related to...

FIGURE 4.9 Structure of fish skin. Image courtesy of Stephanie Anne Hunczak....

FIGURE 4.10 A cycloid scale (left) and a ctenoid scale (right). Note that th...

FIGURE 4.11 The Greater Amberjack,

Seriola dumerili

(Carangidae), exhibits c...

FIGURE 4.12 Fast start predators, such as pike, gars, and the Great Barracud...

FIGURE 4.13 Anterior, posterior, and lateral views of a Smooth Trunkfish

Lac

...

FIGURE 4.14 Nurse Sharks,

Ginglymostoma cirratum

, are a good example of a sh...

FIGURE 4.15 The two competing models that explain how horizontal locomotion ...

FIGURE 4.16 Moving on land – while out of the water amphibious air‐breathing...

FIGURE 4.17 Sailfin Flyingfish,

Parexocoetus brachypterus

, showing the enlar...

Chapter 5

FIGURE 5.1 Basic circulatory pattern found in most fishes.

FIGURE 5.2 Gills and blood vessels of the head of a cod (

Gadus

).

FIGURE 5.3 Main blood vessels of a bony fish.

FIGURE 5.4 Anterior arterial system in ventral view in the Scombridae, showi...

FIGURE 5.5 Crucian Carp are reported to increase the effective length of sec...

FIGURE 5.6 (A) Lateral views of the gill arches of the Walking Catfish (

Clar

...

FIGURE 5.7 Mudskippers, such as

Boleophthalmus pectinirostris

, can exchange ...

FIGURE 5.8 Oxygen dissociation curves. Vertical axes indicate the percent of...

FIGURE 5.9 Schematic representation of the gas bladders of a physostome (A) ...

FIGURE 5.10 Herbivorous fishes, such as Bermuda Chub (

Kyphosus sectatrix)

wh...

FIGURE 5.11 Variation in intestinal length and other features among carnivor...

FIGURE 5.12 Partitioning of the energy consumed by a fish. Only energy not r...

Chapter 6

FIGURE 6.1 Dorsal views of brains of representative fishes: (A) shark (Squal...

FIGURE 6.2 Dorsal view of a fish reflexively moving away from a sudden stimu...

FIGURE 6.3 Mechanoreception involves sensory hair cells, which are found in ...

FIGURE 6.4 The inner ear of fishes.

FIGURE 6.5 (A) Approximate location of lateral line in many fishes. (B) Cros...

FIGURE 6.6 Fishes demonstrate a continuum of hearing capabilities. Those wit...

FIGURE 6.7 Schematic diagram of the structure of ampullary (A) and tuberous ...

FIGURE 6.8 The elongate, flattened rostrum of the Paddlefish (

Polyodon spath

...

FIGURE 6.9 The small pores on the snout of this Tiger Shark (

Galeocerdo cuvi

...

FIGURE 6.10 The elephantnose fishes of Africa (

Gnathonemus

, Mormyridae) have...

FIGURE 6.11 Electric organs, often made of modified muscles along the trunk,...

FIGURE 6.12 The jamming avoidance response (JAR) of two

Eigenmannia virescen

...

FIGURE 6.13 Cross‐sectional view of the eye of a teleost.

FIGURE 6.14 The divided pupil and oblong lens of the four‐eyed fishes,

Anabl

...

FIGURE 6.15 Each eye of the deep‐sea Brownsnout Spookfish

(Dolichopteryx lon

...

FIGURE 6.16 A deep‐sea dragonfish with photophores that emit reddish light c...

FIGURE 6.17 (A) External view of the nares of a fish. (B) The obvious flap o...

FIGURE 6.18 The chin barbels of goatfishes (Mullidae) have a high density of...

Chapter 7

FIGURE 7.1 (A) The anal fin of a normal male

Gambusia

is elongated to form t...

FIGURE 7.2 Mummichog (

F. heteroclitus

) in the northern part of their range h...

FIGURE 7.3 (A) Atlantic Bluefin Tuna (

Thunnus thynnus

) rely on warm, efficie...

FIGURE 7.4 Maintaining osmotic balance in fresh versus seawater. (A) Freshwa...

FIGURE 7.5 (A) In addition to the abundant mitochondria that provide the ene...

FIGURE 7.6 The yellowish clusters are aggregations of macrophages (a type of...

Chapter 8

FIGURE 8.1 Patterns of sexual determination and maturation among fishes. Sex...

FIGURE 8.2 Bigger fish produce more eggs, both within and among species. Fis...

FIGURE 8.3 (A) Initial phase Stoplight Parrotfish,

Sparisoma viride

, have ch...

FIGURE 8.4 Terminal phase Bluehead Wrasse have the characteristic blue head ...

FIGURE 8.5 Parthenogenesis in Mexican livebearers. (A) In gynogenesis, a tri...

FIGURE 8.6 Kype (hooked lower jaw) on a migrating male land‐locked Atlantic ...

FIGURE 8.7 Breeding tubercles in fishes. Males of at least 25 different fami...

FIGURE 8.8 Turbidity‐influenced light transmission is causing hybridization ...

FIGURE 8.9 A male seahorse (family Syngnathidae) with a brood pouch full of ...

FIGURE 8.10 Male swordtail (

Xiphophorus helleri

, Poeciliidae) with an extend...

FIGURE 8.11 A pregnant female Shiner Perch,

Cymatogaster aggregata

, showing ...

Chapter 9

FIGURE 9.1 Events during, and terminology describing, the early life history...

FIGURE 9.2 Stages during the early life history of the Horse Mackerel,

Trach

...

FIGURE 9.3 Behavioral, physiological, and anatomical events during the poste...

FIGURE 9.4 A recently hatched, marine teleost larva, as represented by a 6 m...

FIGURE 9.5 Development of scales in the Black Crappie,

Pomoxis nigromaculatu

...

FIGURE 9.6 Juvenile anemonefish (

Amphiprion

) disperse from the anemone where...

FIGURE 9.7 (A) Approximate developmental and behavioral milestones of Atlant...

FIGURE 9.8 Larval diversity in marine fishes. Fish larvae often bear little ...

FIGURE 9.9 The general sequence of movement of marine larvae from offshore t...

FIGURE 9.10 Juvenile Brook Trout take greater risks to acquire food much nee...

FIGURE 9.11 (A) Atlantic Salmon parr in the Machias River, Maine. (B) An Atl...

FIGURE 9.12 Some landlocked Atlantic Salmon (

Salmo salar

) stocked as smolts ...

FIGURE 9.13 Sequence of events during smolting in the Coho Salmon,

Oncorhync

...

FIGURE 9.14 Progressive eye migration in a developing Summer Flounder,

Paral

...

FIGURE 9.15 (A) Metamorphosis from a pelagic to a benthic life in flatfishes...

FIGURE 9.16 Spiny dogfish (

Squalus acanthias

) is a commonly used dissection ...

FIGURE 9.17 Adults of the five species of Pacific salmon, such as this Socke...

FIGURE 9.18 The correspondence between growth zones on an otolith and habita...

FIGURE 9.19 Growth curves and their statistical description. The plotted lin...

FIGURE 9.20 Salmonids differ in the minimum size at which they develop the n...

Chapter 10

FIGURE 10.1 Regions and physical features of the deep‐sea environment relati...

FIGURE 10.2 Size differences in male versus female anglerfishes. A 6.2 mm pa...

FIGURE 10.3 Extreme movements of the head and mouth during swallowing in the...

FIGURE 10.4 An adult female Wolftrap Angler,

Lasiognathus amphirhamphus

(One...

FIGURE 10.5 A 50 cm long whipnose anglerfishes presumably foraging just abov...

FIGURE 10.6 United States Navy personnel hold a 23‐foot (7.0 m), 300 lb (140...

FIGURE 10.7 Open ocean and migratory coastal pelagic species of the Californ...

FIGURE 10.8 Keels and tails in open‐water fishes. The evolution of mackerels...

FIGURE 10.9 The eel‐like bodies of gunnels such as this Penpoint Gunnel,

Apo

...

FIGURE 10.10 The depressed, terete body shape of sculpins (family Cottidae) ...

FIGURE 10.11 Convergence in body form among unrelated fishes that occupy swi...

FIGURE 10.12 North and south polar regions. General oceanic circulation patt...

FIGURE 10.13 Body form and habitat types of common Antarctic nototheniid fis...

FIGURE 10.14 The Antarctic icefishes (Channichthyidae) are the only known ve...

FIGURE 10.15 Life cycle of annual cyprinodontoids, as shown by the Venezuela...

FIGURE 10.16 Devils Hole, Nevada, natural home of the Devils Hole Pupfish, t...

FIGURE 10.17 Cave fishes from three different orders, showing convergent los...

FIGURE 10.18

C. thamicola

, a waterfall‐climbing, cave‐dwelling fish from Tha...

Chapter 11

FIGURE 11.1 Periods of occurrence of major jawless fish taxa based on the fo...

FIGURE 11.2 Conodonts. (A) Conodont apparatus. The various elements (A–G) oc...

FIGURE 11.3 The earliest known fishes were jawless pteraspidomorphs with arm...

FIGURE 11.4 Other jawless fishes are placed in the superclasses Anaspidomorp...

FIGURE 11.5 Cephalaspidomorphs (superclass Osteostracomorphi) were diverse j...

FIGURE 11.6 One view of relationships among early agnathous fishes, modern j...

FIGURE 11.7 Periods of occurrence of major jawed (gnathostome) fish taxa bas...

FIGURE 11.8 Placoderms. (A) The coccosteomorph

Coccosteus

, (B) the ptyctodon...

FIGURE 11.9 Acanthodians. (A)

Climatius

, a primitive acanthodian with multip...

FIGURE 11.10 One view of relationships among bony fishes (class Osteichthyes...

FIGURE 11.11 Dorsal (A) and lateral (B) views of the reconstructed elpistost...

FIGURE 11.12 A cladogram of relationships among sarcopterygians and tetrapod...

FIGURE 11.13 Cladogram (strict consensus tree) of relationships among sarcop...

FIGURE 11.14 Phylogenetic relationships and fossil occurrence among the 24 c...

FIGURE 11.15 Extinct and extant lungfishes. (A)

Scaumenacia

, a Late Devonian...

FIGURE 11.16

Eusthenopteron foordi

, a well‐known osteolepidiform and member ...

FIGURE 11.17 Presumed key traits that characterized the sarcopterygian ances...

FIGURE 11.18 Comparative pelvic appendages of: (A)

Eusthenopteron

, a Devonia...

FIGURE 11.19 Actinopterygian fishes at different grades of development. (A)

FIGURE 11.20 Morphological (and ecological) convergence in fish evolution. P...

FIGURE 11.21

Thoracopterus magnificus

, a 6 cm‐long perleidiform from the Tri...

FIGURE 11.22

Leedsichthys problematicus

, perhaps the world’s largest fish ev...

FIGURE 11.23 Phylogenetic relationships among actinopterygian fishes. Pholid...

FIGURE 11.24 Diversification of the dorsal fin in modern teleosts. (A) Primi...

FIGURE 11.25 The phylogeny of paired fin locations in teleosts. The location...

FIGURE 11.26 Diversity in the body form of Paleozoic sharks from the two ext...

FIGURE 11.27 Examples of similar structure among some ancient and modern cho...

FIGURE 11.28 Tooth replacement in chondrichthyans. (A) Cross‐section through...

FIGURE 11.29 Extinct holocephalans. (A)

Ischyodus

, a Jurassic callorhinchid ...

Chapter 12

FIGURE 12.1 Phylogenetic relationships among living chondrichthyans. Relatio...

FIGURE 12.2 Taxonomic distribution and representative orders of over 1250 sp...

FIGURE 12.3 Reconstructing the jaws and estimating the size of the extinct M...

FIGURE 12.4 The Giant Freshwater Whipray

Himantura chaophraya

. This endanger...

FIGURE 12.5 Sawfishes are among the most imperilled marine and estuarine fis...

FIGURE 12.6 The role of scales in drag reduction in sharks. (A) Scanning ele...

FIGURE 12.7 The track of Nicole, a 4‐m White Shark that had been seen over a...

FIGURE 12.8 Head and jaw movements associated with feeding in the White Shar...

FIGURE 12.9 Cookie cutter sharks. (A)

Isistius brasiliensis

, the Cookie Cutt...

FIGURE 12.10 Representative tooth types of modern sharks: (A) Nurse Shark; (...

FIGURE 12.11 Pavement or molariform, crushing teeth characterize sharks and ...

FIGURE 12.12 The Megamouth Shark,

Megachasma pelagios

. This 4–5 m long zoopl...

FIGURE 12.13 Brain size in sharks. (A) Sharks have relatively large brains f...

FIGURE 12.14 Placental viviparity in advanced sharks. A new‐born Atlantic Sh...

FIGURE 12.15 Sexual dimorphism in the skin thickness of sharks. On the left ...

FIGURE 12.16 Modern holocephalans. (A)

Chimaera cubana

, a 50‐cm‐long Caribbe...

Chapter 13

FIGURE 13.1 An adult lancelet, Branchiostoma longirostrum; actual length, 43...

FIGURE 13.2 Hagfishes. (A) Adult Atlantic Hagfish,

Myxine glutinosa

, 38 cm l...

FIGURE 13.3 A single hagfish can produce prodigious quantities of slime when...

FIGURE 13.4 Lampreys. (A) An adult parasitic Sea Lamprey (

Petromyzon marinus

FIGURE 13.5 Reconstruction of the Carboniferous lamprey,

Mayomyzon pieckoens

...

FIGURE 13.6 Comparative life histories of a species pair of European lamprey...

FIGURE 13.7 The living African Coelacanth,

Latimeria chalumnae

, an extant me...

FIGURE 13.8 Marjorie Courtenay‐Latimer’s drawing and description of the firs...

FIGURE 13.9 Coelacanths are as cuddly as pandas. (A) The Coelacanth Conserva...

FIGURE 13.10 Modern lungfishes. (A) An African lungfish,

Protopterus annecte

...

FIGURE 13.11 An African lungfish estivating in its mud and mucus cocoon, vie...

FIGURE 13.12 A young African lungfish. The arrow indicates the external gill...

FIGURE 13.13 A seventh lungfish,

Ompax spatuloides

? This is the illustration...

FIGURE 13.14 (A) A 29 cm long bichir,

Polypterus palmas polli

, from the Ivor...

FIGURE 13.15 Sturgeons. (A) An Atlantic Sturgeon,

Acipenser oxyrhynchus

. Not...

FIGURE 13.16 Paddlefishes. (A) The North American Paddlefish,

Polyodon spath

...

FIGURE 13.17 Gars. (A) A Spotted Gar,

Lepisosteus oculatus

, showing the dist...

FIGURE 13.18 An Alligator Gar caught in Louisiana by the Baton Rouge Fish an...

FIGURE 13.19 (A) The Bowfin,

Amia calva

, a member of a monotypic order endem...

Chapter 14

FIGURE 14.1 Increased genetic information in recent decades has resulted in ...

FIGURE 14.2 Elopomorphs. (A) A Tarpon,

Megalops atlanticus

. Citron / Wikimed...

FIGURE 14.3 A Gulper or Pelican Eel,

Eurypharynx pelecanoides

. Ironically, t...

FIGURE 14.4 Osteoglossomorphs. (A) A mormyrid elephantfish,

Gnathonemus pete

...

FIGURE 14.5 The Asian Arawana or Golden Dragonfish,

Scleropages formosus

. Ov...

FIGURE 14.6

Knightia alta

, an Eocene herring from the Green River formation ...

FIGURE 14.7 The giant and imperiled Asian Carp,

Catlocarpio siamensis

, nativ...

FIGURE 14.8 The extinct Harelip Sucker,

Moxostoma lacerum

. Once abundant in ...

FIGURE 14.9  A juvenile Robust Redhorse,

Moxostoma robustum

. Growing to larg...

FIGURE 14.10 Piranhas,

Serrasalmus

spp., are representative of the speciose ...

FIGURE 14.11 Selected catfishes, showing some of the array of body types and...

FIGURE 14.12 Large catfishes. (A) The Mekong Giant Catfish, one of the world...

FIGURE 14.13 Two individual

Orthosternarchus tamandua

, an apteronotid knifef...

FIGURE 14.14  Gymnotiform electric knifefishes – an Electric Eel (

Electropho

...

FIGURE 14.15 Protacanthopterygians. (A) An osmerid, the Capelin,

Mallotus vi

...

FIGURE 14.16 Neck flexibility in the Australian salamanderfish,

Lepidogalaxi

...

FIGURE 14.17 Phylogeny of the salmonids. A cladogram of most living salmonid...

FIGURE 14.18 A Golden Trout,

Oncorhynchus aguabonita

, the California state f...

FIGURE 14.19 The esocid Muskellunge,

Esox masquinongy

, one of the largest pr...

FIGURE 14.20 Aulopiforms, from deep and shallow water. (A) Giganturid telesc...

FIGURE 14.21 A Spothead Lanternfish,

Diaphus metopoclampus

. The round struct...

FIGURE 14.22 Lampriformes. (A) An opah or moonfish,

Lampris regius

. This lar...

FIGURE 14.23 Paracanthopterygiians. (A) A percopsid, the Trout‐perch,

Percop

...

FIGURE 14.24  Spawning behavior of the Pirate Perch,

Aphrododerus sayanus

, a...

Chapter 15

FIGURE 15.1 Three different interpretations of the relationships among the a...

FIGURE 15.2 Berycipmorphs. Red light does not penetrate to the depths at whi...

FIGURE 15.3 Formerly included among the beryciphorms, squirrelfishes (family...

FIGURE 15.4 (A) A Pacific Pearlfish,

Encheliophis dubius

, entombed in a blac...

FIGURE 15.5 Batrachoidiforms – The Oyster Toadfish (

Opsanus tao

) is easily v...

FIGURE 15.6

(A)

A striped mullet,

Mugil cephalus

. From Jordan (1905 ). (B) A ...

FIGURE 15.7 Atherinomorphs. (A) A belonid needlefish,

Tylosurus crocodilus

. ...

FIGURE 15.8 Gasterosteiforms and Syngnathiforms. (A) A Three‐spined Stickleb...

FIGURE 15.9 Synbranchiforms such as the Asian Swamp Eel (

Monopterus albus

) a...

FIGURE 15.10 Scorpaeniforms. (A) A Canary Rockfish,

Sebastes pinniger

, one o...

FIGURE 15.11 Representative percoid fishes. (A) A centropomid Snook,

Centrop

...

FIGURE 15.12 (A) The tangerine darter,

Percina aurantiaca

, the second larges...

FIGURE 15.13 Banggai Cardinalfish, endemic to the Banggai Islands of Indones...

FIGURE 15.14 (A) Dolphinfish, dorado, or mahimahi,

Coryphaena hippurus

, exhi...

FIGURE 15.15 (A) This female Cobia,

Rachycentron canadum

(Rachycentridae), w...

FIGURE 15.16 (A) A Mutton Snapper (

Lutjanus analis

, Lutjanidae) resting behi...

FIGURE 15.17 Butterflyfishes (A, Chaetodontidae,) and angelfishes B, (Pomaca...

FIGURE 15.18 A sphyraenid, the Great Barracuda,

Sphyraena barracuda

.

FIGURE 15.19 (A) Diversity in body shape among African cichlids. These fishe...

FIGURE 15.20 Pomacentrid damselfishes. (A) the Sergeant Major,

Abudefduf sax

...

FIGURE 15.21 Representative labroids. (A) A California Sheephead (

Semicossyp

...

FIGURE 15.22 Zoarcoids. (A) A zoarcoid Atlantic wolffish,

Anarhichas lupus

. ...

FIGURE 15.23 The Patagonian Toothfish,

Dissostichus eleginoides

, a large, lo...

FIGURE 15.24 There are over 400 species in the family Blenniidae; the fish i...

FIGURE 15.25 Clingfishes (A, family Gobiesocidae) and many gobies (B, family...

FIGURE 15.26 A – A Foxface Rabbitfish (

Siganus vulpinus

, Siganidae).; B‐...

FIGURE 15.27 (A) Striped Marlin (

Kajikia audax

) are among the larger members...

FIGURE 15.28 Snakeheads. (A) The Northern Snakehead,

Channa argus

. This Sout...

FIGURE 15.29 A pleuronectiform, the bothid Fourspot Flounder,

Paralichthys o

...

FIGURE 15.30 Lophiiformes. (A) Striated Frogfish (

Antennarius striatus

, fami...

FIGURE 15.31 Ceratioid anglerfishes. Shown are females from the 11 ceratioid...

FIGURE 15.32 Tetraodontiforms. (A) A Gray Triggerfish,

Balistes capriscus

(f...

Chapter 16

FIGURE 16.1 “Aggressive” mimicry in the Striated Frogfish

Antennarius striat

...

FIGURE 16.2 Variations on a theme: convergence in morphology among fast star...

FIGURE 16.3 Some fishes have evolved morphological and behavioral traits tha...

FIGURE 16.4 Some fishes avoid detection by predators by blending in with obj...

FIGURE 16.5 The Comet (

Calloplesiops altivelis

) is a tropical reef fish with...

FIGURE 16.6 Examples and functions of disruptive coloration in fishes. (A) T...

FIGURE 16.7 The functional morphology of mirror sides in fishes. (A) A clear...

FIGURE 16.8 (A) The advantage to fishes of hovering in shade. On a sunny day...

FIGURE 16.9 Countershading, silvery sides, and turbidity from phytoplankton ...

FIGURE 16.10 Is this the face of death? (A) The general features of a predat...

FIGURE 16.11 Lionfishes, native to Indo‐Pacific regions, are among the few f...

FIGURE 16.12 Predator avoidance has shaped body morphology in zooplanktivoro...

FIGURE 16.13 Flying fish become airborne to escape predators. NOAA / Public ...

FIGURE 16.14 A 3 m long hammerhead shark captured and bit a piece off of a l...

FIGURE 16.15 Examples of the responses of minnows under attack. Responses in...

FIGURE 16.16 Sharp, blade‐like teeth of the Great Barracuda allow it to cut ...

FIGURE 16.17 When members of the family Diodontidae, such as pufferfish, bal...

Chapter 17

FIGURE 17.1 Spawning nests or bowers of African cichlids. Male cichlids cons...

FIGURE 17.2 Some of the cods (Gadidae) are known to vocalize during courtshi...

FIGURE 17.3 Indigo Hamlets (

Hypoplectrus indigo

) are simultaneous hermaphrod...

FIGURE 17.4 Spawning frequently involves multiple males and a single female....

FIGURE 17.5 Fertilization occurs in the mouth of some female African cichlid...

FIGURE 17.6 Well‐developed (near‐term) embryos in the ovary of a Mexican goo...

FIGURE 17.7 (A) A preserved male Nurseryfish,

Kurtus gulliveri

, with eggs at...

FIGURE 17.8 Parental care in the Spraying Characin,

Copella

sp. Eggs in this...

FIGURE 17.9 Parental care in cichlid fishes. Two of the more striking forms ...

FIGURE 17.10 (A) Cuckoo Catfish pairs (darker fish) follow close behind spaw...

FIGURE 17.11 Variations in life history and behavior result in alternative m...

FIGURE 17.12 Convict Cichlids raise young as socially monogamous pairs of pa...

FIGURE 17.13 Parthenogenesis has only rarely been observed among vertebrates...

FIGURE 17.14 A nighttime photograph of a male ponyfish,

Leiognathus elongatu

...

FIGURE 17.15 Some temperate freshwater fishes rival tropical fishes for brig...

FIGURE 17.16 Lateral and frontal displays in fishes. During agonistic intera...

FIGURE 17.17 Exaggerated swimming display of the Gray Reef Shark,

Carcharhin

...

FIGURE 17.18 Types and activities of fish aggregations. Shoals contain fish ...

FIGURE 17.19 Garden eels (Congridae) live in colonies of several hundred ind...

FIGURE 17.20 A Yellowstripe Goatfish,

Mulloidichthys flavolineatus

at Kona, ...

FIGURE 17.21 Some gobies live in mutualistic relationships with burrowing sh...

FIGURE 17.22 Anemonefish (subfamily Amphiprioninae) move among the tentacles...

Chapter 18

FIGURE 18.1 Day (A) versus night (B) distribution of families in a rocky ree...

FIGURE 18.2 A daytime resting school of juvenile French and White Grunts hov...

FIGURE 18.3 Light availability, dark adaptation, species changeover, and pre...

FIGURE 18.4 Zooplantivorous fishes often follow the vertical migrations of t...

FIGURE 18.5 Thin, elongate fishes, such as this Monkeyface Prickleback,

Cebi

...

FIGURE 18.6 Endogenous, circatidal activity rhythm of Shanny,

Lipophrys phol

...

FIGURE 18.7 Genetically based, local adaptation in Fraser River Sockeye Salm...

FIGURE 18.8 Sticklebacks have a daily rhythm of photosensitivity that is max...

FIGURE 18.9 The seasonal progression of events for many fishes in large trop...

FIGURE 18.10 Diadromy takes three general forms: anadromy, catadromy, and am...

FIGURE 18.11 Many familiar diadromous fishes are either anadromous (such as ...

FIGURE 18.12 The latitudinal distribution and frequency of different forms o...

FIGURE 18.13 Pacific salmon begin their lives in streams, move downstream in...

FIGURE 18.14 Characteristic life history of a Pacific salmon, as seen in the...

FIGURE 18.15 Yellow phase American eels in a cave spring in northern Florida...

Chapter 19

FIGURE 19.1 The earth’s important tectonic features. Dashed lines denote mar...

FIGURE 19.2 Cladogram of the

Scomberomorus regalis

group of Spanish mackerel...

FIGURE 19.3 Ranges of the regalis group of Spanish mackerels (

Scomberomorus

)...

FIGURE 19.4 A neighbor joining tree (A) and parsimony network (B) based on m...

FIGURE 19.5 The sardines support the dispersal model of fish speciation and ...

FIGURE 19.6 Distribution of marine populations of 11 species of needlefishes...

FIGURE 19.7 Biogeographic provinces of the shallow tropical Indo‐Pacific as ...

FIGURE 19.8 The ranges of seven Indian‐West Pacific species of Spanish macke...

FIGURE 19.9 Distribution of coral reef fishes (shaded areas) in the tropical...

FIGURE 19.10 Parsimony network for mtDNA cytochrome

b

showing that the Golds...

FIGURE 19.11 Map of the Indo‐Pacific showing warm‐temperate biogeographic pr...

FIGURE 19.12 Main ichthyological provinces in North and Middle America (from...

FIGURE 19.13 South American ichthyological provinces (from Lévêque et al. 20...

FIGURE 19.14 Main ichthyological provinces in Africa (based on Lévêque & Pau...

FIGURE 19.15 (A) Main ichthyological provinces in western Europe (from Lévêq...

FIGURE 19.16 (A) Wallace’s and Weber’s lines separating the Oriental and Aus...

FIGURE 19.17 Osteoglossidae and their distribution (inset below). (A)

Hetero

...

FIGURE 19.18 Summary of archaic freshwater fish distributions.

FIGURE 19.19 Distribution of the pickerels, Esocidae.

FIGURE 19.20 Distribution of the darters and perches, Percidae.

FIGURE 19.21 Distribution of the sunfishes (Centrarchidae) and cichlids (Cic...

FIGURE 19.22 Summary of recent distributions of primary freshwater fishes of...

FIGURE 19.23 Distribution of the minnows and carps, Cyprinidae.

FIGURE 19.24 Distribution of the suckers, Catostomidae.

FIGURE 19.25 Summary of otophysan distribution.

FIGURE 19.26 Distributional limits of certain primary and secondary freshwat...

FIGURE 19.27 Phylogeographic data for the Lake Whitefish (

Coregonus clupeafo

...

FIGURE 19.28 The relationship between nucleotide diversity and latitude for ...

FIGURE 19.29 Drainage basins of western North America that support at least ...

Chapter 20

FIGURE 20.1 Bigger fish produce more eggs, both within and among species. Si...

FIGURE 20.2 Strong selection against large adults in a fish population, such...

FIGURE 20.3  (A) Many fishes, such as the Pumpkinseed sunfish (

Lepomis gibbo

...

FIGURE 20.4 Four major groupings of commonly co‐occurring life history chara...

FIGURE 20.5 Heavy fishing pressure has pushed the evolution of harvested spe...

FIGURE 20.6 The Cui‐ui,

Chasmistes cujus

(Catostomidae), is an endangered su...

FIGURE 20.7 Largemouth Bass demonstrate ontogenetic niche shifts as they gro...

FIGURE 20.8 Four morphs of Arctic Char that differ anatomically, behaviorall...

FIGURE 20.9 American Eel (

Anguilla rostrata)

are distributed in coastal and ...

FIGURE 20.10 Seahorses and other syngnathids show limited dispersal. This li...

Chapter 21

FIGURE 21.1 Foraging guilds of soft bottom fishes on the southern California...

FIGURE 21.2 Habitat choice in stream fishes as demonstrated by vertical segr...

FIGURE 21.3 The characid Dorado (

Salminus brasiliensis

), a popular gamefish ...

FIGURE 21.4 Goliath Grouper, like many marine fishes, occupy productive insh...

FIGURE 21.5 Vertical zonation of fishes in a kelp bed. Species portrayed are...

FIGURE 21.6 A triggerfish, wrasse, and various damselfishes in the Maldives....

FIGURE 21.7 The various processes that operate to determine the diversity an...

FIGURE 21.8 Priority effects. The foraging behavior of species that settle i...

FIGURE 21.9 Bluegill, Pumpkinseed, and Green Sunfish (photo) eat similar foo...

FIGURE 21.10 Oceanic prey fishes are often trapped between predatory birds f...

FIGURE 21.11 Survivorship curves in theory and practice. (A) Three general f...

FIGURE 21.12 (A) A River Otter (

Lontra canadensis)

consumes a Saddleback Gun...

FIGURE 21.13 The distribution of prey fishes in shallow streams reflects the...

FIGURE 21.14 Growth of the predatory dinoflagellate,

Pfiesteria piscicida

, r...

FIGURE 21.15 Evidence of Sea Lamprey (

Petromyzon marinus

) feeding on an Atla...

FIGURE 21.16 Relative numbers and types of herbivores as a function of latit...

FIGURE 21.17 Reef‐dwelling rabbitfish have narrow, elongated snouts which al...

FIGURE 21.18 Food webs involving ciguatera fish poisoning in a reef communit...

FIGURE 21.19 Damselfish territoriality helps maintain high algal species div...

FIGURE 21.20 Algal productivity increases in relation to density of inverteb...

FIGURE 21.21 Effects of fish predation on zooplankton assemblage structure. ...

FIGURE 21.22 Ecosystem level implications of overfishing on coral reefs. Ove...

FIGURE 21.23 Trophic cascades. (A) The various components of a trophic casca...

FIGURE 21.24 Disruptions of large‐scale, oceanic trophic cascades can result...

FIGURE 21.25 The relationship between parrotfish grazing and algal growth in...

FIGURE 21.26 A trophic cascade with unexpected results. Extensive gill‐netti...

FIGURE 21.27 Food webs involving fishes. (A) A relatively simple food web in...

FIGURE 21.28 Salmon‐derived nutrients and food webs based on anadromous fish...

FIGURE 21.29 (A) Bite marks on a Pacific coral head from parrotfish grazing....

FIGURE 21.30 Yellowfin shiner (

Notropis lutipinnus

) spawning over a nest mou...

FIGURE 21.31 A pictorial summary of ecosystem services provided by fishes to...

Chapter 22

FIGURE 22.1 Extinction rates and causes. (A) Fish extinctions in North Ameri...

FIGURE 22.2 Is the rare Devils Hole Pupfish a genetically distinct species, ...

FIGURE 22.3 Impacts of bottom trawling on gravel (A) and mud (B) habitats, S...

FIGURE 22.4 The U.S. Fish and Wildlife Service placed logs in Redd Creek cha...

FIGURE 22.5 Endangered fishes of the upper Colorado River. Prior to impoundm...

FIGURE 22.6 Commercial catches of Chinook Salmon and Steelhead (sea‐run Rain...

FIGURE 22.7 The Snail Darter (

Percina tanasi

) rose to fame when its discover...

FIGURE 22.8 Effects of suspended sediments on young fishes. Threatened Spotf...

FIGURE 22.9 The Aral Sea, once the fourth largest lake in the world, is a me...

FIGURE 22.10 Both native and non‐native species are involved in faunal homog...

FIGURE 22.11 Brown Trout (

Salmo trutta

) are among the most widely transplant...

FIGURE 22.12 Effects of Nile Perch introduction on the food web of Lake Vict...

FIGURE 22.13 Lionfishes (

Pterois

, Scorpaenidae) are now common on tropical w...

FIGURE 22.14 Experimental evidence of competitive displacement of a native b...

FIGURE 22.15 The elongated and barbed abdominal spine of the Spiny and Fishh...

FIGURE 22.16 Trends in various indicators of abundances of Northeast Atlanti...

FIGURE 22.17 The Gulf of California and a reconstruction of the presumed sea...

FIGURE 22.18 Evolution of body shape in exploited Lake Whitefish in Lesser S...

FIGURE 22.19 Fishing down food webs. Since about 1950, most of the world’s m...

FIGURE 22.20 (A) Atlantic salmon net pens in a fjord in Norway. In 2012, Nor...

FIGURE 22.21 Aquarium collecting of tropical fishes such as the Achilles Tan...

FIGURE 22.22 The complementary roles of three scientific fields (phylogeneti...

FIGURE 22.23 Functions of marine protected areas from a fisheries management...

Guide

Cover Page

Title Page

Copyright Page

Dedication

Preface to the Third Edition

Preface to the Second Edition

Preface to the First Edition

Acknowledgments

About the Companion Website

Table of Contents

Begin Reading

Appendix I Appendix 1 ‐ Glossary of Genetic/Molecular Terminology

References

Index

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The Diversity of Fishes

Biology, Evolution and Ecology

Third Edition

DOUGLAS E. FACEY

Emeritus Professor of Biology, Saint Michael’s College, Vermont, USA

BRIAN W. BOWEN

Research Professor, Hawaii Institute of Marine Biology, University of Hawaii at Manoa, Honolulu, Hawaii, USA

BRUCE B. COLLETTE

Research Associate, NOAA, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA

GENE S. HELFMAN

Emeritus Professor of Ecology, Odum School of Ecology, University of Georgia, Athens, Georgia, USA

This edition first published 2023© 2023 John Wiley & Sons Ltd

Edition HistoryGene S. Helfman, Bruce B. Collette, Douglas E. Facey (1e, 1997); Gene S. Helfman, Bruce B. Collette, Douglas E. Facey, and Brian W. Bowen (2e, 2009)

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Library of Congress Cataloging‐in‐Publication DataNames: Facey, Douglas E., author. | Bowen, Brian W. (Brian William), 1957‐ author. | Collette, Bruce B., author. | Helfman, Gene S., author.Title: The diversity of fishes : biology, evolution and ecology / Douglas E. Facey, Brian W. Bowen, Bruce B. Collette, Gene S. Helfman.Description: Third edition. | Hoboken, NJ : Wiley, 2023. | Revised edition  of: The diversity of fishes / Gene Helfman … [et al.] 2nd ed. 2009. |  Includes bibliographical references and index.Identifiers: LCCN 2022030874 (print) | LCCN 2022030875 (ebook) | ISBN  9781119341918 (cloth) | ISBN 9781119341802 (adobe pdf) | ISBN  9781119341833 (epub) Subjects: LCSH: Fishes. | Fishes—Variation. | Fishes—Adaptation.Classification: LCC QL615 .F33 2023 (print) | LCC QL615 (ebook) | DDC  597–dc23/eng/20220810LC record available at https://lccn.loc.gov/2022030874LC ebook record available at https://lccn.loc.gov/2022030875

Cover Design: WileyCover Images: Courtesy of Gene Helfman; Luiz A. Rocha, California Academy of Sciences; Jon Hyde & Kimberly Sultze, www.hydesultze.com; Andrew Nagy

To our parents, for their encouragement of our nascent interest in things biological;To our spouses – Janice, RuthEllen, Sara, and Judy – for their patience and understanding during the production of this volume;And to students, colleagues, and fish lovers for their efforts to preserve biodiversity for future generations.

Preface to the Third Edition

Thank you to all who helped make the first two editions of The Diversity of Fishes successful. We greatly appreciate the feedback and suggestions from those who have used them. And to those who have been asking us about an update – thanks for your patience. Here is what you can expect in the third edition.

We have done our best to update information while recognizing that no textbook can represent the most current information in any large field. This book is intended as a starting point, not the final destination. Therefore, we encourage those using the text to consult more current sources for updated information.

This edition includes more color photographs to better demonstrate the diversity and beauty of fishes that attract many to the field. We include photographs from public agencies, in part to acknowledge the valuable contributions to Ichthyology of those in natural resources and management organizations.

Reorganization and consolidation of some topics has reduced the total number of chapters to 22.

Each chapter begins with a Summary that provides a broad overview of the content of that chapter. This may be particularly useful for those using the text for a course and who do not intend to utilize some chapters in detail – students can read the summary of each of those chapters.

Molecular genetics has transformed many aspects of ichthyology over the last few decades, and this is reflected throughout the text. Important concepts are introduced in

Chapters 1

and

2

, supported by an Appendix of terminology at the end of the book, and specific contributions of molecular genetics to the field of ichthyology are included in many chapters.

Structure and function are addressed together rather than treated in separate chapters. For example,

Chapter 3

addresses structure and function of the head (e.g. bones, muscles, breathing, jaw suspension, feeding, dentition), and

Chapter 4

addresses structure and function of the trunk (e.g. bones, muscles, integument, scales, fins, locomotion).

In chapters addressing the history of the fishes (

Chapter 11

) and systematics of the major groups of fishes (

Chapters 12

15

), we have largely adopted the approach of Nelson et al. (2016), which includes consideration of molecular phylogenetics. However, we have retained the approach of Nelson (2006) where conclusions from molecular data are quite different from those based on morphological evidence (see

Chapter 15

, for example). We note differences between the approaches and encourage readers to consult current sources for updated information and perspectives, as our collective understanding of the relationships among groups of fishes continues to evolve.

Fishes as predators and prey are considered together in one chapter (

Chapter 16

). Many fishes are both predators and prey, and many physical and behavioral adaptations are rather similar and, in some cases, may have developed in response to one another.

Our chapter on Zoogeography and Phylogeography (

Chapter 19

) uses global maps that more accurately represent the comparative sizes of oceans and landmasses than maps used in prior editions.

Fish populations are covered in

Chapter 20

, including Population Ecology, Population Dynamics and Regulation, and Population Genetics.

Fish interactions with other species within fish assemblages and broader communities, and their impacts on ecosystems are addressed in

Chapter 21

. This is a long chapter but avoids the redundancy of addressing similar types of interactions and impacts in multiple chapters.

As in the past, we conclude with a chapter on Conservation (

Chapter 22

), but this now includes Conservation Genetics.

Preface to the Second Edition

The first edition of The diversity of fishes was successful beyond our wildest dreams. We have received constant and mostly positive feedback from readers, including much constructive criticism, all of which convinces us that the approach we have taken is satisfactory to ichthyological students, teachers, and researchers. Wiley-Blackwell has validated that impression: by their calculations, The diversity of fishes is the most widely adopted ichthyology textbook in the world.

However, ichthyology is an active science, and a great deal of growth has occurred since this book was first published in 1997. Updates and improvements are justified by active and exciting research in all relevant areas, including a wealth of new discoveries (e.g., a second coelacanth species, 33 more megamouth specimens, several new record tiniest fishes, and exciting fossil discoveries including some that push back the origin of fishes many million years and another involving a missing link between fishes and amphibians), application of new technologies (molecular genetics, transgenic fish), and increased emphasis on conservation issues (e.g., Helfman 2007). Websites on fishes were essentially nonexistent when the first edition was being produced; websites now dominate as an instant source of information. Many of the volumes we used as primary references have themselves been revised. Reflective of these changes, and of shortcomings in the first edition, is the addition of a new chapter and author. Genetics received insufficient coverage, a gross omission that has been corrected by Brian Bowen‛s contribution of a chapter devoted to that subject and by his suggested improvements to many other chapters. Brian‛s contributions were aided by extensive and constructive comments from Matthew Craig, Daryl Parkyn, Luiz Rocha, and Robert Toonen. He is especially grateful to John Avise, Robert Chapman, and John Musick for their guidance and mentorship during his professional career, and most of all to his wife, RuthEllen, for her forbearance and support.

Among the advances made in the decade following our initial publication, a great deal has been discovered about the phylogeny of major groups, especially among jawless fishes, sarcopterygians, early actinopterygians, and holocephalans. In almost all taxa, the fossil record has expanded, prompting reanalysis and sometimes culminating in conflicting interpretations of new findings. A basic textbook is not the appropriate place to attempt to summarize or critique the arguments, opinions, and interpretations. We have decided to accept one general compilation and synthesis. As in the 1997 edition, where we adopted with little adjustment the conclusions and terminology of Nelson (1996), we here follow Nelson (2006), who reviews the recent discoveries and clearly presents and assesses the many alternative hypotheses about most groups. Instructors who used our first edition will have to join us in learning and disseminating many changed names as well as rearrangements among taxa within and among phylogenies, especially Chapters 11–13. Science is continually self-correcting. We should applaud the advances and resist the temptation to comfortably retain familiar names and concepts that have been modified in light of improved knowledge.

Also, we have now adopted the accepted practice of capitalizing common names.

Acknowledgments

Thanks especially to the many students and professionals who corrected errors in the first edition (J. Andrew, A. Clarke, D. Hall, G.D. Johnson, H. Mattingly, P. Motta, L.R. Parenti, C. Reynolds, C. Scharpf, E. Schultz, M.L.J. Stiassny, and S. Vives proved particularly alert editors). Their suggestions alone led to many changes, to which we have added literally hundreds of new examples, facts, and updates. Wiley-Blackwell has provided a website for this second edition, www.wiley.com/go/helfman