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Vertebrate Life


Eleventh Edition

Harvey Pough, Author William E. Bemis, Author Betty Anne McGuire, and Author Christine M. Janis

14 November 2022

ISBN: 9780197558621

656 pages

In Stock

Price: £110.99

If you are a lecturer interested in adopting this title for your course, please contact your local campus representative to arrange a local price.


Widely praised for its comprehensive coverage and exceptionally clear writing style, this best-selling text explores how the anatomy, physiology, ecology, and behavior of animals interact to produce organisms that function effectively in their environments and how lineages of organisms change through evolutionary time.

  • Incorporation of the dagger symbol (†) to indicate extinct taxa, a helpful convention in a field that requires constant comparisons between extinct and extant taxa
  • Includes more than 2,000 photographs and illustrations, of which about 800 are new
  • Consistent phylogenetic approach

New to this edition

  • Learning Objectives presented at the start of each section within a chapter to highlight key take-away concepts for the section
  • 45 new interactive figures in the Enhanced e-book
  • Re-development of background materials for the study of vertebrate evolution: Exemplified by Figure 1.2, and the many new time trees and cladograms
  • Many updated examples throughout the book use key fossils vertebrates to explore new phylogenetic interpretations and explain how particular anatomical features, such as the mammalian middle ear, evolved (see Figure 22.11)
  • Reorganization of key topics to improve continuity and connections among chapters along with extensive new cross-referencing to link topics between chapters
  • Enhanced coverage and reorganization of chapters on sharks (Chapter 6), ray-finned fishes (Chapter 7), sarcopterygians (Chapter 8), early tetrapods, lissamphibians, and amniotes (Chapter 9), dinosaurs (Chapter 18 and 19), synapsids (Chapter 22), therians (Chapter 23), and primates (Chapter 24)
  • Consolidation and reorganization of chapters to better highlight the role of thermoregulation and comparative physiology in vertebrate evolution (Chapter 14)
  • Incorporation of new behavioral and ecological information throughout the book to highlight the lives of vertebrates
  • Updated Discussion Questions at the end of each chapter
  • A selection of new and relevant references for each chapter to help users connect with the constant stream of new and exciting information on vertebrate biology
  • Development of new and informative labels that appear directly on the figures, providing instant captions to orient students and serve instructors who depend on illustrations for teaching
  • "Balloons" call attention to key points in many figures

About the Author(s)

Harvey Pough, Author William E. Bemis, Author Betty Anne McGuire, and Author Christine M. Janis

F. Harvey Pough, Professor Emeritus at Rochester Institute of Technology, is a herpetologist, specializing in environmental and evolutionary physiology, a past president of the American Society of Ichthyologists and Herpetologists, and the senior author of textbooks on Herpetology and Vertebrate Zoology. He has taught courses in Animal Behavior, Ecology, Herpetology, Human Biology, Introductory Biology, Physiological Ecology, and Vertebrate Zoology.

William E. Bemis is Professor of Ecology and Evolutionary Biology at Cornell University and Faculty Curator of Ichthyology at the Cornell University Museum of Vertebrates. He has studied the anatomy, systematics, and evolution of extant and fossil vertebrates for 50 years with a focus on fishes. He currently teaches Vertebrate Biology, Ichthyology, and Herpetology.

Betty McGuire is a retired Senior Lecturer from the Department of Ecology and Evolutionary Biology at Cornell University. She has studied social behavior, reproduction, and ecology of small mammals and currently studies behavior of domestic dogs. She coauthored textbooks on Animal Behavior and Human Biology, and taught courses in Vertebrate Biology, Mammalogy, Human Biology, Animal Behavior, Evolution, and Introductory Biology.

Christine M. Janis is Professor Emerita at Brown University, USA, and currently an Honorary Professor at the University of Bristol, UK. She is a mammalian paleobiologist who has studied the feeding and locomotion of Cenozoic mammals, especially ungulates (hoofed mammals) and kangaroos, and their paleobiology in the context of climatic and environmental change. She has taught courses in Comparative Anatomy and Vertebrate Paleontology.

Table of Contents

    Preface xv
    Chapter 1
    Diversity, Classification, and Evolution of Vertebrates
    1.1 The Vertebrate Story
    —Binominal nomenclature
    —Extant vertebrate groups
    1.2 Phylogenetic Systematics
    1.3 Applying Phylogenetic Criteria
    —Evaluating possible phylogenies
    —Molecules and morphology
    —The problem of dating
    —Dagger (†) convention adopted in this book
    1.4 Using Phylogenetic Trees
    —Extant phylogenetic brackets
    —Crown and stem groups
    1.5 Genetic Mechanisms of Evolutionary Change
    —Phenotypes and fitness
    —Developmental regulatory genes
    1.6 Epigenetic Effects
    1.7 Earth History and Vertebrate Evolution
    Chapter 2
    What Is a Vertebrate??
    2.1 Vertebrates in Relation to Other Animals?
    2.2 Characteristics of Chordates?
    —Chordate origins and evolution?
    —Extant nonvertebrate chordates?
    2.3 What Distinguishes a Vertebrate??
    2.4 Vertebrate Embryonic Development?
    —Development of the body?
    —Development of the pharyngeal region?
    —Development of the brain?
    —Other neurogenic tissues of vertebrates?
    2.5 Vertebrate Tissues?
    —Adult tissue types
    —Mineralized tissues
    2.6 Vertebrate Organ Systems
    —Integumentary system
    —Skeletal system
    —Muscular system
    —Nervous system and sense organs
    —Endocrine system
    —Respiratory system
    —Circulatory system
    —Digestive system
    —Excretory and reproductive systems
    Chapter 3
    Jawless Vertebrates and the Origin of Gnathostomes
    3.1 Earliest Evidence of Vertebrates
    —Enigmas: †Conodonts and †Tullimonstrum
    —-Early mineralized tissues
    —Environment of early vertebrate evolution
    3.2 Cyclostomes: Extant Jawless Vertebrate
    —Characters of cyclostomes
    —Hagfishes: Myxiniformes
    —Lampreys: Petromyzontiformes
    3.3 Jawless Osteognathostomes
    3.4 Gnathostome Body Plan
    —Gnathostome skeletons
    —What about soft anatomical features?
    3.5 Origin of Jaws
    —Hypotheses of jaw origins
    —Importance of the nose
    —Selective value of jaws
    3.6 Origin of Paired Appendages
    —Fin development and the lateral somitic frontier
    —Advantages of fins
    3.7 Extinct Paleozoic Jawed Fishes
    Chapter 4
    Living in Water
    4.1 Aquatic Environment
    —Obtaining oxygen from water using gill
    —Obtaining oxygen from air using lungs and other respiratory structure
    —Adjusting buoyancy
    4.2 Sensory World of Aquatic Vertebrates
    —Chemosensation: Olfaction and taste
    —Detecting water displacement
    —Hearing and equilibrium
    —Electroreception and electrogenesis
    4.3 Maintaining an Internal Environment
    —Nitrogenous wastes and kidney
    —Regulation of ions and body fluids
    4.4 Osmoregulation in Different Environments
    —Marine cartilaginous fishes and coelacanths
    —Marine teleosts
    —Freshwater teleosts and lissamphibians
    —Euryhaline vertebrates
    Chapter 5
    Geography and Ecology of the Paleozoic
    5.1 Deep Time
    —The Precambrian world
    —The Paleozoic
    5.2 Continental Geography
    —Continental drift and plate tectonics
    —Shifting continents of the Paleozoic
    —Shifting continents and changing climates
    5.3 Paleozoic Climates
    5.4 Paleozoic Ecosystems
    —Aquatic life
    —Terrestrial flora
    —Terrestrial fauna
    5.5 Extinctions
    Chapter 6
    Origin and Radiation of Chondrichthyans
    6.1 Acanthodii
    6.2 Chondrichthyes
    —Habitats and diversity
    —Placoid scales
    —Cartilaginous skeleton
    —Teeth and tooth plates
    —Jaws and jaw suspension
    —Internal fertilization and claspers
    —Distinctive soft tissue and physiological features
    6.3 Euchondrocephali and Chimaeriformes
    —Biology of extant Chimaeriformes
    6.4 Elasmobranchii, Euselachii, and Neoselachii
    —Selachii: Sharks
    —Batomorphi: Skates and rays
    6.5 Biology of Neoselachii
    —Bioluminescence and biofluorescence
    —-Hypoxia and the epaulette shark
    —Endothermal heterothermy
    —Elasmobranch brains
    —Social networks and migration in sand tiger sharks
    6.6 Declining Elasmobranch Populations
    —Conservation and sawfishes
    —Threats to chondrichthyans
    —Vulnerabilities of chondrichthyans
    —Ecological impacts of shark population declines
    —Policies to protect sharks
    Chapter 7
    Origin of Osteichthyes and Radiation of Actinopterygian
    7.1 Osteichthyes, Actinopterygii, and Sarcopterygi
    —Osteichthyan character
    —Fin adaptations
    —Other differences between actinopterygians and sarcopterygians
    7.2 Actinopterygii: Basal Group
    —Neopterygii: Holostei
    —Neopterygii: Teleoste
    7.3 Characters of Teleostei
    7.4 Teleostei: Basal Groups
    —Basal euteleosts
    7.5 Teleostei: Acanthopterygii
    —Basal acanthopterygians
    7.6 Swimming and Hydrodynamics
    —Generating forward thrust
    —Modes of locomotion
    —Speed and drag
    —Steering, stopping, and staying in place
    7.7 Reproduction and Development
    —Sex change in teleosts
    7.8 Ecology of Marine Teleosts
    —Black-water diving and larval teleosts
    —The photic zone and its subdivisions
    —Coral reef fishes
    —Pelagic and deep-sea fishes
    Chapter 8
    Sarcopterygians and the Origin of Tetrapods
    8.1 Phylogenetic Concepts of Tetrapoda and Characters for Sarcopterygii
    8.2 The Miguasha Lagerstätte and the “Good Fossil Effect”
    8.3 Actinistia
    8.4 Dipnomorpha
    8.5 Tetrapodomorpha
    —Basal tetrapodomorphs
    8.6 Moving onto Land
    —How did fins become limbs?
    —Body support and locomotion of early tetrapods
    8.7 Paleoecology of Devonian Tetrapodomorphs
    Chapter 9
    Origins of Lissamphibia and Amniota
    9.1 Paleozoic Tetrapods and the Origins of Extant Groups
    —Origins of Lissamphibia
    —Reptiliomorpha and the origin of amniotes
    —Paleozoic diversification of amniotes
    9.2 Characters of Amniotes
    —Skeletal characters
    —The amniotic egg
    —Other soft-tissue characters of amniotes
    9.3 Diversification of Amniotes
    —Temporal fenestration: Synapsids and diapsid
    —Ankle evolution in amniote
    Chapter 10
    Geography and Ecology of the Mesozoic
    10.1 Continental Geography and Climates
    —Continental movements
    —Climate shifts
    10.2 Terrestrial Ecosystems
    10.3 Marine Ecosystems
    —Faunal composition: Apex predators
    —Other clades
    10.4 Extinctions
    —Triassic and Jurassic extinctions
    —Cretaceous extinctions
    Chapter 11
    Living on Land
    11.1 Support on Land
    —Axial skeleton
    —Axial muscle
    —Appendicular skeleton
    —Size and scaling
    11.2 Locomotion
    11.3 Eating
    11.4 Breathing Air
    11.5 Pumping Blood Uphill
    11.6 Sensory Systems
    —Hearing and equilibrium
    11.7 Conserving Water in a Dry Environment
    —Cutaneous water loss
    —Respiratory water loss
    —Excretory water loss
    Chapter 12
    12.1 Diversity of Lissamphibians
    —Synapomorphies of Lissamphibia
    12.2 Life Histories of Lissamphibians
    —Mating and reproduction in salamanders
    —Anuran mating and reproduction
    —Anuran metamorphosis
    —The ecology of tadpoles
    —Caecilian reproduction and development
    12.3 Respiration and Circulation
    —Cutaneous respiration and blood flow
    —Blood flow in larvae and adults
    12.4 Water Relations
    —Uptake and storage of water
    —Cutaneous water loss
    —Behavioral control of cutaneous water loss
    12.5 Crypsis, Warning Colors, Toxins, and Venoms
    —Skin glands and toxins
    —Toxicity and diet
    —Venomous lissamphibians
    12.6 Why Are Lissamphibians Vanishing?
    —Chytrid fungi
    —Synergisms and domino effects
    Chapter 13
    Synapsids and Sauropsids: Two Ways of Living on the Land
    13.1 Conflicts between Locomotion and Respiration
    13.2 Lungs and Lung Ventilation: Supplying Oxygen to the Blood
    —Synapsid lungs
    —Sauropsid lungs
    13.3 Circulatory Systems: Supplying Oxygen to Tissues
    —Systemic arches of mammals and birds
    —Hearts with a ventricular septum: Mammals and birds
    —Hearts without a ventricular septum: Turtles and lepidosaurs
    —Shunting blood when the heart has a ventricular septum: Crocodylians
    13.4 Getting Rid of Wastes: The Kidneys
    —Nitrogenous waste products
    —Nitrogen excretion by synapsids: The mammalian kidney
    —Nitrogen excretion by sauropsids: Renal and extrarenal routes
    Chapter 14
    Ectothermy and Endothermy: Two Ways of Regulating Body Temperature
    14.1 Why Regulate Body Temperature?
    14.2 Ectothermal Thermoregulation
    —Energy exchange and mechanisms of ectothermy
    —Thermal ecology of ectotherms
    14.3 Endothermal Thermoregulation
    —Mechanisms of endothermal thermoregulation?
    14.4 Pure Ectothermy and Pure Endothermy Lie at the Extremes of a Continuum
    —Endothermal ectotherms
    —Heterothermal endotherms: Torpor and hibernation
    —Heterothermal endotherms: Hyperthermia and life in the desert
    14.5 Evolution of Endothermy
    —How did endothermy evolve?
    —Evaluating the models
    —Many factors
    14.6 Thermoregulation, Energy Use, and Body Size
    —Energy requirements
    —Body size
    —Gigantothermy and the body temperatures of dinosaurs
    14.7 Ectotherms, Endotherms, and Ecosystems
    Chapter 15
    15.1 Characters and Diversity of Lepidosaurs
    —Rhynchocephalians and the biology of tuatara
    —Squamata: Lizards
    —Squamata: Serpentes
    15.2 Foraging Modes
    15.3 Skull Kinesis and Feeding
    —Feeding specializations of snakes
    —Venom and fangs
    —Hearts and stomachs
    15.4 Predator Avoidance and Defense
    —Crypsis, aposematism, and mimicry
    —Venom and poisons as defense mechanisms
    15.5 Social Behavior
    —Courtship and territoriality
    —Sociality and parental care
    15.6 Reproductive Modes
    —Oviparity and viviparity
    —Sex determination
    15.7 Climate Change
    Chapter 16
    16.1 Form and Function
    —Shell and skeleton
    —Head retraction
    —Lung ventilation
    —Evolution of the turtle body plan
    16.2 Diversity
    16.3 Social Behavior, Communication, and Courtship
    16.4 Reproduction
    —Environmental sex determination
    —Parental care
    —Hatching and the behavior of baby turtles
    16.5 Navigation and Migration
    —Navigation by adult sea turtles
    —Navigation by hatchling and juvenile sea turtles
    16.6 Turtles in Trouble
    —Life history
    —Turtles are both delicious and considered medicinal
    —Turtles are in demand as pets
    —Sea turtles face extra risks
    Chapter 17
    17.1 Diversity of Extant Crocodylians
    —Distribution of extant crocodylians
    17.2 The Crocodylomorph Lineage
    17.3 Predatory Behavior and Diet
    17.4 Communication and Social Behavior
    17.5 Reproduction and Parental Care
    —Environmental sex determination
    —Parental care
    17.6 Threats to and from Crocodylians
    —Threats from crocodylians
    —Threats to crocodylians
    —Reconciling humans and crocodylians
    Chapter 18
    Avemetatarsalia and the Origin of Dinosauria
    18.1 Characters and Systematics of Avemetatarsalia
    18.2 †Pterosaurs: Vertebrates Achieve Powered Flight
    —Structure of †pterosaurs
    —Reproduction, eggs, and parental care
    —Did the evolution of birds doom †pterosaurs
    18.3 Dinosaurs: One of the Most Successful Tetrapod Radiations
    —The structure of dinosaurs
    18.4 †Ornithischia
    —Social behavior of †ornithischian dinosaurs
    —Nesting and parental care by †ornithischians
    —Social behavior of †ornithischian dinosaurs
    —Nesting and parental care by †ornithischians
    18.5 †Sauropodomorpha
    —Social behavior of †sauropod
    —Nesting and parental care by †sauropodomorphs?
    Chapter 19
    Theropods and the Origin of Birds
    19.1 Characters and Systematics of Theropods
    —Phylogenetic overview of Theropoda
    —†Coelophysoids: Early theropods
    —Community ecology of theropods
    —Social behavior of theropods
    19.2 †Archaeopteryx, Mesozoic Avialans, and the Mosaic Evolution of Avian Characters
    —Discovery of †Archaeopteryx
    —Cretaceous avialans
    —Mosaic evolution of some avialan characters
    —Other avian features
    —Body size
    19.3 Evolution of Powered Flight
    —How-and why-birds got off the ground
    —Gliding and flying by other Mesozoic paravians
    19.4 Reproduction and Parental Care by Theropods
    —Eggs and nests
    —Parental care of hatchlings
    Chapter 20
    Geography and Ecology of the Cenozoic
    20.1 Continental Geography and Climates
    —Continental movements
    —Cenozoic climates
    20.2 Cenozoic Ecosystems
    —Fossil Lake
    —Freshwater habitats
    —Marine habitats
    —Terrestrial flora
    —Terrestrial fauna
    20.3 The Great American Biotic Interchange
    —Terrestrial vertebrates of North and South America
    —Faunal interchange
    —Marine fauna and isthmian pairs
    20.4 Extinctions
    Chapter 21
    Extant Birds
    21.1 Diversity of Aves
    21.2 Structural Specializations for Flight and Bipedalism
    —Body size
    —Streamlining and weight reduction
    21.3 Wings and Flight
    —Flight mechanics
    —Wing shape and flight
    21.4 Feet and Locomotion
    —Hopping, walking, and running
    21.5 Bills, Feeding, and Digestion
    —Bills, cranial kinesis, and tongues
    —Digestive tract
    21.6 Sensory Systems
    21.7 Communication
    —Visual displays
    21.8 Reproduction
    —Reproductive organs and insemination
    —Egg structure
    —Maternal effects
    —-Sex determination
    -Hatching and developmental state of young
    21.9 Parental Care
    —Nest building
    —Feeding young
    —Interspecific brood parasitism
    21.10 Orientation, Navigation, and Migration
    —Navigational abilities
    —Using multiple cues during navigation
    —Seasonal migration
    21.11 Conservation
    Chapter 22
    Synapsids and the Origin of Mammals
    22.1 Synapsid Evolution
    —Cranial skeleton and teeth
    —Phylogenetic history of synapsids
    22.2 Jaw Joints and Middle Ear Bones
    22.3 Other Mammalian Features
    —Specializations of the palate and tongue for swallowing
    —Facial muscles
    —Lactation, nursing, and suckling
    —Brain and senses
    —Internal anatomy
    22.4 Basal Mammalian Clades
    Chapter 23
    23.1 Therian Features and Origins of Marsupialia and Placentalia
    —Therian skeletons
    23.2 Diversity of Marsupials
    —Marsupials and the Australian fauna
    23.3 Diversity of Placentals
    23.4 Reproduction
    —Urogenital tracts
    —Evolution of therian viviparity
    23.5 Teeth and Feeding Specializations
    —Cusps and lophs
    —Carnivores and herbivores: Differences in jaw muscles
    —Digestive tracts
    23.6 Locomotion
    —Limbs: Speed versus power
    —Cursorial adaptations of ungulate limbs
    —Powered flight of bats
    —Cetacean evolution
    23.7 Trophy Hunting and Extinction Risk
    —Bighorn sheep: A case study
    —Endangering the endangered: The effect of perceived rarity
    —The extinction vortex
    Chapter 24
    Primate Evolution and the Emergence of Humans
    24.1 Primate Origins and Diversification
    —Basal primates
    —New World monkeys
    —Old World monkeys and apes
    24.2 Origin and Evolution of Hominoidea
    24.3 Origin and Evolution of Hominini
    —Distinctive features of hominins
    —Early hominins
    24.4 The Genus Homo
    —†Homo habilis
    —†Homo erectus
    —†Dmanisi hominins
    —†Denisovan hominins
    —†Homo longi
    —Island species and miniaturization
    —†Homo naledi
    —†Homo bodoensis
    —Origin and radiation of Homo sapiens
    —What happened to the humans who were already there?
    24.5 Evolution of Human Characters
    —Large brains
    —Speech and language
    24.6 Humans and Other Vertebrates
    —Humans as superpredators and environmental disruptors
    —Megafaunal extinctions
    —Is this the Anthropocene?


"“I really like the summary component of Pough et al. This will be extremely helpful for students to focus on the big picture points moving forward.” -Carly Anne York, Lenoir-Rhyne University "

"“I think Vertebrate Life has over many iterations of deeply scholarly editions has achieved an excellent treatment of the comparative evolution of adaptive form and function across the tree of vertebrate life. It excels at placing the comparative morphology and physiology of major vertebrate lineages into a clear phylogenetic context with outstanding integration of both extinct and extant forms through the history of space and time on Earth since the early Paleozoic. [In the upcoming edition,] I like the continued expansion to more mixed media illustrations. The increased use of multiple colors in displays and use of more photographs to complement drawn scientific illustrations continues to move in a productive direction for this chapter, and this new edition in general.” -Andrew M. Shedlock, University of Charleston "

"“Vertebrate Life really has been the top undergraduate textbook for a survey course on the evolution and form and function of vertebrate animals, for the better part of two decades. The new edition [has] upped the ante a bit in terms of clarity and coverage.” -Jonathan Weinbaum, Southern Connecticut State University "

"“[The new edition of Vertebrate Life] updates a previous edition of an excellent text with better organization and new information. [It has a] balance between presenting interesting information without overloading detail.” -Udo Savalli, Arizona State University "

"“Pough et al. has a wonderful variety of species that the students will find compelling. It is written in a very accessible writing style, [with a variety of] engaging species examples. The figures will greatly enhance student understanding with well-chosen and interesting examples.” -Claire Kendal-Wright, Chaminade University "

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