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 Contents
Preface xv Paleontology 29
Fossilization and Fossils 30
Recovery and Restoration 32
CHAPTERo ne From Animal to Fossil 35
Introduction 1 Dating Fossils 37
Stratigraphy 37
Comparative Vertebrate Morphology 1 Index Fossils 38
Designs of Students 2 Radiometric Dating 39
Vertebrate Design—Form and Function 3 Geological Ages 39
Grand Design 4 Tools of the Trade 41
Historical Predecessors—Evolution 4 The Question 41
The Process behind the Change 5 The Function 42
Linnaeus 5 The Biological Role 46
Naturalists 6 Overview 47
J-B. de Lamarck 6
Acquired Characteristics 7
Upward to Perfection 7 CHAPTERtwo
Natural Selection 8
A. R. Wallace 8 Origin of Chordates 48
Charles Darwin 9
Critics and Controversy 10 Chordate Phylogeny 48
Historical Predecessors—Morphology 11 Chordate Characteristics 50
Georges Cuvier 11 Notochord 51
Richard Owen 12 Pharyngeal Slits 52
Endostyle or Thyroid Gland 53
Why Are There No Flying Elephants? 14
Dorsal and Tubular Nerve Cord 54
Morphological Concepts 15 Postanal Tail 54
Similarities 15 Chordate Body Plan 54
Symmetry 16 Protochordates 54
Segmentation 16 General Protochordate Features 55
Evolutionary Morphology 19 Hemichordata 56
Function and Biological Role 19 Enteropneusta—“Acorn Worms” 57
Preadaptation 20 Pterobranchia 60
Evolution as Remodeling 21 Hemichordate Phylogenetic Affinities
to Chordates 60
Phylogeny 21 Hemichordate Phylogenetic Affinities
Of Bean Stalks and Bushes 21 to Echinoderms 61
Simplification 23 Cephalochordata 61
Patterns of Phylogeny 23 Urochordata 66
Grades and Clades 24 Ascidiacea—“Sea Squirts” 68

vii
 Larvacea (Appendicularia) 70 Salientia (Anura)–Frogs 107
Thaliacea 73 Gymnophiona (Apoda)–Caecilians 108
Overview of Protochordates 73 Lepospondyls 108
Chordate Origins 74 Amniotes 108
Chordates from Annelids and Arthropods 75 Stem-Amniotes 110
Chordates from Echinoderms 76 Sauropsids 110
Auricularian Hypothesis 76 Mesosaurs 110
Larval Echinoderm to Chordate Tadpole 77 Reptilia 111
Chordate Origins and Phylogeny 78 Synapsida 119
Dorsoventral Inversion 78 Pelycosauria 119
Therapsida 120
Overview 80
Mammalia 121
Overview 126
CHAPTERthree
The Vertebrate Story 82 CHAPTERfo u r
Introduction 82 Biological Design 128
Innovations 83 Introduction: Size and Shape 128
Vertebral Column 83
Head 84 Size 129
Origin of Vertebrates 84 Relationships Among Length, Area, and Volume 129
Step 1: Prevertebrate 85 Surface Area 133
Step 2: Agnathan 85 Volume and Mass 133
Step 3: Gnathostome 86
Vertebrate Classification 86
Shape 134
Allometry 134
Agnathans 86 Transformation Grids 134
Living Agnathans 86
On the Consequences of Being
Myxinoidea 86
Petromyzontiformes 88 the Right Size 137
Early Vertebrate Fossils 89 Biomechanics 137
Conodonts 89 Fundamental Principles 138
Ostracoderms 90 Basic Quantities—Length, Time, and Mass 138
Pteraspidomorphi 93 Units 139
Other Ostracoderms (Osteostracans, Derived Quantities—Velocity, Acceleration,
Anaspids, Thelodonts) 93 Force, and Relatives 139
Overview of Agnathan Evolution 93 Reference Systems 140
Gnathostomes 94 Center of Mass 140
Placodermi 94 Vectors 140
Chondrichthyes 95 Basic Force Laws 141
Elasmobranchii—Sharks and Rays 96 Free Bodies and Forces 142
Holocephali—Chimaeras 97 Torques and Levers 142
Land and Fluid 144
Teleostomi 97 Life on Land: Gravity 144
Acanthodii 97 Life in Fluids 145
Osteichthyes 98 Machines 148
Actinopterygii 99 Strength of Materials 148
Sarcopterygii 101 Loads 149
Overview of Fish Phylogeny 104 Biological Design and Biological Failure 149
Tissue Response to Mechanical Stress 151
Tetrapods 104
Responsiveness of Bone 152
Primitive Tetrapods 104
Labyrinthodonts 104 Biophysics and Other Physical Processes 155
Lissamphibia—Modern Amphibians 106 Diffusion and Exchange 156
Urodela (Caudata)–Salamanders 107 Pressures and Partial Pressures 156

viii Contents
 Countercurrent, Concurrent, Maturation 197
and Crosscurrent Exchange 156 Metamorphosis 197
Optics 158 Heterochrony 198
Depth Perception 158 Peramorphosis 199
Accommodation 159 Paedomorphosis 199
Overview 159 Ontogeny and Phylogeny 201
Biogenetic Law 201
Von Baer’s Law 202
CHAPTERfive Overview of the Biogenetic Laws 202
Hox Genes and Their Kingdoms 204
Life History 161 Egg to Adult 204
Shaping Up: Positions and Parts 204
Introduction 161 Evolutionary Significance 204
Early Embryology 163 Epigenomics 205
Induction 206
Fertilization 163
Phylogeny 206
Cleavage 164
Amphioxus 164 Overview 210
Fishes 165
Amphibians 165
Reptiles and Birds 165 CHAPTERsi x
Mammals 166
Overview of Cleavage 167 Integument 212
Gastrulation and Neurulation 167
Amphioxus 169 Embryonic Origin 213
Fishes 169 General Features of the Integument 213
Amphibians 171
Dermis 213
Reptiles and Birds 171
Epidermis 215
Mammals 173
Phylogeny 216
Organogenesis 176
Integument of Fishes 216
Histogenesis 177
Primitive Fishes 217
Epithelium 177
Chondrichthyes 217
Covering and Lining Epithelium 179
Bony Fishes 218
Glandular Epithelium 180
Integument of Tetrapods 219
Connective Tissues 180
Amphibians 219
General Connective Tissues 181
Reptiles 220
Special Connective Tissues 181
Birds 221
Bone Development and Growth 182
Mammals 228
Endochondral Bone Development 183
Intramembranous Bone Development 184 Specializations of the Integument 233
Comparative Bone Histology 186 Nails, Claws, Hooves 233
Bone Remodeling and Repair 186 Horns and Antlers 234
Joints 187 Baleen 235
Neural Crest and Ectodermal Placodes 189 Scales 236
Extraembryonic Membranes 190 Dermal Armor 236
Mucus 237
Reptiles and Birds 190
Color 238
Mammals 191
Eutherian Placenta 192 Overview 239
Other Placentae 192
Overview of Early Embryonic
CHAPTERseven
Development 193
Skeletal System: The Skull 241
Development of the Coelom
and Its Compartments 196 Introduction 242

Contents ix
Chondrocranium 242 Sternum 299
Embryology 242 Gastralia 299

Splanchnocranium 244 Embryonic Development 301

Embryology 244 Fishes 301
Origin of Jaws 246 Tetrapods 302
Types of Jaw Attachments 247 Phylogeny 304
Dermatocranium 248 Fishes 304
Parts of the Dermatocranium 249 Agnathans 304
Dermal Bone Series 249 Gnathostomes 304
Tetrapods 309
Overview of Skull Morphology 250 Early Tetrapods 309
Braincase 250 Amniotes 312
Jaws 253
Hyoid Apparatus 253 Form and Function 314
Fluid Environment 314
Cranial Kinesis 253 Terrestrial Environment 315
Phylogeny of the Skull 255 Design of Vertebrae 317
Direction of the Neural Spine 317
Agnathans 255
Height of the Neural Spine 317
Early Vertebrates 255
Regionalization of the Vertebral Column 318
Ostracoderms 255
Cyclostomes 255 Overview 321
Gnathostomes 255
Fishes 256
Early Tetrapods 263 CHAPTERni ne
Primitive Amniotes 264
Modern Reptiles 268 Skeletal System: The
Birds 272 Appendicular Skeleton 325
Synapsids 274
Overview of Skull Function and Design 284 Introduction 325
Prey Capture 285 Basic Components 326
Feeding in Water 285 Fins 326
Feeding in Air 287 Limbs 326
Swallowing 288
Origin of Paired Fins 327
Overview 288
Gill-Arch Theory 327
Cranial Neural Crest 288 Fin-Fold Theory 328
Emergence of Mammals 289 Embryonic Development of Tetrapod Limbs 330
Evolutionary Modifications of Immature Forms:
Akinesis in Mammals 291 Phylogeny 331
Composite Skull 292 Fishes 331
Agnathans 331
Placoderms 331
CHAPTERei g h t Chondrichthyans 333
Acanthodians 334
Skeletal System: The Axial Bony Fishes 334
Skeleton 294 Tetrapods 336
Pectoral Girdle 336
Introduction 294 Pelvic Girdle 339
Manus and Pes 339
Basic Components 295
Vertebrae 295 Evolution of the Appendicular System 346
Regions of the Vertebral Column 295 Dual Origin of the Pectoral Girdle 346
Centra 295 Adaptive Advantage of Lobe Fins 347
Ribs 297 Onto the Land 347

x Contents
Form and Function 348 Axial Musculature 394
Swimming 349 Appendicular Musculature 397
Terrestrial Locomotion 350 Cranial Musculature 405
Early Gaits 350 Branchiomeric Musculature 405
Early Modes of Locomotion 350 Hypobranchial Musculature 408
Cursorial Locomotion 353 Overview 411
Aerial Locomotion 358
Origin of Bird Flight 367
Fossorial Locomotion 369 CHAPTERel even
Overview 370
The Respiratory System 413
t en
CHAPTER Introduction 413

The Muscular System 372 Respiratory Organs 416

Gills 416
Introduction 372 Gas Bladders 416
Lungs 416
Organization of Muscles 373
Swim Bladders 416
Classification of Muscles 373 Cutaneous Respiratory Organs 417
Skeletal Muscle 374 Accessory Air-breathing Organs 417
Cardiac Muscle 375 Breathing and Embryos 418
Smooth Muscle 375
Structure of Skeletal Muscles 375 Ventilatory Mechanisms 421
Tendons 376 Cilia 421
Basis of Muscle Contraction 376 Muscular Mechanisms 421
Resting and Active Muscle 376 Water Ventilation: Dual Pump 421
Molecular Mechanisms of Contraction 376 Air Ventilation: Buccal Pump 422
Air Ventilation: Aspiration Pump 423
Muscle Function 377
Muscle Fibers 377 Phylogeny 424
Tension-Length Curves for a Single Muscle Fiber 377 Agnathans 424
Properties of Muscle Fibers 377 Elasmobranchs 426
Muscle Organs and Fibers 379 Bony Fishes 427
Whole Muscle Force Generation 379 Overview of Fish Respiration 427
Tension-Length Curves for a Whole Muscle 380 Gills 427
Graded Force 380 Lungs and Swim Bladders 428
Cross-Sectional Area 383 Amphibians 430
Fiber Orientation 383 Amphibian Larvae 430
Velocity of Shortening 385 Amphibian Adults 432
Distance of Shortening 385 Reptiles 433
Bone–Muscle Lever Systems 385 Mammals 435
Sequencing of Muscle Actions 387 Ventilation 435
Overview of Muscle Mechanics 388 Gas Exchange 437
Muscle Actions 388 Birds 437
Muscle Homologies 390
Form and Function 440
Embryonic Origin of Muscles 391 Patterns of Gas Transfer 440
Postcranial Musculature 392 Rates of Gas Transfer 443
Appendicular Musculature 392 Breathing in Water 444
Axial Musculature 393 Breathing in Air 444
Cranial Musculature 393
Evolution of Respiratory Organs 444
Jaw and Pharyngeal Musculature 393
Extrinsic Eye Muscles 393 Acid-Base Regulation 444
Ventilation 446
Comparative Anatomy 394 Ciliary Pumps 446
Postcranial Musculature 394 Muscular Pumps 446

Contents xi
 Water-to-Land Transition 446 Components of the Digestive System 504
Air-breathing Organs 446 Buccal Cavity 504
Advantages of Movement to Land 447 Boundaries 504
Air-breathing Mechanisms 448 Palate 505
Bird Lungs and Air Sacs 449 Teeth 506
Overview 450 Tongue 516
Pharynx 517
Alimentary Canal 520
CHAPTERtwelve Esophagus 521
Stomach 521
The Circulatory System 451 Intestines 523
Introduction 451 Cloaca 525
Specializations of the Alimentary Canal 525
Cardiovascular System 452 Vascularization of the Gastrointestinal Tract 527
Blood 452 Fishes 527
Arteries, Veins, and Capillaries 452 Tetrapods 528
Arteries 453 Associated Glands of Digestion 531
Hemodynamics of Circulation 453 Oral Glands 531
Veins 454 Liver 533
Microcirculation 454 Pancreas 533
Single and Double Circulation 455 Function and Evolution
Embryonic Development of the Cardiovascular
of the Digestive System 535
System 456
Phylogeny of the Cardiovascular System 457 Absorption 535
Arterial Vessels 461 Feces 535
Venous Vessels 466 Mechanical Breakdown of Food 536
Hearts 473 Mastication 536
Basic Vertebrate Heart 473 Gizzards 536
Fishes 475 Chemical Breakdown of Food 536
Amphibians 478 Foregut Fermentation 537
Reptiles 480 Hindgut Fermentation 540
Birds and Mammals 489 Foregut versus Hindgut Fermenters 540
Cardiovascular System: Matching Design Size and Fermentation 541
to Environmental Demands 490 Digesting Toxins 542
Accessory Air-breathing Organs 491 Feeding and Fasting 542
Diving Birds and Mammals 491 Overview 543
Heart Flow 492
Ontogeny of Cardiovascular Function 492
Fetal Circulation in Placental Mammals 492 fo u rteen
CHAPTER
Changes at Birth 492
Heat Transfer 494 The Urogenital System 545
Lymphatic System 496 Introduction 545
Lymphatic Vessels 496
Lymphatic Tissue 498
Urinary System 545
Form and Function 498 Embryonic Development 546
Nephrotome to Nephric Tubules 546
Overview 499 Tripartite Concept of Kidney Organization 547
Kidney Phylogeny 550
CHAPTERthirteen Fishes 550
Tetrapods 551
The Digestive System 503 Kidney Function and Structure 553
Excretion: Removing the Products
Introduction 503 of Nitrogen Metabolism 553
Preview 503 Osmoregulation: Regulating Water and Salt Balance 554

xii Contents
 Evolution 562 Endocrine Coordination 613
Preadaptation 562 Mammalian Reproduction 613
Origin of Vertebrates 562 Male 613
Reproductive System 563 Female 613
Metamorphosis in Frogs 616
Structure of the Mammalian Reproductive System 563
Fundamentals of Hormonal Control 620
Embryonic Development 563
Functional and Structural Linkage 620
Gonads and Gametes 563
Target Tissue Responses 621
Reproductive Tracts 565
The Endocrine System and the Environment 621
Overview 565
Female Reproductive System 567 Evolution 622
Ovary 567
Genital Ducts 567 Overview 624
Oviduct 569
Uterus 570
Male Reproductive System 572
Testis 572 si xteen
CHAPTER
Genital Ducts 572 The Nervous System 625
Copulatory Organs 576
Cloaca 582 Introduction 625
Urinary Bladder 586 Types of Cells within the Nervous System 625
Function and Evolution 588 Neuroglia 625
Potency and Fertility 588 Neurons 625
External and Internal Fertilization 588 Transmission of Information 626
Delays in Gestation 589 Neurosecretory Cells 628
Overview 589 Peripheral Nervous System 628
Spinal Nerves 629
Cranial Nerves 630
f i f t een
CHAPTER
Evolution 637
The Endocrine System 592 Functions of the Peripheral Nervous System 638
Spinal Reflexes 638
Survey of Endocrine Glands 592 The Autonomic Nervous System 641
Thyroid Gland 592
Central Nervous System 646
Structure and Phylogeny 592
Function 594 Embryology 646
Ultimobranchial Body and Parathyroid Gland 596 Spinal Cord 649
Ultimobranchial Body 596 Spinal Reflexes 649
Parathyroid Gland 597 Spinal Tracts 650
Form and Function 598 Brain 651
Adrenal Gland 598 Phylogeny 651
Structure and Phylogeny 598 Form and Function 654
Function 601 Functional Associations of Parts
Pancreatic Islets 602 of the Central Nervous System 666
Structure and Phylogeny 602 Limbic System 666
Function 602 Overview 669
Pituitary Gland 604
Structure 604
Phylogeny 606
Function 607 CHAPTERseventeen
Gonads 610
Pineal Gland 610
Sensory Organs 671
Secondary Endocrine Glands 611 Introduction 671
Gastrointestinal Tract 611
Kidneys 612 Components of a Sensory Organ 672

Contents xiii
General Sensory Organs 672 Evolutionary Analysis 722
Free Sensory Receptors 672 Historical Constraints 722
Encapsulated Sensory Receptors 673 Primitive and Advanced 722
Associated Sensory Receptors 673 Diversity of Type/Unity of Pattern 723
Proprioception 673 Mosaic Evolution 725
Mechanisms of Perceiving Stimuli Morphology and Modules 725
from General Sensory Receptors 674
Mode and Tempo of Evolution 727
Special Sensory Organs 674 Remodeling 728
Chemoreceptors 674 Embryonic Changes 729
Nasal Passages 675 Hox Genes 729
Vomeronasal Area 677 Evolutionary Significance 730
Mouth 678
The Promise of Vertebrate Morphology 730
Radiation Receptors 679
Photoreceptors 681
Infrared Receptors 691
Mechanoreceptors 693
Lateral Line System 694
Vestibular Apparatus 695 APPENDIX A
Auditory System 696
Functions of the Ear 701 Vector Algebra 
Electroreceptors 709
Structure and Phylogeny 709 APPENDIX B
Form and Function 709
Additional Special Sensory Organs 712 International Units (SI) 
Overview 712
APPENDIX C
ei g h t e e n
CHAPTER Common Greek and Latin Combining
Conclusions 714 Forms 

Introduction 714
APPENDIX D
Structural Analysis 717
Classification of Chordates Linnaean 
Functional Analysis 718
How Does It Work? 718 Classification of Chordates Cladistic 
Functional Coupling, Functional Compromise 719
Multiple Functions 720 Glossary 
Performance 721 Credits 
Ecological Analysis 722 Index 

xiv Contents
 Preface

If you are a student coming to the study of vertebrates for within a phylogenetic context. But, today we expect our stu-
the first time, several introductory remarks may be helpful, dents to develop academic and professional skills beyond just
especially on how this textbook will support your work. First, facility with anatomical terminology. In general, we expect
the discipline of vertebrate biology is diverse and inclusive. It our students to develop skills in critical thinking and a facil-
brings together themes from molecular biology, genes and ity with scientific concepts. Each of us will find our own
genomes, evolution and embryology, biomechanics and way of composing a course in vertebrate morphology that
experimental physiology, and it incorporates continuing and serves such course objectives. This textbook was written to
astonishing new fossils into the vertebrate story. Much of support such course objectives as individual instructors build
what you have met in earlier courses you will meet again here their courses. It is flexible. You can mix and match, change
in an integrated way. order to suit your course, and give emphasis to those sys-
Second, to unify these themes, I have again written and tems that most suit the organization of your course. Because
revised this seventh edition within the unifying framework of each chapter integrates form, function, and evolution perti-
form, function, and evolution. The first few chapters set this nent to that system, each chapter is coherent within itself.
up, and the subsequent chapters treat vertebrates system by Where information or concepts are treated in greater detail
system. You may notice that each of these subsequent chap- outside a particular chapter, they are cross referenced to help
ters begins with a discussion of morphology, followed by a guide the student and clarify the discussion. Although dis-
discussion of function and evolution. Each chapter is therefore cussed in earlier editions, let me repeat the specific strategy
self-contained—form, function, evolution. built into this textbook to improve student success and to
Third, as a student you likely enter this course after help them develop skills in critical thinking and conceptual
some background in the sciences, perhaps expecting to equip understanding.
yourself with practical knowledge useful later in professional
schools or in health-related careers. Certainly this course, in For the Student
part, delivers such practical information. But because ver-
tebrate morphology is an integrative discipline, it brings A number of strategic features within the textbook enhance
together physiology, embryology, behavior, and ecology its usefulness for students. It is richly illustrated with fig-
and also deploys modern methods of systematics and new ures that include new information and provide fresh per-
finds in paleontology. Consequently, you will move beyond spectives. Each chapter opens with an outline. Important
memorizing facts in isolation or as an end in themselves, concepts and major anatomical terms are boldfaced. Cross
and instead begin to meet and understand larger concepts references direct students to other areas of the text where
to which the morphology testifies. What may come as a sur- they can refresh their understanding or clarify an unfamil-
prise is that many theories, especially evolutionary theories iar subject. Each chapter concludes with a chapter overview,
within vertebrate biology, are still unsettled and unresolved, which draws attention to some of the concepts developed
inviting a new idea or fresh approach open to anyone. This is within the chapter. Box Essays are included along the way
one of the reasons I have included various controversies, and in most chapters. Their purpose is to present subjects or his-
support your efforts to become engaged in the thinking and torical events that students should find interesting and, per-
scientific process. haps from time to time, even fun. A glossary of definitions is
For faculty who have used this textbook before, you included at the end of the book.
will find it retains a familiar and inviting organization with In addition to its practical features, the textbook also
the science updated and the student support enhanced. For uses selected topics within vertebrate structure to develop
those coming to this textbook for the first time, you will student skills in critical thinking and mastery of concepts
notice that the morphology receives generous treatment within a coherent framework.

xv
Critical Thinking integrated organism, a connected system of parts and their
functions. This too was recognized within the nineteenth
Within the sciences, critical thinking is the ability to mar- century, suggesting constraints on evolutionary modifica-
shal factual information into a logical, reasoned argument. tion. Vertebrate morphology provides compelling examples
Especially if accompanied by a laboratory, a course in vertebrate of how an integrated organism might evolve. For example, a
morphology delivers hands-on experience with the anatomy of remarkable fossil record documents an undeniable change in
representative animals. Students can be directly engaged in the jaw articulation within synapsids, seeing the two participat-
discovery of vertebrate form. But they can be encouraged to go ing bones (articular, quadrate) of basal synapsids replaced by
beyond this. Instructors can lead students into larger issues— two different bones in derived groups, including mammals.
How does it function? How did it evolve? For example, early Fossil intermediates between the two conditions mark the
on in the textbook, students are introduced to “Tools of the anatomical changes, but they also suggest how functional
Trade,” methods by which we empirically examine how parts changes, which must accompany evolving systems, also
work and how we can place organisms within a phylogenetic change without disrupting performance.
context. After a discussion of basic morphology, each chapter Within many vertebrate systems, the close coupling of
discusses how these systems work and how they evolved. form and function with lifestyle is illustrated. Built on a basic
I have deliberately included new, neglected, or com- vertebrate plan, the tetrapod locomotor system illustrates the
peting views on function and evolution. Many of these ideas close relationship between limbs and axial skeleton, and the type
come from Europe, where they have been known for a long of locomotion—flight, cursorial, burrowing. The cardiovascular
time. Personally, I find many of these ideas compelling, even system, especially in organisms that exploit water and air, illus-
elegant. Others strike me, frankly, as thin and unconvinc- trates the close relationship between vascular morphology and
ing. Despite my own skepticism, a few contrary ideas are the physiological flexibility that permits. The basic concepts of
included. My purpose is to get students to think about issues form, function, and adaptive evolution parade before us as we
of form, function, and evolution. move from system to system in vertebrate morphology.
Several theories on the evolution of jaws are discussed, Evolution proceeds most often by remodeling, modi-
as are several theories of the origin of paired fins. Often stu- fication of a basic underlying plan, not by all new construc-
dents expect that today we have the final answers. Students tion. This is illustrated in the skeletal system, as well as
implore, “Just tell me the answer.” The debate about dino- within the cardiovascular (aortic arches) system.
saur physiology is a wonderful opportunity to show students
the ongoing process of scientific investigation. Most have
seen the Hollywood films and expect the issue settled. But Organizational Strategy
we know that science is an ongoing process of refinement, and Rationale
challenge, and sometimes revolutionary change. One Box
Essay sets forth the early case for dinosaur endothermy. That I have written this book within the unifying framework of
debate spawned further investigation that now returns to form, function, and evolution. These are common themes
challenge such a view of dinosaurs as “hot-blooded” beasts. that run throughout. The vertebrate groups are organized
The second Box Essay on dinosaur endothermy presents phylogenetically, and their systems discussed within such a
this newer and contrary evidence, and thereby showcases context. Morphology is foremost, but I have developed and
how, even in extinct animals, it is possible to test hypotheses integrated an understanding of function and evolution into
about their physiology, morphology, and lifestyles. the discussion of anatomy of the various systems. The first
five chapters prepare the way.
Chapter 1 introduces the discipline, evaluates the intel-
Concepts lectual predecessors to modern morphology, defines central
Vertebrate morphology also helps develop an appreciation concepts, and alerts students to misunderstandings they may
and understanding of the scientific concepts that unite biol- unknowingly bring with them to the study of evolutionary
ogy and reflect on “how” science works. As John A. Moore processes. Chordates and their origins are covered in chapter
put it, science is a “way of knowing” (Moore, American 2. Considerable attention is given to the neglected protochor-
Zoologist, 1988). Comparative morphology throws into clear dates and their evolution. This sets the stage for an extended
relief differences and similarities between organisms. The discussion of the cast of characters in the vertebrate radiation,
concepts of homology, analogy, and homoplasy help us which occupies us for the remainder of the book, beginning
understand the basis of these comparative features. Many of next in chapter 3. Here we discuss vertebrates, their origins,
the concepts were birthed in the nineteenth century and have and basic taxonomic relationships. Chapter 4 introduces
grown into the guiding themes of biology today. Evolution, basic concepts of biomechanics and biophysics, preparing for
defined as descent with modification through time, is one their use later in understanding aspects of vertebrate design
of the foundation concepts in biology. Vertebrate morphol- and function. Chapter 5 includes a summary of descriptive
ogy provides a showcase of adaptive change on the basic embryology and concludes with a discussion of the role
vertebrate body plan. But evolution is change in a highly embryonic processes play in vertebrate evolutionary events.

xvi Preface
 The remaining chapters develop each major system. Turning over Chordates. New developmental genet-
Besides carrying overall themes, each chapter internally ics, discussed in the previous editions, informs us that the
follows a consistent organization. Each begins with a basic immediate chordate ancestors flipped over, reversing dor-
introduction to the morphology, and then proceeds to dis- sal and ventral surfaces. That view seems to hold still and
cuss function and evolution. This way, the overall themes therefore remains the surprising basis of the chordate body
are repeated in each chapter, bringing consistency of presen- plan today.
tation to each chapter and coherence throughout. Updated and Revised. Countless changes and revisions
throughout this new edition have been made, some major,
some small. These changes have corrected misinformation,
New and Expanded updated information, and often better clarified an explana-
in the Seventh Edition tion. For this I am indebted to students, reviewers, and col-
leagues for bringing these suggestions to my attention.
New fossil finds, modern experimental research, and new
Serving the Student. Features of the textbook have been
phylogenies continue to enrich vertebrate biology, some-
further expanded to make its presentation more clear and
times solving old questions or surprising us with a new
inviting. The use of color brightens these sections of the
understanding of how vertebrates function and how they
book. Color has also been used to better correlate and com-
evolved. Much of this is added to this new edition.
pare structures between figures in these chapters. Where fea-
Feathered Dinosaurs. New and remarkable fossil finds
sible, within color signatures, for example, I have added more
of dinosaurs, especially in China, continue to be discov-
color to the illustrations. Many illustrations are new, revised,
ered, some showing evidence of surface feathers. In other
or relabeled to improve clarity. For example, besides those
words, feathers evolved before birds. This means that these
illustrations mentioned earlier, new/revised figures illustrate a
skin specializations addressed biological roles before they
feathered dinosaur, clarify embryonic development of the uro-
addressed flight. This is discussed in the chapter on integu-
genital system; and various changes have been made in figures
ment (chapter 6) with new supportive illustrations.
elsewhere. Scientific references are available to the students,
Shark Skin. In addition to favorably affecting fluid flow
online, if they would like to follow up or read more about
across their surface, shark placoid scales also bristle, lift up,
a particular subject. The accompanying laboratory dissection
when boundary layer separation begins to occur to reduce its
guide (authored with E. J. Zalisko) is closely cross-referenced
effects. This newly discovered feature of shark skin is now
to this textbook. In addition, selective functional laborato-
also included in chapter 6.
ries are available, online, to provide students with firsthand
Evo-Devo. I have built on the genetic section on evo-
experience of working between the anatomy and its functional
lution and development (chapter 5) introduced in earlier
and evolutionary significance.
editions. Examples throughout show how master control
Serving Instructors. This seventh edition—new,
genes (Hox genes) and developmental genes preside over the
revised, updated—can serve as reference and resource sup-
construction of the vertebrate body and its various systems.
port for the course you put together on vertebrates. In
For example, in chapter 8 wonderful experimental work in
addition to this, resources are available to you online. The
mice using “knockout” techniques has shown how various
functional laboratories may be downloaded and used as they
Hox genes control differentiation of the mammalian axial
supplement your course. PowerPoint images, chapter by
column. In the concluding chapter, I emphasize how these
chapter, are available online along with additional images
special evo-devo gene sets provide the basis for understant-
from McGraw-Hill that can be used to compose lectures
ing the genetic mechanisms underlying major evolutionary
and laboratory presentations.
changes.
Phylogenetic Relationships. Thanks to continuing use
of improved genetic and morphological data sets, phyloge- Supplements
netic relationships are becoming better resolved, and natural
groups are emerging from this analysis with better clarity.
Comparative Vertebrate Anatomy:
This is the basis for revision in chapter 3, but these updated
A Laboratory Dissection Guide
phylogenies are carried forward throughout the book or new
phylogenies added, as for example with the phylogeny of the Newly revised, Comparative Vertebrate Anatomy: A Laboratory
axial column (chapter 8). Dissection Guide, Seventh Edition, by Kenneth V. Kardong
Lungs and the Rise of Archosaurs. The especially efficient and Edward J. Zalisko, is now available. At the end of this
lungs of birds are well known with air sacs and one-way flow dissection guide, the authors include a Student Art Notebook.
of air. But new experimental evidence (chapter 11) identifies This notebook, promoted by students, is a reprinted collection
similar one-way air flow, even without air sacs, in crocodiles. of the most important and commonly used dissection figures
If true of archosaurs in general, it may represent a respira- in the current edition of the laboratory manual. It addresses
tory adaptation to low oxygen levels in the early Mesozoic a frustration inherent in most dissection guides, especially
and account for the rise of Archosaurs. when comparing homologous systems between representative

Preface xvii
animals, of having to flip between text and distantly placed gain access to powerful web tools for learning—including
illustrations. This laboratory manual weaves the functional full text search, notes and highlighting, and email tools for
and evolutionary concepts from this textbook, Vertebrates: sharing notes between classmates. www.CourseSmart.com
Comparative Anatomy, Function, Evolution, into the morpho-
logical details of the laboratory exercises. Using icons, the lab-
oratory manual identifies cross references to this textbook, so
Art and Artists
students can quickly move from the dissection guide to this Please indulge me a final moment of lament. For illustra-
textbook to consult the expanded treatment of function and tions, modern textbooks have substituted photographs
evolution. Each chapter of the dissection guide first introduces and figures made of computer generated plastic figures,
the system, makes comparisons, and demonstrates common especially when rending molecular events. Nothing wrong
themes in the animal systems. Then the written text carefully with this but what has been eliminated is involvement of
guides students through dissections, which are richly illus- the human touch, directly, namely the artist. Preparing this
trated. Anatomical terms are boldfaced and concepts itali- textbook has given me the chance, at no expense to students
cized. The dissection guide is written so that instructors have by the way, of engaging some of the best artists of our day.
the flexibility to tailor-make the laboratory to suit their needs. They bring a sharp eye and traditional talent to rendering of
enlightening pieces of art. Many have contributed, but one
Website for Vertebrates: Comparative is L. Laszlo Meszoly (Harvard University), who has contrib-
Anatomy, Function, Evolution, Seventh uted special figures to this and earlier editions. Another is
Edition Kathleen M. Bodley, whose remarkable ability to render soft
tissue in particular has enriched our dissection guide and is
A website for this textbook, available at www.mhhe.com/ quite stunning. Her work also graces the cover of this text-
kardong7e, includes further useful information upon which book. My thanks to these two in particular. This is a won-
instructors can depend and students can consult. Here can derful tradition of scientific illustration they carry forward.
be found the functional laboratories, helpful in a linked
laboratory if available, or helpful selectively in lecture. End-
of-chapter selected references, giving students a start into Acknowledgments
the literature, are located here.
I am indebted to reviewers, students, and colleagues who
have generously shared with me their suggestions to improve
Biology Digitized Video Clips this edition of the textbook. My hope is that these colleagues
will see, if not their point of view, at least their influence
McGraw-Hill is pleased to offer digitized biology video
within this edition, and accept my sincere thanks for their
clips on DVD. Licensed from some of the highest-quality
thoughtful suggestions and criticisms. For their special help
science video producers in the world, these brief segments
in this edition and earlier editions I recognize:
range from about five seconds to just under three min-
utes in length and cover all areas of general biology, from Daniel Blackburn
cells to ecosystems. Engaging and informative, McGraw- Trinity College
Hill’s digitized biology videos will help capture students’
Richard W. Blob
interest while illustrating key biological concepts and pro-
Clemson University
cesses. Includes video clips on mitosis, Darwin’s finches,
amoeba locomotion, tarantula defense, nematodes, bird/ Carol Britson
water buffalo mutualism, echinoderms, and much more! University of Mississippi
ISBN: 978-0-07-312155-0 (MHID: 0-07-312155-X)
Stephen Burnett
Clayton State University
Electronic Textbook George Cline
CourseSmart is a new way for faculty to find and review Jacksonville State University
eTextbooks. It’s also a great option for students who are
C. G. Farmer
interested in accessing their course materials digitally and
University of Utah
saving money. CourseSmart offers thousands of the most
commonly adopted textbooks across hundreds of courses T. H. Frazetta
from a wide variety of higher education publishers. It is the University of Illinois
only place for faculty to review and compare the full text
Nick Geist
of a textbook online, providing immediate access without
Sonoma State University
the environmental impact of requesting a print copy. At
CourseSmart, students can save up to 50% off the cost of Ira F. Greenbaum
a print book, reduce their impact on the environment, and Texas A & M University

xviii Preface
Maria Laura Habegger It has been a special pleasure for me to work previously
University of South Florida with several especially supportive and helpful colleagues. In
particular, I note the extensive help of Christine M. Janis in
Christine M. Janis
several difficult chapters, as well as the patient and especially
Brown University
informative education I received on regenerating bird feathers
Amy W. Lang from P. F. A. Maderson and W. J. Hillenius. I also want to
University of Alabama note especially the help from Ira R. Greenbaum for engaging in
a pleasurable and productive exchange on all things vertebrate.
Jon M. Mallatt
For answering my queries, supplying me with their
Washington State University
critical thoughts, and/or for earlier participation in this and
Sue Ann Miller previous editions, I gratefully recognize the following: Neil
Hamilton College F. Anderson, Alejandra Arreola, Miriam A. Ashley-Ross,
Philip J. Motta Ann Campbell Burke, Walter Bock, Warren W. Burggren,
University of South Florida Anindo Choudhury, Michael Collins, Mason Dean, Ken P.
Dial, Alan Feduccia, Adrian Grimes, Maria Laura Habegger,
Barbara Pleasants Linda Holland, Marge Kemp, Amy W. Lang, William T.
Iowa State University Maple, Jessie Maisano, David N. M. Mbora, Philip Motta,
Calvin A. Porter David O. Norris, R. Glenn Northcutt, Kevin Padian, Kathryn
Xavier University of Louisiana Sloan Ponnock, Michael K. Richardson, Timothy Rowe, John
Ruben, J. Matthias Starck, James R. Stewart, Billie J. Swalla,
Tamara L. Smith Steven Vogel, Alan Walker, and Bruce A. Young.
Westridge School I am indebted to the patient, able, and supportive
Jeffrey Thomas people at McGraw-Hill who were so important in bring-
Queens University Charlotte ing this revised seventh edition along. I thank again the
McGraw-Hill field staff who link the summary effort of all
David Varricchio who helped in this revision to faculty and students who use
Montana State University it. In turn, these field reps return your comments of what
Mindy Walker you do and do not like, and thereby aid in the improve-
Rockhurst University ment of this textbook, making it a shared work in prog-
ress. Lori Bradshow was indispensible as Developmental
Andrea Ward
Editor. It was a pleasure to work with Ligo Alex and her
Adelphi University
talented team of copyeditors at SPi-Global.
Jeanette Wyneken To friends and family I remain grateful and thank them
Florida Atlantic University for their support during various editions of this textbook.

Preface xix
This page intentionally left blank
 C H A P T E R
1
Introduction

Comparative Vertebrate Morphology Evolutionary Morphology

Designs of Students Function and Biological Role
Vertebrate Design—Form and Function Preadaptation
Grand Design Evolution as Remodeling

Historical Predecessors—Evolution Phylogeny

The Process behind the Change Of Bean Stalks and Bushes
Linnaeus Simplification
Naturalists Patterns of Phylogeny
J-B. de Lamarck Grades and Clades
Acquired Characteristics
Upward to Perfection Paleontology
Natural Selection Fossilization and Fossils
A. R. Wallace Recovery and Restoration
Charles Darwin From Animal to Fossil
Critics and Controversy Dating Fossils
Stratigraphy
Historical Predecessors—Morphology Index Fossils
Georges Cuvier Radiometric Dating
Richard Owen Geological Ages

Why Are There No Flying Elephants? Tools of the Trade

The Question
Morphological Concepts The Function
Similarities The Biological Role
Symmetry
Segmentation Overview

Comparative Vertebrate Morphology

Comparative morphology deals with anatomy and its signifi- For example, different fishes have different tail shapes.
cance. We focus on animals, in particular vertebrate animals, In the homocercal tail, both lobes are equal in size, making
and the significance these organisms and their structure may the tail symmetrical (figure 1.1a). In the heterocercal tail,
hold. The use of “comparison” in comparative morphology is found in sharks and a few other groups, the upper lobe is
not just a convenience. It is a tool. Comparison of structures elongated (figure 1.1b). Why this difference? The homocer-
throws similarities and differences into better relief. Com- cal tail is found in teleost fishes—salmon, tuna, trout,
parison emphasizes the functional and evolutionary themes and the like. These fishes have a swim bladder, an air-filled
vertebrates carry within their structures. Comparison also sac that gives their dense bodies neutral buoyancy. They nei-
helps formulate the questions we might ask of structure. ther sink to the bottom nor bob to the surface, so they need

1
 centimeter of muscle fiber cross section. Rather than test-
ing all vertebrate muscles, a time-consuming process, we
usually assume that other muscles of similar cross section
produce a similar force (other things being equal). The
discovery of force production in some muscles is extrapo-
(a) Homocercal tail
lated to others. In medicine, the comparative effects of
drugs on rabbits or mice are extrapolated to tentative
use in humans. Of course, the assumed similarities upon
which an extrapolation is based often do not hold in our
analysis. Insight into the human female reproductive
Pectoral
cycle is best obtained if we compare the human cycle with
(b) Heterocercal tail fin those in higher primates because primate reproductive
cycles, including the human one, differ significantly from
FIGURE 1.1 Homocercal and heterocercal fish those of other mammals.
tails. Form differs because function differs. (a) Sweeping, Extrapolation allows us to make testable predictions.
side-to-side movements of the homocercal tail, common Where tests do not support an extrapolation, science is
in fishes with neutral buoyancy, drive the body forward. well served because this forces us to reflect on the assump-
(b) Swimming strokes of the heterocercal tail propel the fish
tions behind the comparison, perhaps to re-examine the
forward, and motion of the long extended upper lobe imparts an
upward lift to the posterior end of the fish. Sharks, which are a
initial analysis of structures and to return with improved
good deal denser than water, need the upward forces provided hypotheses about the animals or systems of interest. Com-
by the extended lobe of the tail to counteract a tendency parison itself is not just a quick and easy device. The point
to sink. to emphasize is this: Comparison is a tool of insight that
guides our analysis and helps us set up hypotheses about the
basis of animal design.

not struggle to keep their vertical position in the water.

Sharks, however, lack swim bladders, and so tend to sink. Designs of Students
The extended lobe of their heterocercal tail provides lift dur- Such philosophical niceties, however, usually do not entice
ing swimming to help counteract this sinking tendency. So, students into their first course in morphology. Most stu-
the differences in structure, homocercal versus heterocer- dents first venture into a course in vertebrate morphol-
cal, are related to differences in function. Why an animal ogy on their way into some other profession. Customarily,
is constructed in a particular way is related to the functional morphology courses prepare students headed into techni-
requirements the part serves. Form and function are cou- cal fields such as human medicine, dentistry, or veterinary
pled. Comparison of parts highlights these differences and medicine. Vertebrate form and function will be the founda-
helps us pose a question. Functional analysis helps answer tion for these medical fields. As will, for example, diagnostic
our question and gives us a better understanding of animal medicine that benefits from the development of anatomi-
design. Functional morphology is the discipline that relates cally and functionally correct prosthetic devices to replace
a structure to its function. injured body parts lost to disease and trauma.
Comparative analysis thus deploys various methods to In addition, morphology is important to taxonomists
address different biological questions. Generally, compara- who use the structure of animals to define characters. In
tive analysis is used either in a historical or a nonhistorical turn, these characters are used as the basis for establishing
context. When we address historical questions, we exam- relationships between species.
ine evolutionary events to work out the history of life. For Morphology is central to evolutionary biology as
example, on the basis of the comparison of characters, we well. Many scientists, in fact, would like to see a discipline
may attempt to construct classifications of organisms and devoted to the combined subject, namely, evolutionary
the evolutionary phylogeny of the group. Often such his- morphology. Evidence of past evolutionary changes is
torical comparisons are not restricted to classification alone inscribed in animal structure. Within the amphibian limb
but center on the process of evolution behind morphological are the structural reminders of its fish-fin ancestry; within
units, such as jaws, limbs, or eyes. the wing of a bird are the evidences of its derivation from
When we make nonhistorical comparisons, as is the reptilian forelimb. Each modern group living today car-
frequently the case, we look outside an evolutionary con- ries forward mementos of the evolutionary course traveled
text, with no intention of concluding with a classification by its ancestors. For many biologists, a study of the morpho-
or elucidation of an evolutionary process. Nonhistorical logical products of the past gives insight into the processes
comparisons are usually extrapolative. For example, by that produced them, insight into the natural forces that
testing a few vertebrate muscles, we may demonstrate drove evolutionary changes, and insight into the limitations
that they produce a force of 15 N (newtons) per square of evolutionary change.

2 Chapter One
 BOX ESSAY 1.1
The Scientific Method—What They
Tell You and What They Don’t

Formally, the scientific method includes of transmission and years later win him a transplanted legs were brown and some
formulation of a hypothesis, design of a share of the Nobel Prize. were white. It then dawned on him that
test, carrying out of an experiment, analy- Loewi’s experiment was as simple as the stump of the transplanted limb might
sis of results, corroboration or falsification it was elegant. He removed the heart and in some instances include nearby neural
of the hypothesis, and formulation of a new associated vagus nerve from the body of a crest cells, but not in all instances. Neural
hypothesis. In practice, science does not frog and isolated them in a beaker of saline. crest cells form first on top of the nerve
follow such a stately and linear sequence. Next he stimulated the free vagus nerve, tube and then normally disperse about the
Broken equipment, uncooperative animals, causing the heart rate to slow. Loewi then embryo. He tried limbs with and without
paperwork, and committee meetings all con- took this saline and poured it over another accompanying neural crest cells. That was
spire against the well-laid plans of mice, men, isolated frog heart from which the vagus it. Those brown leghorn limbs with neu-
and women. It is more than the “expected had been removed. The rate of this heart ral crest cells produced brown feathers.
unexpected” that affects experiments and also slowed, providing clear evidence that a Those without lacked pigment cells and
tests one’s blood pressure. The intellectual chemical produced by the stimulated vagus were white. Eastlick, who started out
questions themselves do not always find sat- nerve controlled heart rate. Transmission working on muscles, confirmed what a few
isfying answers. Accidents, chance, and even between nerve (vagus) and organ (heart) had guessed at the time, namely, that one
dreams are part of the creative process. was brought about by chemical agents, not derivative of neural crest cells is pigment
Otto Loewi shared the 1936 Nobel by electrical currents. cells that give feathers their color.
Prize in medicine with Henry Dale for As a young cell biologist, Herbert Alexander Fleming (1881–1955), while
demonstrating that nerve impulses pass Eastlick began a series of experiments to studying bacteria, noticed that when molds
from one nerve cell to the next in series pursue his interest in embryonic develop- occasionally contaminated cultures, the
across the space between them, the syn- ment of young muscle. He transplanted bacteria next to the molds failed to grow.
apse, by a chemical transmitter. Early in the still formative hindlimbs of a chick to Hundreds of students and fellow bacteri-
the twentieth century, opinion had been the side of a host chick while the host was ologists before Fleming had seen molds
divided between those physiologists who still developing in its egg. The transplanted and likely noticed the stunted growth of
felt that this neuron-to-neuron trans- hindlimbs were usually received and grew bacteria. But it was Fleming’s curiosity that
mission was chemical and those who felt well enough on the side of the host chick precipitated the serious question, “What
that it was electrical. A definitive experi- to allow study. One day, when a local sup- causes this reaction?” In answering it, he
ment settling the issue was needed. One plier was temporarily out of the white leg- discovered that molds produced penicil-
night, when he was deep in sleep, the horn eggs Eastlick had used, he substituted lin, a bacterial inhibitor. Fleming’s question
definitive experiment came to Loewi and brown leghorns, a breed with brown feath- opened the way for development of a new
woke him. Relieved and satisfied, he went ers. After three days of incubation, one egg branch of pharmacology and a new indus-
back to sleep looking forward to the next was opened and both leg-forming areas of try. His answer established the basis of dis-
day. When awaking the next morning, he a brown leghorn were transplanted to a ease control through antibiotics.
remembered dreaming the experiment white leghorn host. Results were puzzling. Testing of a well-crafted hypoth-
but had forgotten what it was. Several The right transplanted leg from the brown esis forms the center of the scientific
frustrating weeks passed until, once again leghorn developed brown feathers, the method. But where the next hypothesis
deep in sleep, Loewi dreamt the same left transplanted leg from the same brown comes from cannot always be predicted.
dream, and the experimental design came leghorn developed white feathers. What A thought in the middle of the night, an
back. Leaving nothing to chance this time, caused these contrary results? experiment gone wrong, a close observa-
he got up, dressed, and in the middle of Eastlick checked his notes, repeated tion of the ordinary, these too may inspire
the night went to his laboratory to begin his experiments, and used great care in a new scientific hypothesis and are part of
the experiment that would settle the issue performing more transplants. Still some the method of science.

Vertebrate Design—Form and Function that the wings of birds, tails of fishes, or hair of mammals
Morphology offers more than charitable assistance to other arose for the adaptive advantages each structure provided,
disciplines. The study of morphology provides its own plea- and so they were favored by natural selection. Certainly this
sure. It raises unique questions about structure and offers a is true, but it is only a partial explanation for the presence of
method to address these questions. In brief, vertebrate mor- these respective features in bird, fish, and mammal designs.
phology seeks to explain vertebrate design by elucidating the The external environment in which an animal design must
reasons for and processes that produce the basic structural serve certainly brings to bear evolutionary pressures on its
plan of an organism. For most scientists today, evolutionary survival, and thus on those anatomical features of its design
processes explain form and function. We might hear it said that convey adaptive benefits.

Introduction 3
 Internal structure itself also affects the kinds of designs But, I am getting ahead of the story. We have not
that do or do not appear in animals. No terrestrial vertebrate had an easy intellectual journey in reaching the clarity of
rolls along on wheels. No aerial vertebrate flies through the morphological concepts we seem to enjoy at the moment.
air powered by a rotary propeller. Natural selection alone The principles were not always so obvious, the evidence
cannot explain the absence of wheels in vertebrates. It is not always so clear. In fact, some issues prevalent over 100
quite possible to imagine that wheels, were they to appear years ago remain unresolved. The significance of underly-
in certain terrestrial vertebrates, would provide considerable ing structure to the evolution of design, central to much
adaptive advantages and be strongly favored by natural selec- of biology early in the nineteenth century, is only recently
tion. In part, the explanation lies in the internal limitations of being re-examined for its potential contribution to modern
the structure itself. Rotating wheels could not be nourished morphology. Morphology has often been internally beset by
through blood vessels nor innervated with nerves without unhappy contentions between those scientists centered on
quickly twisting these cords into knots. Wheels and propellers structure and those centered on evolution. To some extent,
fall outside the range of structural possibility in vertebrates. the fundamental principles of both structure and evolution
Structure itself contributes to design by the possibilities it have grown from different intellectual sources and different
creates; evolution contributes to design by the favored struc- intellectual outlooks. To understand this, we need to exam-
tures it preserves. We must consult both structure and evolu- ine the historical development of morphology. Later in this
tion to understand overall design. That is why we turn to the chapter, we examine the intellectual roots of theories about
discipline of morphology. It is one of the few modern sci- structure. But first, let’s look to the intellectual roots of the-
ences that addresses the natural unity of both structure (form ories about evolution.
and function) and evolution (adaptation and natural selec-
tion). By wrapping these together in an integrated approach,
morphology contributes a holistic analysis of the larger issues Historical Predecessors—Evolution
before contemporary biology. Morphology is concerned cen-
The concept of evolution is tied to the name Charles Darwin
trally with the emergent properties of organisms that make
(figure 1.2). Yet most persons are surprised to learn that
them much more than the reduced molecules of their parts.
Darwin was not first, nor was he ever foremost, in proposing
that organisms evolve. In fact, the idea of change through
Grand Design time in animals and plants dates back to ancient schools
of Greek philosophy. Over 2,500 years ago, Anaximander
Vertebrate design is complex, often elegant, and sometimes
developed ideas about the course of change from fishlike
remarkably precise. To many early-day morphologists, this
and scaly animals to land forms. Empedocles saw original
complexity, this elegance, and this precision implied the direct
creatures come together in oddly assembled ways—humans
intervention of a divine hand in guiding the production of
such sophisticated designs. However, not everyone was con-
vinced. Spectacular mountain ranges do not require recourse
to divine intervention to explain them. Plate tectonics offers
a natural explanation. Under pressure from colliding tectonic
plates, the Earth’s crust crumples to produce these ranges.
With knowledge, scientific explanations uncover the myster-
ies that shroud geological events.
Similarly, biology has found satisfying natural expla-
nations to replace what were once assumed to be direct
divine causes. Modern principles of evolution and structural
biology offer a fresh approach to vertebrate design and an
insight into the processes responsible for producing that
design. Just as processes of plate tectonics help geologists
understand the origin of the Earth’s surface features, struc-
tural and evolutionary processes help biologists understand
the origin of plant and animal life. Life on Earth is a product
of these natural processes. Humans are not exempt nor are
FIGURE 1.2 Charles Darwin (1809–1882), about
we given special dispensation from these processes. Like our
30 years old and three years back from his voyage aboard
fellow vertebrates, humans too are products of our evolu-
H.M.S. Beagle. Although The Origin of Species was still just a few
tionary past and basic structural plan. The study of morphol- notebooks in length and several decades away from publica-
ogy, therefore, brings us an understanding of the integrated tion, Darwin had several accomplishments behind him, including
processes that forged us. To understand the processes behind his account of The Voyage of the Beagle, a collection of scientific
our design is to understand the product, namely, humans observations. At this time, he was also engaged to his cousin
themselves, both what we are and what we can become. Emma Wedgwood, with whom he would live a happy married life.

4 Chapter One
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