A

I n t r o d u c t i o n

K e n n e t h
Second

Edition

t o

K a r d o n g

4.6 billion years-ago

Second

Edition

Kenneth V. Kardong
W a s h i n g t o n State U n i v e r s i t y

The McGraw-Hill Companies

G r aw
Hill
u

H i g h e r

E d u c a t i o n

AN INTRODUCTION TO BIOLOGICAL EVOLUTION SECOND EDITION

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Library of Congress Cataloging-in-Publication Data
Kardong, Kenneth V.
An introduction to biological evolution / Kenneth V. Kardong. 2nd ed.
p. cm.
Includes index.
ISBN 978-0-07-305077-5 ISBN 0-07-305077-6 (hard copy: alk. paper)
1. Evolution (Biology)Textbooks. I. Title.
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576.8dc22
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2006028519

Kenneth Kardong is professor in the School of

Biological Sciences at Washington State University. He
obtained his advanced degrees from the University of
Washington (Seattle) and the University of Illinois
(Champaign-Urbana). In addition to teaching histology,
embryology, and introductory biology, he has also piloted courses in vertebrate evolution and Darwinian
principles of evolution for biology teachers, science
majors, and students in business and the humanities.
Kardong is a recipient of various teaching awards, and
sets high goals for both himself and his students. First,
he expects that students develop a respect for discovered knowledge based on rational techniques of investigation, which include thinking critically, gathering
logical information, and developing reasoned arguments based on sound evidence. Second, because information changes and knowledge evolves, Kardong
encourages students to become life-long learners. This
means that they understand the resources that are available to them, apply critical judgments to cherished beliefs, and have the intellectual honesty to evolve along
with their disciplines. Finally, Kardong believes that
students must take responsibility for their own learning.
Because learning is an active processnot a passive
endeavor in which teachers give and students receive
students must be active and engaged participants in
their own learning. In order to achieve these goals,

Kardong, together with colleagues, developed "seminar

teams," interactive, collaborative-based learning groups
for undergraduate students. Not only was this strategy
effective in small courses, but Kardong was able to
adapt it to larger undergraduate courses as well.
For many years, Kardong has safely engaged undergraduate students in his research program, which is
broadly centered on the evolution of complex systems,
but more specifically focuses on the process involved in
the evolution of venomous snakes from nonvenomous
ancestors. In order to gain a first-hand understanding of
how information is gathered and upon which foundations scientific interpretations are based, Kardong's
students analyze venom pharmacology, examine the
biomechanics of the complex snake jaw apparatus, and
study the predatory behavior of nonvenomous and venomous snakes. Because this research is done at a high
level of expectation, Kardong's undergraduates occasionally have the opportunity to publish their work i f
the animals cooperate and the students themselves are
not defeated by the unexpected surprises that attend
authentic research. Besides holding productive international scientific collaborations with colleagues in the
Netherlands, France, Belgium, Canada, and Japan, Kardong has also authored or co-authored over 90 scientific publications, including the textbook, Vertebrates:
Anatomy, Function, Evolution.

To Jason and Tami,

to Kyle and Darcie,
and to their good friends and company

Chapter 1

Evolution of Evolution

Chapter 2

Time

Chapter 3

Heredity

Chapter 4

E m e r g e n c e o f Life

Chapter 5

Diversity o f Life

Chapter 6

Evidence of Evolution

Chapter 7

Selection

Chapter 8

Variation: Spice o f Life

Chapter 9

Speciation

18
40

76

Human

Chapter 15 H u m a n

144

176

Life H i s t o r y Strategies

Chapter 13 Extinctions

98

158

C h a p t e r 12 Life in G r o u p s

Chapter 14

60

122

Chapter 10 Co-Evolution
Chapter 11

196

206
226

E v o l u t i o n : T h e Early Years

248

Evolution: Building Modern

Humans

Chapter 1 6 Evolutionary Biology: Today and Beyond

Chapter 17 Afterword

318

296

270

Preface xi
Chapter 1

Evolution of Evolution 2
Introduction 3
Philosophical Schools 3
Fact, Course, Mechanism 5
Fixity of Species 6
Linnaeus 6
Naturalists 6
Change of Species 7
J-B. de Lamarck 7
Upward to Perfection 8

Chapter 2

Time

18
Introduction 19
Dating Fossils 20
Stratigraphy 21
Index Fossils 22
Radiometric Dating 23
Geological Ages 25

Chapter 3

Heredity

"The Age of Bacteria" 28

Fossils and Fossilization 28
Recovery and Reconstruction 30
Lewis and Clark-In Search of Mastodons 32
From Animal to Fossil 36
Missing Fossils 38
Overview 38

40

Introduction 41
Inheritance by Intuition 41
Early Intuition 41
Blending Inheritance 42
Mendelian Inheritance 43
Gregor Mendel 43
Testcross 47
Mendelian Principles of Inheritance 48
Mendel's Achievement 48
Chapter 4

The Mechanism of Evolution:

Natural Selection 9
A. R. Wallace 9
Charles Darwin 10
A Most Dangerous Voyage 12
Critics and Controversy 13
Captain and Commander 13
Overview 14

Chromosomes 50
Cell Division 50
Mendel Amended 52
Gene Linkage 52
Multiple Alleles 52
Multiple GenesPolygenes 53
Population Genetics 53
Overview 55

Emergence of Life 60
Introduction 61
Major Transitions of Life 61
Inorganic to Organic Evolution
(4 billion years ago) 62
Cell - Prokaryotic. Heterotroph
(3.5 billion years ago) 64
Cell - Prokaryotic, Autotroph
(2.7 billion years ago) 64
Cell - Prokaryote to Eukaryote
(2 billion years ago) 65
Multicellularity 66
Brushing Up on Dental Caries 67
Major Transitions of Life and Consequences 67
Ozone 67
Pollutant 67

Eukaryotic Origins 67
Chemical CodingFrom Genotype to
Phenotype 69
DNA 69
RNA 69
Genes 70
Cell Metabolism 71
Metabolic Pathways 72
Carbon Fixation 72
Photosynthesis 73
What Is Life? 73
An RNA World 73
A Protein World 73
Overview 74

CONTENTS

Chapter 5

Diversity o f Life 7 6
Introduction 77
Prokaryotes 77
Bacteria (Eubacteria) 77
Archaea (Archaebacteria) 79
E. coliFriend and Foe 79
Eukaryotes 79
Protists 79

Chapter 6

Directional Selection 133

Disruptive Selection 134
Sexual Selection 134
From Mate to Meal 138
Overview 140

Variation: Spice o f Life 1 4 4

Introduction 145
Tulips 145
Mixing It Up 146
Recombination 146
Sex 146
Mutations 147
Early Work 147

Chapter 9

Vestigial and Atavistic Structures 111

Distributional Evidence 113
The Course of Evolution 119
Overview 119

Selection 122
Introduction 123
Artificial versus Natural Selection 124
Artificial Selection 124
Natural Selection 128
The Phenotype Takes a Beating 128
Types of Natural Selection 132
Stabilizing Selection 132

Chapter 8

Plants 80
Fungi 85
Animals 85
Fungus among Us 86
Environment 92
Overview 95

Evidence of Evolution 98
Introduction 99
The Fact of Evolution 100
The Fossil Record 100
Comparative Anatomy 102
Comparative Embryology 106
Human AppendixOut of a Job 111

Chapter 7

vii

Speciation

Mistakes Happen 147

Point Mutations 148
Gene Duplication 149
Chromosomal Mutations 149
Sickle-Cell Anemia: Disease
against Disease 150
Hox Genes and Their Kingdoms
Overview 155

158

Introduction 159
Species Definitions 160
Biological Species 160
Morphospecies 160
Paleospecies 160
Agamospecies 160
The Process of Species Formation 161
Four Steps to Speciation 161
Isolation and Diversification 163
Accentuated Reproductive and Ecological
Isolation 163

Reproductive Isolating Mechanisms 164

Prezygotic Mechanisms 164
Postzygotic Mechanisms 166
Natural Selection and RIMs 166
Patterns of Speciation 167
Clines 167
Ring Species 169
Flaming Retreats 169
Parallelism and Convergence 172
Latitudinal Gradients of Species Diversity 172
Overview 173

viii

CONTENTS

Chapter 10 Co-Evolution

176

Introduction 177
Competition 177
SymbiosisGood, Bad, and Ugly 178
Plant-Animal Co-Evolution 179
Spines and Thorns 179
Arms Race 180
Chemical Warfare 180
Mutualism 181
Commensalism 184
Protective Coloration and Shape 185
Camouflage 185
Warning Coloration (Aposematic) 186

C h a p t e r 11

Life H i s t o r y Strategies

196

Introduction 197
Life History Traits 197
Lizards 197
Guppies 198
Roundabout with Parasites 199

C h a p t e r 12

Life in C r o u p s

Extinctions

Time and Energy Budgets 201

Abiotic Factors 201
Biotic Factors 202
Overview 204

206

Introduction 207
Alarm Calls 207
Individual Selection and Group Selection 209
Altruism versus Selfish Behavior 210
Kin Selection 210
Inclusive Fitness 210
SexWhat Good Is It? 211
Coefficient of Relationship 212
Levels of Selection 213

C h a p t e r 13

Mimicry 187
Batesian Mimicry 187
Milkweeds and Monarchs on the Move 191
Mullerian Mimicry 192
Other Types of Mimicry 192
Overview 193
Linkage and Liaisons 193
Remodeling 194
Complex 195

Microevolution and Macroevolution 214

Quantum Evolution 214
Punctuated Equilibrium 216
Consequences of Punctuated Equilibrium 217
Rapid Evolution 218
On the Edge 219
Macro Changes at Micro Levels 221
Overview 223

226

Introduction 227
Uniform Extinctions 228
Co-Evolution 228
Islands 229
Red Queen 232
Assessment of Uniform Extinctions 233
Mass Extinctions 234
Causes of Dinosaur Extinctions 235

Mass ExtinctionsCase Studies 236

The North Pole Is Headed South 238
Plate Tectonics 238
Ice Ages 242
Cosmic Collisions 243
Overview 245