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

Introduction 372
Organization of Muscles 373
Classification of Muscles 373
Skeletal Muscle 374
Cardiac Muscle 375
Smooth Muscle 375
Structure of Skeletal Muscles 375
Tendons 376
Basis of Muscle Contraction 376
Resting and Active Muscle 376
Molecular Mechanisms of Contraction
Muscle Function 377
Muscle Fibers 377
Tension-Length Curves for a Single Muscle Fiber 377
Properties of Muscle Fibers 377
Muscle Organs and Fibers 379
Whole Muscle Force Generation 379
Tension-Length Curves for a Whole Muscle 380
Graded Force 380
Cross-Sectional Area 383
Fiber Orientation 383
Velocity of Shortening 385
Distance of Shortening 385
Bone–Muscle Lever Systems 385
Sequencing of Muscle Actions 387
Overview of Muscle Mechanics 388
Muscle Actions 388
Muscle Homologies 390
Embryonic Origin of Muscles
Postcranial Musculature 392
Appendicular Musculature 392
Axial Musculature 393
Cranial Musculature 393
Jaw and Pharyngeal Musculature
Extrinsic Eye Muscles 393
Comparative Anatomy 394
Postcranial Musculature 394
Gills 416
Gas Bladders 416
Lungs 416
Swim Bladders 416
Cutaneous Respiratory Organs 417
Accessory Air-breathing Organs 417
Breathing and Embryos 418
Ventilatory Mechanisms 421
Cilia 421
Muscular Mechanisms 421
Water Ventilation: Dual Pump 421
376 Air Ventilation: Buccal Pump 422
Air Ventilation: Aspiration Pump 423
Phylogeny 424
Agnathans 424
Elasmobranchs 426
Bony Fishes 427
Overview of Fish Respiration 427
Gills 427
Lungs and Swim Bladders 428
Amphibians 430
Amphibian Larvae 430
Amphibian Adults 432
Reptiles 433
Mammals 435
Ventilation 435
Gas Exchange 437
Birds 437
Form and Function 440
391 Patterns of Gas Transfer 440
Rates of Gas Transfer 443
Breathing in Water 444
Breathing in Air 444
Evolution of Respiratory Organs 444
393
Acid-Base Regulation 444
Ventilation 446
Ciliary Pumps 446
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
Preadaptation 562
Origin of Vertebrates 562
Reproductive System 563
Structure of the Mammalian Reproductive System 563
Embryonic Development 563
Gonads and Gametes 563
Reproductive Tracts 565
Overview 565
Female Reproductive System 567
Ovary 567
Genital Ducts 567
Oviduct 569
Uterus 570
Male Reproductive System 572
Testis 572
Genital Ducts 572
Copulatory Organs 576
Cloaca 582
Urinary Bladder 586
Function and Evolution 588
Potency and Fertility 588
External and Internal Fertilization 588
Delays in Gestation 589
Overview 589
f i f t een
CHAPTER
The Endocrine System
Survey of Endocrine Glands
Thyroid Gland 592
Structure and Phylogeny 592
Function 594
Ultimobranchial Body and Parathyroid Gland
Ultimobranchial Body 596
Parathyroid Gland 597
Form and Function 598
Adrenal Gland 598
Structure and Phylogeny 598
Function 601
Pancreatic Islets 602
Structure and Phylogeny 602
Function 602
Pituitary Gland 604
Structure 604
Phylogeny 606
Function 607
Gonads 610
Pineal Gland 610
Secondary Endocrine Glands 611
Gastrointestinal Tract 611
Kidneys 612
Endocrine Coordination 613
Mammalian Reproduction 613
Male 613
Female 613
Metamorphosis in Frogs 616
Fundamentals of Hormonal Control 620
Functional and Structural Linkage 620
Target Tissue Responses 621
The Endocrine System and the Environment 621
Evolution 622
Overview 624
si xteen
CHAPTER
The Nervous System 625
Introduction 625
Types of Cells within the Nervous System 625
Neuroglia 625
Neurons 625
Transmission of Information 626
Neurosecretory Cells 628
Peripheral Nervous System 628
Spinal Nerves 629
Cranial Nerves 630
Evolution 637
592 Functions of the Peripheral Nervous System 638
Spinal Reflexes 638
592 The Autonomic Nervous System 641
Central Nervous System 646
Embryology 646
596 Spinal Cord 649
Spinal Reflexes 649
Spinal Tracts 650
Brain 651
Phylogeny 651
Form and Function 654
Functional Associations of Parts
of the Central Nervous System 666
Limbic System 666
Overview 669
CHAPTERseventeen
Sensory Organs 671
Introduction 671
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
the first time, several introductory remarks may be helpful,
especially on how this textbook will support your work. First,
the discipline of vertebrate biology is diverse and inclusive. It
brings together themes from molecular biology, genes and
genomes, evolution and embryology, biomechanics and
experimental physiology, and it incorporates continuing and
astonishing new fossils into the vertebrate story. Much of
what you have met in earlier courses you will meet again here
in an integrated way.
Second, to unify these themes, I have again written and
revised this seventh edition within the unifying framework of
form, function, and evolution. The first few chapters set this
up, and the subsequent chapters treat vertebrates system by
system. You may notice that each of these subsequent chap-
ters begins with a discussion of morphology, followed by a
discussion of function and evolution. Each chapter is therefore
self-contained—form, function, evolution.
Third, as a student you likely enter this course after
some background in the sciences, perhaps expecting to equip
yourself with practical knowledge useful later in professional
schools or in health-related careers. Certainly this course, in
part, delivers such practical information. But because ver-
tebrate morphology is an integrative discipline, it brings
together physiology, embryology, behavior, and ecology
and also deploys modern methods of systematics and new
finds in paleontology. Consequently, you will move beyond
memorizing facts in isolation or as an end in themselves,
and instead begin to meet and understand larger concepts
to which the morphology testifies. What may come as a sur-
prise is that many theories, especially evolutionary theories
within vertebrate biology, are still unsettled and unresolved,
inviting a new idea or fresh approach open to anyone. This is
one of the reasons I have included various controversies, and
support your efforts to become engaged in the thinking and
scientific process.
For faculty who have used this textbook before, you
will find it retains a familiar and inviting organization with
the science updated and the student support enhanced. For
those coming to this textbook for the first time, you will
notice that the morphology receives generous treatment
xv
within a phylogenetic context. But, today we expect our stu-
dents to develop academic and professional skills beyond just
facility with anatomical terminology. In general, we expect
our students to develop skills in critical thinking and a facil-
ity with scientific concepts. Each of us will find our own
way of composing a course in vertebrate morphology that
serves such course objectives. This textbook was written to
support such course objectives as individual instructors build
their courses. It is flexible. You can mix and match, change
order to suit your course, and give emphasis to those sys-
tems that most suit the organization of your course. Because
each chapter integrates form, function, and evolution perti-
nent to that system, each chapter is coherent within itself.
Where information or concepts are treated in greater detail
outside a particular chapter, they are cross referenced to help
guide the student and clarify the discussion. Although dis-
cussed in earlier editions, let me repeat the specific strategy
built into this textbook to improve student success and to
help them develop skills in critical thinking and conceptual
understanding.
For the Student
A number of strategic features within the textbook enhance
its usefulness for students. It is richly illustrated with fig-
ures that include new information and provide fresh per-
spectives. Each chapter opens with an outline. Important
concepts and major anatomical terms are boldfaced. Cross
references direct students to other areas of the text where
they can refresh their understanding or clarify an unfamil-
iar subject. Each chapter concludes with a chapter overview,
which draws attention to some of the concepts developed
within the chapter. Box Essays are included along the way
in most chapters. Their purpose is to present subjects or his-
torical events that students should find interesting and, per-
haps from time to time, even fun. A glossary of definitions is
included at the end of the book.
In addition to its practical features, the textbook also
uses selected topics within vertebrate structure to develop
student skills in critical thinking and mastery of concepts
within a coherent framework.
Critical Thinking
Within the sciences, critical thinking is the ability to mar-
shal factual information into a logical, reasoned argument.
Especially if accompanied by a laboratory, a course in vertebrate
morphology delivers hands-on experience with the anatomy of
representative animals. Students can be directly engaged in the
discovery of vertebrate form. But they can be encouraged to go
beyond this. Instructors can lead students into larger issues—
How does it function? How did it evolve? For example, early
on in the textbook, students are introduced to “Tools of the
Trade,” methods by which we empirically examine how parts
work and how we can place organisms within a phylogenetic
context. After a discussion of basic morphology, each chapter
discusses how these systems work and how they evolved.
I have deliberately included new, neglected, or com-
peting views on function and evolution. Many of these ideas
come from Europe, where they have been known for a long
time. Personally, I find many of these ideas compelling, even
elegant. Others strike me, frankly, as thin and unconvinc-
ing. Despite my own skepticism, a few contrary ideas are
included. My purpose is to get students to think about issues
of form, function, and evolution.
Several theories on the evolution of jaws are discussed,
as are several theories of the origin of paired fins. Often stu-
dents expect that today we have the final answers. Students
implore, “Just tell me the answer.” The debate about dino-
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
we know that science is an ongoing process of refinement,
challenge, and sometimes revolutionary change. One Box
Essay sets forth the early case for dinosaur endothermy. That
debate spawned further investigation that now returns to
challenge such a view of dinosaurs as “hot-blooded” beasts.
The second Box Essay on dinosaur endothermy presents
this newer and contrary evidence, and thereby showcases
how, even in extinct animals, it is possible to test hypotheses
about their physiology, morphology, and lifestyles.
Concepts
Vertebrate morphology also helps develop an appreciation
and understanding of the scientific concepts that unite biol-
ogy and reflect on “how” science works. As John A. Moore
put it, science is a “way of knowing” (Moore, American
Zoologist, 1988). Comparative morphology throws into clear
relief differences and similarities between organisms. The
concepts of homology, analogy, and homoplasy help us
understand the basis of these comparative features. Many of
the concepts were birthed in the nineteenth century and have
grown into the guiding themes of biology today. Evolution,
defined as descent with modification through time, is one
of the foundation concepts in biology. Vertebrate morphol-
ogy provides a showcase of adaptive change on the basic
vertebrate body plan. But evolution is change in a highly
xvi Preface
integrated organism, a connected system of parts and their
functions. This too was recognized within the nineteenth
century, suggesting constraints on evolutionary modifica-
tion. Vertebrate morphology provides compelling examples
of how an integrated organism might evolve. For example, a
remarkable fossil record documents an undeniable change in
jaw articulation within synapsids, seeing the two participat-
ing bones (articular, quadrate) of basal synapsids replaced by
two different bones in derived groups, including mammals.
Fossil intermediates between the two conditions mark the
anatomical changes, but they also suggest how functional
changes, which must accompany evolving systems, also
change without disrupting performance.
Within many vertebrate systems, the close coupling of
form and function with lifestyle is illustrated. Built on a basic
vertebrate plan, the tetrapod locomotor system illustrates the
close relationship between limbs and axial skeleton, and the type
of locomotion—flight, cursorial, burrowing. The cardiovascular
system, especially in organisms that exploit water and air, illus-
trates the close relationship between vascular morphology and
the physiological flexibility that permits. The basic concepts of
form, function, and adaptive evolution parade before us as we
move from system to system in vertebrate morphology.
Evolution proceeds most often by remodeling, modi-
fication of a basic underlying plan, not by all new construc-
tion. This is illustrated in the skeletal system, as well as
within the cardiovascular (aortic arches) system.
Organizational Strategy
and Rationale
I have written this book within the unifying framework of
form, function, and evolution. These are common themes
that run throughout. The vertebrate groups are organized
phylogenetically, and their systems discussed within such a
context. Morphology is foremost, but I have developed and
integrated an understanding of function and evolution into
the discussion of anatomy of the various systems. The first
five chapters prepare the way.
Chapter 1 introduces the discipline, evaluates the intel-
lectual predecessors to modern morphology, defines central
concepts, and alerts students to misunderstandings they may
unknowingly bring with them to the study of evolutionary
processes. Chordates and their origins are covered in chapter
2. Considerable attention is given to the neglected protochor-
dates and their evolution. This sets the stage for an extended
discussion of the cast of characters in the vertebrate radiation,
which occupies us for the remainder of the book, beginning
next in chapter 3. Here we discuss vertebrates, their origins,
and basic taxonomic relationships. Chapter 4 introduces
basic concepts of biomechanics and biophysics, preparing for
their use later in understanding aspects of vertebrate design
and function. Chapter 5 includes a summary of descriptive
embryology and concludes with a discussion of the role
embryonic processes play in vertebrate evolutionary events.
 The remaining chapters develop each major system.
Besides carrying overall themes, each chapter internally
follows a consistent organization. Each begins with a basic
introduction to the morphology, and then proceeds to dis-
cuss function and evolution. This way, the overall themes
are repeated in each chapter, bringing consistency of presen-
tation to each chapter and coherence throughout.
New and Expanded
in the Seventh Edition
New fossil finds, modern experimental research, and new
phylogenies continue to enrich vertebrate biology, some-
times solving old questions or surprising us with a new
understanding of how vertebrates function and how they
evolved. Much of this is added to this new edition.
Feathered Dinosaurs. New and remarkable fossil finds
of dinosaurs, especially in China, continue to be discov-
ered, some showing evidence of surface feathers. In other
words, feathers evolved before birds. This means that these
skin specializations addressed biological roles before they
addressed flight. This is discussed in the chapter on integu-
ment (chapter 6) with new supportive illustrations.
Shark Skin. In addition to favorably affecting fluid flow
across their surface, shark placoid scales also bristle, lift up,
when boundary layer separation begins to occur to reduce its
effects. This newly discovered feature of shark skin is now
also included in chapter 6.
Evo-Devo. I have built on the genetic section on evo-
lution and development (chapter 5) introduced in earlier
editions. Examples throughout show how master control
genes (Hox genes) and developmental genes preside over the
construction of the vertebrate body and its various systems.
For example, in chapter 8 wonderful experimental work in
mice using “knockout” techniques has shown how various
Hox genes control differentiation of the mammalian axial
column. In the concluding chapter, I emphasize how these
special evo-devo gene sets provide the basis for understant-
ing the genetic mechanisms underlying major evolutionary
changes.
Phylogenetic Relationships. Thanks to continuing use
of improved genetic and morphological data sets, phyloge-
netic relationships are becoming better resolved, and natural
groups are emerging from this analysis with better clarity.
This is the basis for revision in chapter 3, but these updated
phylogenies are carried forward throughout the book or new
phylogenies added, as for example with the phylogeny of the
axial column (chapter 8).
Lungs and the Rise of Archosaurs. The especially efficient
lungs of birds are well known with air sacs and one-way flow
of air. But new experimental evidence (chapter 11) identifies
similar one-way air flow, even without air sacs, in crocodiles.
If true of archosaurs in general, it may represent a respira-
tory adaptation to low oxygen levels in the early Mesozoic
and account for the rise of Archosaurs.
Turning over Chordates. New developmental genet-
ics, discussed in the previous editions, informs us that the
immediate chordate ancestors flipped over, reversing dor-
sal and ventral surfaces. That view seems to hold still and
therefore remains the surprising basis of the chordate body
plan today.
Updated and Revised. Countless changes and revisions
throughout this new edition have been made, some major,
some small. These changes have corrected misinformation,
updated information, and often better clarified an explana-
tion. For this I am indebted to students, reviewers, and col-
leagues for bringing these suggestions to my attention.
Serving the Student. Features of the textbook have been
further expanded to make its presentation more clear and
inviting. The use of color brightens these sections of the
book. Color has also been used to better correlate and com-
pare structures between figures in these chapters. Where fea-
sible, within color signatures, for example, I have added more
color to the illustrations. Many illustrations are new, revised,
or relabeled to improve clarity. For example, besides those
illustrations mentioned earlier, new/revised figures illustrate a
feathered dinosaur, clarify embryonic development of the uro-
genital system; and various changes have been made in figures
elsewhere. Scientific references are available to the students,
online, if they would like to follow up or read more about
a particular subject. The accompanying laboratory dissection
guide (authored with E. J. Zalisko) is closely cross-referenced
to this textbook. In addition, selective functional laborato-
ries are available, online, to provide students with firsthand
experience of working between the anatomy and its functional
and evolutionary significance.
Serving Instructors. This seventh edition—new,
revised, updated—can serve as reference and resource sup-
port for the course you put together on vertebrates. In
addition to this, resources are available to you online. The
functional laboratories may be downloaded and used as they
supplement your course. PowerPoint images, chapter by
chapter, are available online along with additional images
from McGraw-Hill that can be used to compose lectures
and laboratory presentations.
Supplements
Comparative Vertebrate Anatomy:
A Laboratory Dissection Guide
Newly revised, Comparative Vertebrate Anatomy: A Laboratory
Dissection Guide, Seventh Edition, by Kenneth V. Kardong
and Edward J. Zalisko, is now available. At the end of this
dissection guide, the authors include a Student Art Notebook.
This notebook, promoted by students, is a reprinted collection
of the most important and commonly used dissection figures
in the current edition of the laboratory manual. It addresses
a frustration inherent in most dissection guides, especially
when comparing homologous systems between representative
Preface xvii
animals, of having to flip between text and distantly placed
illustrations. This laboratory manual weaves the functional
and evolutionary concepts from this textbook, Vertebrates:
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
students can quickly move from the dissection guide to this
textbook to consult the expanded treatment of function and
evolution. Each chapter of the dissection guide first introduces
the system, makes comparisons, and demonstrates common
themes in the animal systems. Then the written text carefully
guides students through dissections, which are richly illus-
trated. Anatomical terms are boldfaced and concepts itali-
cized. The dissection guide is written so that instructors have
the flexibility to tailor-make the laboratory to suit their needs.
Website for Vertebrates: Comparative
Anatomy, Function, Evolution, Seventh
Edition
A website for this textbook, available at www.mhhe.com/
kardong7e, includes further useful information upon which
instructors can depend and students can consult. Here can
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
the literature, are located here.
Biology Digitized Video Clips
McGraw-Hill is pleased to offer digitized biology video
clips on DVD. Licensed from some of the highest-quality
science video producers in the world, these brief segments
range from about five seconds to just under three min-
utes in length and cover all areas of general biology, from
cells to ecosystems. Engaging and informative, McGraw-
Hill’s digitized biology videos will help capture students’
interest while illustrating key biological concepts and pro-
cesses. Includes video clips on mitosis, Darwin’s finches,
amoeba locomotion, tarantula defense, nematodes, bird/
water buffalo mutualism, echinoderms, and much more!
ISBN: 978-0-07-312155-0 (MHID: 0-07-312155-X)
Electronic Textbook
CourseSmart is a new way for faculty to find and review
eTextbooks. It’s also a great option for students who are
interested in accessing their course materials digitally and
saving money. CourseSmart offers thousands of the most
commonly adopted textbooks across hundreds of courses
from a wide variety of higher education publishers. It is the
only place for faculty to review and compare the full text
of a textbook online, providing immediate access without
the environmental impact of requesting a print copy. At
CourseSmart, students can save up to 50% off the cost of
a print book, reduce their impact on the environment, and
xviii Preface
gain access to powerful web tools for learning—including
full text search, notes and highlighting, and email tools for
sharing notes between classmates. www.CourseSmart.com
Art and Artists
Please indulge me a final moment of lament. For illustra-
tions, modern textbooks have substituted photographs
and figures made of computer generated plastic figures,
especially when rending molecular events. Nothing wrong
with this but what has been eliminated is involvement of
the human touch, directly, namely the artist. Preparing this
textbook has given me the chance, at no expense to students
by the way, of engaging some of the best artists of our day.
They bring a sharp eye and traditional talent to rendering of
enlightening pieces of art. Many have contributed, but one
is L. Laszlo Meszoly (Harvard University), who has contrib-
uted special figures to this and earlier editions. Another is
Kathleen M. Bodley, whose remarkable ability to render soft
tissue in particular has enriched our dissection guide and is
quite stunning. Her work also graces the cover of this text-
book. My thanks to these two in particular. This is a won-
derful tradition of scientific illustration they carry forward.
Acknowledgments
I am indebted to reviewers, students, and colleagues who
have generously shared with me their suggestions to improve
this edition of the textbook. My hope is that these colleagues
will see, if not their point of view, at least their influence
within this edition, and accept my sincere thanks for their
thoughtful suggestions and criticisms. For their special help
in this edition and earlier editions I recognize:
Daniel Blackburn
Trinity College
Richard W. Blob
Clemson University
Carol Britson
University of Mississippi
Stephen Burnett
Clayton State University
George Cline
Jacksonville State University
C. G. Farmer
University of Utah
T. H. Frazetta
University of Illinois
Nick Geist
Sonoma State University
Ira F. Greenbaum
Texas A & M University
Maria Laura Habegger
University of South Florida
Christine M. Janis
Brown University
Amy W. Lang
University of Alabama
Jon M. Mallatt
Washington State University
Sue Ann Miller
Hamilton College
Philip J. Motta
University of South Florida
Barbara Pleasants
Iowa State University
Calvin A. Porter
Xavier University of Louisiana
Tamara L. Smith
Westridge School
Jeffrey Thomas
Queens University Charlotte
David Varricchio
Montana State University
Mindy Walker
Rockhurst University
Andrea Ward
Adelphi University
Jeanette Wyneken
Florida Atlantic University
It has been a special pleasure for me to work previously
with several especially supportive and helpful colleagues. In
particular, I note the extensive help of Christine M. Janis in
several difficult chapters, as well as the patient and especially
informative education I received on regenerating bird feathers
from P. F. A. Maderson and W. J. Hillenius. I also want to
note especially the help from Ira R. Greenbaum for engaging in
a pleasurable and productive exchange on all things vertebrate.
For answering my queries, supplying me with their
critical thoughts, and/or for earlier participation in this and
previous editions, I gratefully recognize the following: Neil
F. Anderson, Alejandra Arreola, Miriam A. Ashley-Ross,
Ann Campbell Burke, Walter Bock, Warren W. Burggren,
Anindo Choudhury, Michael Collins, Mason Dean, Ken P.
Dial, Alan Feduccia, Adrian Grimes, Maria Laura Habegger,
Linda Holland, Marge Kemp, Amy W. Lang, William T.
Maple, Jessie Maisano, David N. M. Mbora, Philip Motta,
David O. Norris, R. Glenn Northcutt, Kevin Padian, Kathryn
Sloan Ponnock, Michael K. Richardson, Timothy Rowe, John
Ruben, J. Matthias Starck, James R. Stewart, Billie J. Swalla,
Steven Vogel, Alan Walker, and Bruce A. Young.
I am indebted to the patient, able, and supportive
people at McGraw-Hill who were so important in bring-
ing this revised seventh edition along. I thank again the
McGraw-Hill field staff who link the summary effort of all
who helped in this revision to faculty and students who use
it. In turn, these field reps return your comments of what
you do and do not like, and thereby aid in the improve-
ment of this textbook, making it a shared work in prog-
ress. Lori Bradshow was indispensible as Developmental
Editor. It was a pleasure to work with Ligo Alex and her
talented team of copyeditors at SPi-Global.
To friends and family I remain grateful and thank them
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-
cance. We focus on animals, in particular vertebrate animals,
and the significance these organisms and their structure may
hold. The use of “comparison” in comparative morphology is
not just a convenience. It is a tool. Comparison of structures
throws similarities and differences into better relief. Com-
parison emphasizes the functional and evolutionary themes
vertebrates carry within their structures. Comparison also
helps formulate the questions we might ask of structure.
For example, different fishes have different tail shapes.
In the homocercal tail, both lobes are equal in size, making
the tail symmetrical (figure 1.1a). In the heterocercal tail,
found in sharks and a few other groups, the upper lobe is
elongated (figure 1.1b). Why this difference? The homocer-
cal tail is found in teleost fishes—salmon, tuna, trout,
and the like. These fishes have a swim bladder, an air-filled
sac that gives their dense bodies neutral buoyancy. They nei-
ther sink to the bottom nor bob to the surface, so they need

1

(a) Homocercal tail
Pectoral
(b) Heterocercal tail fin
FIGURE 1.1 Homocercal and heterocercal fish
tails. Form differs because function differs. (a) Sweeping,
side-to-side movements of the homocercal tail, common
in fishes with neutral buoyancy, drive the body forward.
(b) Swimming strokes of the heterocercal tail propel the fish
forward, and motion of the long extended upper lobe imparts an
upward lift to the posterior end of the fish. Sharks, which are a
good deal denser than water, need the upward forces provided
by the extended lobe of the tail to counteract a tendency
to sink.
not struggle to keep their vertical position in the water.
Sharks, however, lack swim bladders, and so tend to sink.
The extended lobe of their heterocercal tail provides lift dur-
ing swimming to help counteract this sinking tendency. So,
the differences in structure, homocercal versus heterocer-
cal, are related to differences in function. Why an animal
is constructed in a particular way is related to the functional
requirements the part serves. Form and function are cou-
pled. Comparison of parts highlights these differences and
helps us pose a question. Functional analysis helps answer
our question and gives us a better understanding of animal
design. Functional morphology is the discipline that relates
a structure to its function.
Comparative analysis thus deploys various methods to
address different biological questions. Generally, compara-
tive analysis is used either in a historical or a nonhistorical
context. When we address historical questions, we exam-
ine evolutionary events to work out the history of life. For
example, on the basis of the comparison of characters, we
may attempt to construct classifications of organisms and
the evolutionary phylogeny of the group. Often such his-
torical comparisons are not restricted to classification alone
but center on the process of evolution behind morphological
units, such as jaws, limbs, or eyes.
When we make nonhistorical comparisons, as is
frequently the case, we look outside an evolutionary con-
text, with no intention of concluding with a classification
or elucidation of an evolutionary process. Nonhistorical
comparisons are usually extrapolative. For example, by
testing a few vertebrate muscles, we may demonstrate
that they produce a force of 15 N (newtons) per square
2 Chapter One
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-
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
cycle is best obtained if we compare the human cycle with
those in higher primates because primate reproductive
cycles, including the human one, differ significantly from
those of other mammals.
Extrapolation allows us to make testable predictions.
Where tests do not support an extrapolation, science is
well served because this forces us to reflect on the assump-
tions behind the comparison, perhaps to re-examine the
initial analysis of structures and to return with improved
hypotheses about the animals or systems of interest. Com-
parison itself is not just a quick and easy device. The point
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.
Designs of Students
Such philosophical niceties, however, usually do not entice
students into their first course in morphology. Most stu-
dents first venture into a course in vertebrate morphol-
ogy on their way into some other profession. Customarily,
morphology courses prepare students headed into techni-
cal fields such as human medicine, dentistry, or veterinary
medicine. Vertebrate form and function will be the founda-
tion for these medical fields. As will, for example, diagnostic
medicine that benefits from the development of anatomi-
cally and functionally correct prosthetic devices to replace
injured body parts lost to disease and trauma.
In addition, morphology is important to taxonomists
who use the structure of animals to define characters. In
turn, these characters are used as the basis for establishing
relationships between species.
Morphology is central to evolutionary biology as
well. Many scientists, in fact, would like to see a discipline
devoted to the combined subject, namely, evolutionary
morphology. Evidence of past evolutionary changes is
inscribed in animal structure. Within the amphibian limb
are the structural reminders of its fish-fin ancestry; within
the wing of a bird are the evidences of its derivation from
the reptilian forelimb. Each modern group living today car-
ries forward mementos of the evolutionary course traveled
by its ancestors. For many biologists, a study of the morpho-
logical products of the past gives insight into the processes
that produced them, insight into the natural forces that
drove evolutionary changes, and insight into the limitations
of evolutionary change.
 BOX ESSAY 1.1
The Scientific Method—What They
Tell You and What They Don’t

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

Vertebrate Design—Form and Function
Morphology offers more than charitable assistance to other
disciplines. The study of morphology provides its own plea-
sure. It raises unique questions about structure and offers a
method to address these questions. In brief, vertebrate mor-
phology seeks to explain vertebrate design by elucidating the
reasons for and processes that produce the basic structural
plan of an organism. For most scientists today, evolutionary
processes explain form and function. We might hear it said
that the wings of birds, tails of fishes, or hair of mammals
arose for the adaptive advantages each structure provided,
and so they were favored by natural selection. Certainly this
is true, but it is only a partial explanation for the presence of
these respective features in bird, fish, and mammal designs.
The external environment in which an animal design must
serve certainly brings to bear evolutionary pressures on its
survival, and thus on those anatomical features of its design
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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 Part 2

T HE Willoweses came back to London about the second week in

September. For many years the children’s schooling had governed the
date of their return; and when the children had grown too old for school,
the habit had grown too old to be broken. There was also a further reason.
The fallen leaves, so Henry and Caroline thought, made the country
unhealthy after the second week in September. When Laura was younger
she had sometimes tried to argue that, even allowing the unhealthiness of
fallen leaves, leaves at that time of year were still green upon the trees. This
was considered mere casuistry. When they walked in Kensington Gardens
upon the first Sunday morning after their return, Caroline would point along
the tarnishing vistas and say: ‘You see, Lolly, the leaves are beginning to
fall. It was quite time to come home.’
It was useless to protest that autumn begins earlier in London than it
does in the country. That it did so, Laura knew well. That was why she
disliked having to come back; autumn boded her no good, and it was hard
that by a day’s train-journey she should lose almost a month’s reprieve.
Obediently looking along the tarnishing vistas, she knew that once again
she was in for it.
What It was exactly, she would have found hard to say. She sometimes
told herself that it must be the yearly reverberation of those miserable first
months in London when her sorrow for her father’s death was still fresh. No
other winter had been so cold or so long, not even the long cold winters of
the war. Yet now her thoughts of Everard were mellowed and painless, and
she had long ago forgiven her sorrow. Had the coming of autumn quickened
in her only an experienced grief she would not have dreaded it thus, nor felt
so restless and tormented.
Her disquiet had no relevance to her life. It arose out of the ground with
the smell of the dead leaves: it followed her through the darkening streets; it
confronted her in the look of the risen moon. ‘Now! Now!’ it said to her:
and no more. The moon seemed to have torn the leaves from the trees that it
might stare at her more imperiously. Sometimes she tried to account for her
uneasiness by saying that she was growing old, and that the year’s death
reminded her of her own. She compared herself to the ripening acorn that
feels through windless autumnal days and nights the increasing pull of the
earth below. That explanation was very poetical and suitable. But it did not
explain what she felt. She was not wildly anxious either to die or to live;
why, then, should she be rent by this anxiety?
At these times she was subject to a peculiar kind of day-dreaming, so
vivid as to be almost a hallucination: that she was in the country, at dusk,
and alone, and strangely at peace. She did not recall the places which she
had visited in holiday-time, these reproached her like opportunities
neglected. But while her body sat before the first fires and was cosy with
Henry and Caroline, her mind walked by lonely sea-bords, in marshes and
fens, or came at nightfall to the edge of a wood. She never imagined herself
in these places by daylight. She never thought of them as being in any way
beautiful. It was not beauty at all that she wanted, or, depressed though she
was, she would have bought a ticket to somewhere or other upon the
Metropolitan railway and gone out to see the recumbent autumnal graces of
the country-side. Her mind was groping after something that eluded her
experience, a something that was shadowy and menacing, and yet in some
way congenial; a something that lurked in waste places, that was hinted at
by the sound of water gurgling through deep channels and by the voices of
birds of ill-omen. Loneliness, dreariness, aptness for arousing a sense of
fear, a kind of ungodly hallowedness—these were the things that called her
thoughts away from the comfortable fireside.
In this mood she would sometimes go off to explore among the City
churches, or to lose herself in the riverside quarters east of the Pool. She
liked to think of the London of Defoe’s Journal, and to fancy herself back
in the seventeenth century, when, so it seemed to her, there were still
darknesses in men’s minds. Once, hemmed in by the jostling tombstones at
Bunhill Fields, she almost pounced on the clue to her disquiet; and once
again in the goods-yard of the G.W.R., where she had gone to find, not her
own secret, but a case of apples for Caroline.
As time went on Laura grew accustomed to this recurrent autumnal
fever. It was as much a sign of the season as the falling leaves or the first
frost. Before the end of November it was all over and done with. The next
moon had no message for her. Her rambles in the strange places of the mind
were at an end. And if she still went on expeditions to Rotherhithe or the
Jews’ Burying-Ground, she went in search for no more than a little
diversion. Nothing was left but cold and sleet and the knowledge that all
this fuss had been about nothing. She fortified herself against the
dismalness of this reaction by various small self-indulgences. Out of these
she had contrived for herself a sort of mental fur coat. Roasted chestnuts
could be bought and taken home for bedroom eating. Second-hand book-
shops were never so enticing; and the combination of east winds and
London water made it allowable to experiment in the most expensive soaps.
Coming back from her expeditions, westward from the city with the sunset
in her eyes, or eastward from a waning Kew, she would pause for a
sumptuous and furtive tea, eating marrons glacés with a silver fork in the
reflecting warm glitter of a smart pastry-cook’s. These things were exciting
enough to be pleasurable, for she kept them secret. Henry and Caroline
would scarcely have minded if they had known. They were quite indifferent
as to where and how she spent her afternoons; they felt no need to question
her, since they could be sure that she would do nothing unsuitable or
extravagant. Laura’s expeditions were secret because no one asked her
where she had been. Had they asked, she must have answered. But she did
not examine too closely into this; she liked to think of them as secret.
One manifestation of the fur-coat policy, however, could not be kept
from their knowledge, and that manifestation slightly qualified their trust
that Laura would do nothing unsuitable or extravagant.
Except for a gradual increment of Christmas and birthday presents,
Laura’s room had altered little since the day it ceased to be the small spare-
room and became hers. But every winter it blossomed with an unseasonable
luxury of flowers, profusely, shameless as a greenhouse.
‘Why, Lolly! Lilies at this time of year!’ Caroline would say, not
reproachfully, but still with a consciousness that in the drawing-room there
were dahlias, and in the dining-room a fern, and in her own sitting-room,
where she did the accounts, neither ferns nor flowers. Then Laura would
thrust the lilies into her hands; and she would take them to show that she
had not spoken with ill-will. Besides, Lolly would really see more of them
if they were in the drawing-room. And the next day she would meet Laura
on the stairs carrying azaleas. On one occasion even Henry had noticed the
splendour of the lilies: red lilies, angular, authoritative in form and colour
like cardinal’s hats.
‘Where do these come from?’ Caroline had asked, knowing well that
nothing so costly in appearance could come from her florist.
 ‘From Africa,’ Laura had answered, pressing the firm, wet stalks into her
hand.
‘Oh well, I daresay they are quite common flowers there,’ said Caroline
to herself, trying to gloss over the slight awkwardness of accepting a trifle
so needlessly splendid.
Henry had also asked where they came from.
‘From Anthos, I believe,’ said Caroline.
‘Ah!’ said Henry, and roused the coins in his trousers pocket.
‘It’s rather naughty of Lolly. Would you like me just to hint to her that
she mustn’t be quite so reckless?’
‘No. Better not. No need for her to worry about such things.’
Husband and wife exchanged a glance of compassionate understanding.
It was better not. Much better that Lolly should not be worried about money
matters. She was safe in their hands. They could look after Lolly. Henry
was like a wall, and Caroline’s breasts were like towers.
They condoned this extravagance, yet they mistrusted it. Time justified
them in their mistrust. Like many stupid people, they possessed acute
instincts. ‘He that is unfaithful in little things ...’ Caroline would say when
the children forgot to wind up their watches. Their instinct told them that
the same truth applies to extravagance in little things. They were wiser than
they knew. When Laura’s extravagance in great things came it staggered
them so completely that they forgot how judiciously they had suspected it
beforehand.
It befell in the winter of 1921. The war was safely over, so was their
silver wedding, so was Marion’s first confinement. Titus was in his third
year at Oxford, Sibyl was at last going grey, Henry might be made a judge
at any moment. The Trade Returns and the Stock Exchange were not all that
they should be, and there was always the influenza. But Henry was doing
well enough to be lenient to his investments, and Aunt Lucilla and her
fortune had been mercifully released. In the coming spring Caroline
proposed to have the house thoroughly done up. The lesser renovations she
was getting over beforehand, and that was why Laura had gone out before
the shops shut to show Mr. Bunting a pair of massy candlesticks and to
inquire how much he would charge for re-plating them. His estimate was
high, too high to be accepted upon her own responsibility. She decided to
carry the candlesticks back and consult Caroline.
 Mr. Bunting lived in the Earls Court Road, rather a long way off for such
a family friend. But she had plenty of time for walking back, and for
diversion she thought she would take a circuitous route, including the two
foxes who guard the forsaken approach in Holland Park and the lane beside
the Bayswater Synagogue. It was in Moscow Road that she began to be
extravagant. But when she walked into the little shop she had no particular
intention of extravagance, for Caroline’s parcel hung remindingly upon her
arm, and the shop itself, half florist and half greengrocer, had a simple
appearance.
There were several other customers, and while she stood waiting to be
served she looked about her. The aspect of the shop pleased her greatly. It
was small and homely. Fruit and flowers and vegetables were crowded
together in countrified disorder. On the sloping shelf in the window, among
apples and rough-skinned cooking pears and trays of walnuts, chestnuts,
and filberts, was a basket of eggs, smooth and brown, like some larger kind
of nut. At one side of the room was a wooden staging. On this stood jars of
home-made jam and bottled fruits. It was as though the remnants of summer
had come into the little shop for shelter. On the floor lay a heap of earthy
turnips.
Laura looked at the bottled fruits, the sliced pears in syrup, the glistening
red plums, the greengages. She thought of the woman who had filled those
jars and fastened on the bladders. Perhaps the greengrocer’s mother lived in
the country. A solitary old woman picking fruit in a darkening orchard,
rubbing her rough fingertips over the smooth-skinned plums, a lean wiry
old woman, standing with upstretched arms among her fruit trees as though
she were a tree herself, growing out of the long grass, with arms stretched
up like branches. It grew darker and darker; still she worked on,
methodically stripping the quivering taut boughs one after the other.
As Laura stood waiting she felt a great longing. It weighed upon her like
the load of ripened fruit upon a tree. She forgot the shop, the other
customers, her own errand. She forgot the winter air outside, the people
going by on the wet pavements. She forgot that she was in London, she
forgot the whole of her London life. She seemed to be standing alone in a
darkening orchard, her feet in the grass, her arms stretched up to the pattern
of leaves and fruit, her fingers seeking the rounded ovals of the fruit among
the pointed ovals of the leaves. The air about her was cool and moist. There
was no sound, for the birds had left off singing and the owls had not yet
begun to hoot. No sound, except sometimes the soft thud of a ripe plum
falling into the grass, to lie there a compact shadow among shadows. The
back of her neck ached a little with the strain of holding up her arms. Her
fingers searched among the leaves.
She started as the man of the shop came up to her and asked her what
she wished for. Her eyes blinked, she looked with surprise at the gloves
upon her hands.
‘I want one of those large chrysanthemums,’ she said, and turned
towards the window where they stood in a brown jar. There were the apples
and pears, the eggs, the disordered nuts overflowing from their
compartments. There on the floor were the earthy turnips, and close at hand
were the jams and bottled fruits. If she was behaving foolishly, if she looked
like a woman roused out of a fond dream, these were kindly things to
waken to. The man of the shop also had a kind face. He wore a gardener’s
apron, and his hands were brown and dry as if he had been handling earth.
‘Which one would you like, ma’am?’ he asked, turning the bunch of
chrysanthemums about that she might choose for herself. She looked at the
large mop-headed blossoms. Their curled petals were deep garnet colour
within and tawny yellow without. As the light fell on their sleek flesh the
garnet colour glowed, the tawny yellow paled as if it were thinly washed
with silver. She longed for the moment when she might stroke her hand
over those mop heads.
‘I think I will take them all,’ she said.
‘They’re lovely blooms,’ said the man.
He was pleased. He did not expect such a good customer at this late
hour.
When he brought her the change from her pound-note and the
chrysanthemums pinned up in sheets of white paper, he brought also several
sprays of beech leaves. These, he explained, were thrown in with her
purchase. Laura took them into her arms. The great fans of orange tracery
seemed to her even more beautiful than the chrysanthemums, for they had
been given to her, they were a surprise. She sniffed. They smelt of woods,
of dark rustling woods like the wood to whose edge she came so often in
the country of her autumn imagination. She stood very still to make quite
sure of her sensations. Then: ‘Where do they come from?’ she asked.
 ‘From near Chenies, ma’am, in Buckinghamshire. I have a sister living
there, and every Sunday I go out to see her, and bring back a load of foliage
with me.’
There was no need to ask now who made the jams and tied on the
bladders. Laura knew all that she wanted to know. Her course lay clear
before her. Holding the sprays of beech as though she were marching on
Dunsinane, she went to a bookseller’s. There she bought a small guide-book
to the Chilterns and inquired for a map of that district. It must, she
explained, be very detailed, and give as many names and footpaths as
possible. Her eyes were so bright and her demands so earnest that the
bookseller, though he had not that kind of map, was sympathetic, and
directed her to another shop where she could find what she wanted. It was
only a little way off, but closing-time was at hand, so she took a taxi.
Having bought the map she took another taxi home. But at the top of
Apsley Terrace she had one of her impulses of secrecy and told the driver
that she would walk the rest of the way.
There was rather a narrow squeak in the hall, for Caroline’s parcel
became entangled in the gong stand, and she heard Henry coming up from
the wine cellar. If she alarmed the gong Henry would quicken his steps. She
had no time to waste on Henry just then for she had a great deal to think of
before dinner. She ran up to her room, arranged the chrysanthemums and
the beech leaves, and began to read the guide-book. It was just what she
wanted, for it was extremely plain and unperturbed. Beginning as early as
possible with Geology, it passed to Flora and Fauna, Watersheds,
Ecclesiastical Foundations and Local Government. After that came a list of
all the towns and villages, shortly described in alphabetical order. Lamb’s
End had three hundred inhabitants and a perpendicular font. At Walpole St.
Dennis was the country seat of the Bartlet family, faced with stucco and
situated upon an eminence. The almshouses at Semple, built in 1703 by
Bethia Hood, had a fine pair of wrought-iron gates. It was dark as she
pressed her nose against the scrolls and rivets. Bats flickered in the little
courtyard, and shadows moved across the yellow blinds. Had she been born
a deserving widow, life would have been simplified.
She wasted no time over this regret, for now at last she was simplifying
life for herself. She unfolded the map. The woods were coloured green and
the main roads red. There was a great deal of green. She looked at the beech
leaves. As she looked a leaf detached itself and fell slowly. She remembered
squirrels.
The stairs creaked under the tread of Dunlop with the hot-water can.
Dunlop entered, glancing neither at Laura curled askew on the bed nor at
the chrysanthemums ennobling the dressing-table. She was a perfectly
trained servant. Before she left the room she took a deep breath, stooped
down, and picked up the beech leaf.
Quarter of an hour afterwards Laura exclaimed: ‘Oh! a windmill!’ She
took up the guide-book again, and began to read intently.
She was roused by an unaccustomed clash of affable voices in the hall.
She remembered, leapt off the bed, and dressed rapidly for the family
dinner-party. They were all there when she reached the drawing-room. Sibyl
and Titus, Fancy and her Mr. Wolf-Saunders, Marion with the latest news
from Sprat, who, being in the Soudan, could not dine out with his wife.
Sprat had had another boil on his neck, but it had yielded to treatment. ‘Ah,
poor fellow,’ said Henry. He seemed to be saying: ‘The price of Empire.’
During dinner Laura looked at her relations. She felt as though she had
awoken, unchanged, from a twenty-years slumber, to find them almost
unrecognisable. She surveyed them, one after the other. Even Henry and
Caroline, whom she saw every day, were half hidden under their
accumulations—accumulations of prosperity, authority, daily experience.
They were carpeted with experience. No new event could set jarring foot on
them but they would absorb and muffle the impact. If the boiler burst, if a
policeman climbed in at the window waving a sword, Henry and Caroline
would bring the situation to heel by their massive experience of normal
boilers and normal policemen.
She turned her eyes to Sibyl. How strange it was that Sibyl should have
exchanged her former look of a pretty ferret for this refined and waxen
mask. Only when she was silent, though, as now she was, listening to Henry
with her eyes cast down to her empty plate: when she spoke the ferret look
came back. But Sibyl in her house at Hampstead must have spent many
long afternoons in silence, learning this unexpected beauty, preparing her
face for the last look of death. What had been her thoughts? Why was she
so different when she spoke? Which, what, was the real Sibyl: the greedy,
agile little ferret or this memorial urn?
 Fancy’s Mr. Wolf-Saunders had eaten all his bread and was at a loss.
Laura turned to him and asked after her great-nephew, who was just then
determined to be a bus-conductor. ‘He probably will be,’ said his father
gloomily, ‘if things go on as they are at present.’
Great-nephews and great-nieces suggested nephews and nieces.
Resuming her scrutiny of the table she looked at Fancy, Marion, and Titus.
They had grown up as surprisingly as trees since she first knew them, and
yet it did not seem to her that they were so much changed as their elders.
Titus, in particular, was easily recognisable. She caught his eye, and he
smiled back at her, just as he had smiled back when he was a baby. Now he
was long and slim, and his hay-coloured hair was brushed smoothly back
instead of standing up in a crest. But one lock had fallen forward when he
laughed, and hung over his left eye, and this gave him a pleasing, rustic
look. She was glad still to be friends with Titus. He might very usefully abet
her, and though she felt in no need of allies, a little sympathy would do no
harm. Certainly the rustic forelock made Titus look particularly congenial.
And how greedily he was eating that apple, and with what disparagement of
imported fruit he had waved away the Californian plums! It was nice to feel
sure of his understanding and approval, since at this moment he was
looking the greatest Willowes of them all.
Most of the family attention was focussed on Titus that evening. No
sooner had coffee been served than Sibyl began about his career. Had
Caroline ever heard of anything more ridiculous? Titus still declared that he
meant to manage the family brewery. After all his success at Oxford and his
popularity, could anything be more absurd than to bury himself in
Somerset?
His own name was the first thing that Titus heard as he entered the
drawing-room. He greeted it with an approving smile, and sat down by
Laura, carefully crossing his long legs.
‘She spurns at the brewery, and wants me to take a studio in Hampstead
and model bustos,’ he explained.
Titus had a soft voice. His speech was gentle and sedate. He chose his
words with extreme care, but escaped the charge of affectation by
pronouncing them in a hesitating manner.
‘I’m sure sculpture is his métier,’ said Sibyl. ‘Or perhaps poetry.
Anyhow, not brewing. I wish you could have seen that little model he made
of the grocer at Arcachon.’
Marion said: ‘I thought bustos always had wigs.’
‘My dear, you’ve hit it. In fact, that is my objection to this plan for
making me a sculptor. Revive the wig, and I object no more. The head is the
noblest part of man’s anatomy. Therefore enlarge it with a wig.’
Henry thought the conversation was taking a foolish turn. But as host it
was his duty to take part in it.
‘What about the Elgin Marbles?’ he inquired. ‘No wigs there.’
The Peruke and its Functions in Attic Drama, thought Titus, would be a
pretty fancy. But it would not do for his uncle. Agreeably he admitted that
there were no wigs in the Elgin Marbles.
They fell into silence. At an ordinary dinner party Caroline would have
felt this silence to be a token that the dinner party was a failure. But this
was a family affair, there was no disgrace in having nothing to say. They
were all Willoweses and the silence was a seemly Willowes silence. She
could even emphasise it by counting her stitches aloud.
All the chairs and sofas were comfortable. The fire burnt brightly, the
curtains hung in solemn folds; they looked almost as solemn as organ pipes.
Lolly had gone off into one of her day dreams, just her way, she would
never trouble to give a party the least prod. Only Sibyl fidgeted, twisting
her heel about in her satin slipper.
‘What pretty buckles, Sibyl! Have I seen them before?’
Sibyl had bought them second-hand for next to nothing. They came from
Arles, and the old lady who had sold them to her had been such a character.
She repeated the characteristic remarks of the old lady in a very competent
French accent. Her feet were as slim as ever, and she could stretch them out
very prettily. Even in doing so she remembered to ask Caroline where they
were going for the Easter holidays.
‘Oh, to Blythe, I expect,’ said Caroline. ‘We know it.’
‘When I have evicted my tenants and brewed a large butt of family ale, I
shall invite you all down to Lady Place,’ said Titus.
‘But before then,’ said Laura, speaking rather fast, ‘I hope you will all
come to visit me at Great Mop.’
Every one turned to stare at her in bewilderment.
 ‘Of course, it won’t be as comfortable as Lady Place. And I don’t
suppose there will be room for more than one of you at a time. But I’m sure
you’ll think it delightful.’
‘I don’t understand,’ said Caroline. ‘What is this place, Lolly?’
‘Great Mop. It’s not really Great. It’s in the Chilterns.’
‘But why should we go there?’
‘To visit me. I’m going to live there.’
‘Live there? My dear Lolly!’
‘Live there, Aunt Lolly?’
‘This is very sudden. Is there really a place called ...?’
‘Lolly, you are mystifying us.’
They all spoke at once, but Henry spoke loudest, so Laura replied to
him.
‘No, Henry, I’m not mystifying you. Great Mop is a village in the
Chilterns, and I am going to live there, and perhaps keep a donkey. And you
must all come on visits.’
‘I’ve never even heard of the place!’ said Henry conclusively.
‘But you’ll love it. “A secluded hamlet in the heart of the Chilterns,
Great Mop is situated twelve miles from Wickendon in a hilly district with
many beech-woods. The parish church has a fine Norman tower and a
squint. The population is 227.” And quite close by on a hill there is a ruined
windmill, and the nearest railway station is twelve miles off, and there is a
farm called Scramble Through the Hedge....’
Henry thought it time to interrupt. ‘I suppose you don’t expect us to
believe all this.’
‘I know. It does seem almost too good to be true. But it is. I’ve read it in
a guide-book, and seen it on a map.’
‘Well, all I can say is....’
‘Henry! Henry!’ said Caroline warningly. Henry did not say it. He threw
the cushion out of his chair, glared at Laura, and turned away his head.
For some time Titus’s attempts at speech had hovered above the tumult,
like one holy appeasing dove loosed after the other. The last dove was
luckier. It settled on Laura.
 ‘How nice of you to have a donkey. Will it be a grey donkey, like
Madam?’
‘Do you remember dear Madam, then?’
‘Of course I remember dear Madam. I can remember everything that
happened to me when I was four. I rode in one pannier, and you, Marion,
rode in the other. And we went to have tea in Potts’s Dingle.’
‘With sponge cakes and raspberry jam, do you remember?’
‘Yes. And milk surging in a whisky bottle. Will you have thatch or slate,
Aunt Lolly? Slate is very practical.’
‘Thatch is more motherly. Anyhow, I shall have a pump.’
‘Will it be an indoor or an outdoor pump? I ask, for I hope to pump on it
quite often.’
‘You will come to stay with me, won’t you, Titus?’
Laura was a little cast-down. It did not look, just then, as if any one else
wanted to come and stay with her at Great Mop. But Titus was as
sympathetic as she had hoped. They spent the rest of the evening telling
each other how she would live. By half-past ten their conjectures had
become so fantastic that the rest of the family thought the whole scheme
was nothing more than one of Lolly’s odd jokes that nobody was ever
amused by. Henry took heart. He rallied Laura, supposing that when she
lived at Great Mop she would start hunting for catnip again, and become the
village witch.
‘How lovely!’ said Laura.
Henry was satisfied. Obviously Laura could not be in earnest.
When the guests had gone, and Henry had bolted and chained the door,
and put out the hall light, Laura hung about a little, thinking that he or
Caroline might wish to ask her more. But they asked nothing and went
upstairs to bed. Soon after, Laura followed them. As she passed their
bedroom door she heard their voices within, the comfortable fragmentary
talk of a husband and wife with complete confidence in each other and
nothing particular to say.
Laura decided to tackle Henry on the morrow. She observed him during
breakfast and saw with satisfaction that he seemed to be in a particularly
benign mood. He had drunk three cups of coffee, and said ‘Ah! poor
fellow!’ when a wandering cornet-player began to play on the pavement
opposite. Laura took heart from these good omens, and, breakfast being
over, and her brother and the Times retired to the study, she followed them
thither.
‘Henry,’ she said. ‘I have come for a talk with you.’
Henry looked up. ‘Talk away, Lolly,’ he said, and smiled at her.
‘A business talk,’ she continued.
Henry folded the Times and laid it aside. He also (if the expression may
be allowed) folded and laid aside his smile.
‘Now, Lolly, what is it?’
His voice was kind, but business-like. Laura took a deep breath, twisted
the garnet ring round her little finger, and began.
‘It has just occurred to me, Henry, that I am forty-seven.’
She paused.
‘Go on!’ said Henry.
‘And that both the girls are married. I don’t mean that that has just
occurred to me too, but it’s part of it. You know, really I’m not much use to
you now.’
‘My dear Lolly!’ remonstrated her brother ‘You are extremely useful.
Besides, I have never considered our relationship in that light.’
‘So I have been thinking. And I have decided that I should like to go and
live at Great Mop. You know, that place I was talking about last night.’
Henry was silent. His face was completely blank. Should she recall
Great Mop to him by once more repeating the description out of the guide-
book?
‘In the Chilterns,’ she murmured. ‘Pop. 227.’
Henry’s silence was unnerving her.
‘Really, I think it would be a good plan. I should like to live alone in the
country. And in my heart I think I have always meant to, one day. But one
day is so like another, it’s almost impossible to throw salt on its tail. If I
don’t go soon, I never shall. So if you don’t mind, I should like to start as
soon as possible.’
There was another long pause. She could not make out Henry at all. It
was not like him to say nothing when he was annoyed. She had expected
thunders and tramplings, and those she could have weathered. But thus
becalmed under a lowering sky she was beginning to lose her head.
At last he spoke.
‘I hardly know what to say.’
‘I’m sorry if the idea annoys you, Henry.’
‘I am not annoyed. I am grieved. Grieved and astonished. For twenty
years you have lived under my roof. I have always thought—I may be
wrong, but I have always thought—that you were happy here.’
‘Quite happy,’ said Laura.
‘Caroline and I have done all we could to make you so. The children—
all the children—look on you as a second mother. We are all devoted to
you. And now, without a word of warning, you propose to leave us and go
and live at a place called Great Mop. Lolly! I must ask you to put this
ridiculous idea out of your head.’
‘I never expected you to be so upset, Henry. Perhaps I should have told
you more gradually. I should be sorry to hurt you.’
‘You have hurt me, I admit,’ said he, firmly seizing on this advantage.
‘Still, let that pass. Say you won’t leave us, Lolly.’
‘I’m afraid I can’t quite do that.’
‘But Lolly, what you want is absurd.’
‘It’s only my own way, Henry.’
‘If you would like a change, take one by all means. Go away for a
fortnight. Go away for a month! Take a little trip abroad if you like. But
come back to us at the end of it.’
‘No, Henry. I love you all, but I feel I have lived here long enough.’
‘But why? But why? What has come over you?’
Laura shook her head.
‘Surely you must have some reasons.’
‘I have told you my reasons.’
‘Lolly! I cannot allow this. You are my sister. I consider you my charge.
I must ask you, once for all to drop this idea. It is not sensible. Or suitable.’
‘I have reminded you that I am forty-seven. If I am not old enough now
to know what is sensible and suitable, I never shall be.’
‘Apparently not.’
 This was more like Henry’s old form. But though he had scored her off,
it did not seem to have encouraged him as much as scoring off generally
did. He began again, almost as a suppliant.
‘Be guided by me, Lolly. At least, take a few days to think it over.’
‘No, Henry. I don’t feel inclined to; I’d much rather get it over now.
Besides, if you are going to disapprove as violently as this, the sooner I
pack up and start the better.’
‘You are mad. You talk of packing up and starting when you have never
even set eyes on the place.’
‘I was thinking of going there to-day, to make arrangements.’
‘Well, then, you will do nothing of the kind. I’m sorry to seem harsh,
Lolly. But you must put all this out of your mind.’
‘Why?’
‘It is impracticable.’
‘Nothing is impracticable for a single, middle-aged woman with an
income of her own.’
Henry paled slightly, and said: ‘Your income is no longer what it was.’
‘Oh, taxes!’ said Laura contemptuously. ‘Never mind; even if it’s a little
less, I can get along on it.’
‘You know nothing of business, Lolly. I need not enter into explanations
with you. It should be enough for me to say that for the last year your
income has been practically non-existent.’
‘But I can still cash cheques.’
‘I have placed a sum at the bank to your credit.’
Laura had grown rather pale too. Her eyes shone.
‘I’m afraid you must enter into explanations with me, Henry. After all, it
is my income, and I have a right to know what has happened to it.’
‘Your capital has always been in my hands, Lolly, and I have
administered it as I thought fit.’
‘Go on,’ said Laura.
‘In 1920 I transferred the greater part of it to the Ethiopian Development
Syndicate, a perfectly sound investment which will in time be as good as
ever, if not better. Unfortunately, owing to this Government and all this
socialistic talk the soundest investments have been badly hit. The Ethiopian
Development Syndicate is one of them.’
‘Go on, Henry. I have understood quite well so far. You have
administered all my money into something that doesn’t pay. Now explain
why you did this.’
‘I had every reason for thinking that I should be able to sell out at a
profit almost immediately. During November the shares had gone up from
5¾ to 8½. I bought in December at 8½. They went to 8¾ and since then
have steadily sunk. They now stand at 4. Of course, my dear, you needn’t
be alarmed. They will rise again the moment we have a Conservative
Government, and that, thank Heaven, must come soon. But you see at
present it is out of the question for you to think of leaving us.’
‘But don’t these Ethiopians have dividends?’
‘These,’ said Henry with dignity, ‘are not the kind of shares that pay
dividends. They are—that is to say, they were, and of course will be again
—a sound speculative investment. But at present they pay no dividends
worth mentioning. Now, Lolly, don’t become agitated. I assure you that it is
all perfectly all right. But you must give up this idea of the country.
Anyhow, I’m sure you wouldn’t find it suit you. You are rheumatic——’
Laura tried to interpose.
‘—or will be. All the Willoweses are rheumatic. Buckinghamshire is
damp. Those poetical beech-woods make it so. You see, trees draw rain. It
is one of the principles of afforestation. The trees—that is to say, the rain
——’
Laura stamped her foot with impatience. ‘Have done with your trumpery
red herrings!’ she cried.
She had never lost her temper like this before. It was a glorious
sensation.
‘Henry!’ She could feel her voice crackle round his ears. ‘You say you
bought those shares at eight and something, and that they are now four. So
if you sell out now you will get rather less than half what you gave for
them.’
‘Yes,’ said Henry. Surely if Lolly were business woman enough to grasp
that so clearly, she would in time see reason on other matters.
‘Very well. You will sell them immediately——’
 ‘Lolly!’
‘—and reinvest the money in something quite unspeculative and
unsound, like War Loan, that will pay a proper dividend. I shall still have
enough to manage on. I shan’t be as comfortable as I thought I should be. I
shan’t be able to afford the little house that I hoped for, nor the donkey. But
I shan’t mind much. It will matter very little to me when I’m there.’
She stopped. She had forgotten Henry, and the unpleasant things she
meant to say to him. She had come to the edge of the wood, and felt its cool
breath in her face. It did not matter about the donkey, nor the house, nor the
darkening orchard even. If she were not to pick fruit from her own trees,
there were common herbs and berries in plenty for her, growing wherever
she chose to wander. It is best as one grows older to strip oneself of
possessions, to shed oneself downward like a tree, to be almost wholly earth
before one dies.
As she left the room she turned and looked at Henry. Such was her
mood, she could have blessed him solemnly, as before an eternal departure.
But he was sitting with his back to her, and did not look round. When she
had gone he took out his handkerchief and wiped his forehead.
Ten days later Laura arrived at Great Mop. After the interview with
Henry she encountered no more opposition. Caroline knew better than to
persist against an obstinacy which had worsted her husband, and the other
members of the family, their surprise being evaporated, were indifferent.
Titus was a little taken aback when he found that his aunt’s romantic
proposals were seriously intended. He for his part was going to Corsica. ‘A
banal mountainous spot,’ he said politely, ‘compared with
Buckinghamshire.’
The day of Laura’s arrival was wet and blusterous. She drove in a car
from Wickendon. The car lurched and rattled, and the wind slapped the rain
against the windows; Laura could scarcely see the rising undulations of the
landscape. When the car drew up before her new home, she stood for a
moment looking up the village street, but the prospect was intercepted by
the umbrella under which Mrs. Leak hastened to conduct her to the porch.
So had it rained, and so had the wind blown, on the day when she had come
on her visit of inspection and had taken rooms in Mrs. Leak’s cottage. So,
Henry and Caroline and their friends had assured her, did it rain and blow
all through the winter in the Chilterns. No words of theirs, they said, could
describe how dismal and bleak it would be among those unsheltered hills.
To Laura, sitting by the fire in her parlour, the sound of wind and rain was
pleasant. ‘Weather like this,’ she thought, ‘would never be allowed in
London.’
The unchastened gusts that banged against the side of the house and
drove the smoke down the chimney, and the riotous gurgling of the rain in
the gutters were congenial to her spirit. ‘Hoo! You daredevil,’ said the wind.
‘Have you come out to join us?’ Yet sitting there with no companionship
except those exciting voices she was quiet and happy.
Mrs. Leak’s tea was strong Indian tea. The bread-and-butter was cut in
thick slices, and underneath it was a crocheted mat; there was plum jam in a
heart-shaped glass dish, and a plate of rather heavy jam-puffs. It was not
quite so good as the farmhouse teas she remembered in Somerset, but a
great deal better than teas at Apsley Terrace.
Tea being done with, Laura took stock of her new domain. The parlour
was furnished with a large mahogany table, four horsehair chairs and a
horsehair sofa, an armchair, and a sideboard, rather gimcrack compared to
the rest of the furniture. On the walls, which were painted green, hung a
print of the Empress Josephine and two rather scowling classical landscapes
with ruined temples, and volcanoes. On either side of the hearth were
cupboards, and the fireplace was of a cottage pattern with hobs, and a small
oven on one side. This fireplace had caught Laura’s fancy when she first
looked at the rooms. She had stipulated with Mrs. Leak that, should she so
wish, she might cook on it. There are some things—mushrooms, for
instance, or toasted cheese—which can only be satisfactorily cooked by the
eater. Mrs. Leak had made no difficulties. She was an oldish woman,
sparing of her words and moderate in her demands. Her husband worked at
the sawmill. They were childless. She had never let lodgings before, but till
last year an aunt with means of her own had occupied the parlour and
bedroom which were now Laura’s.
It did not take Laura very long to arrange her belongings, for she had
brought little. Soon after supper, which consisted of rabbit, bread and
cheese, and table beer, she went upstairs to bed. Moving about her small
cold bedroom she suddenly noticed that the wind had fallen, and that it was
no longer raining. She pushed aside a corner of the blind and opened the
window. The night air was cold and sweet, and the full moon shone high
overhead. The sky was cloudless, lovely, and serene; a few stars glistened
there like drops of water about to fall. For the first time she was looking at
the intricate landscape of rounded hills and scooped valleys which she had
chosen for learning by heart.
Dark and compact, the beech-woods lay upon the hills. Alighting as
noiselessly as an owl, a white cat sprang up on to the garden fence. It
glanced from side to side, ran for a yard or two along the top of the fence
and jumped off again, going secretly on its way. Laura sighed for happiness.
She had no thoughts; her mind was swept as clean and empty as the
heavens. For a long time she continued to lean out of the window, forgetting
where she was and how she had come there, so unearthly was her
contentment.
Nevertheless her first days at Great Mop gave her little real pleasure. She
wrecked them by her excitement. Every morning immediately after
breakfast she set out to explore the country. She believed that by eating a
large breakfast she could do without lunch. The days were short, and she
wanted to make the most of them, and making the most of the days and
going back for lunch did not seem to her to be compatible. Unfortunately,
she was not used to making large breakfasts, so her enthusiasm was
qualified by indigestion until about four P.M., when both enthusiasm and
indigestion yielded to a faintish feeling. Then she turned back, generally by
road, since it was growing too dark to find out footpaths, and arrived home
with a limp between six and seven. She knew in her heart that she was not
really enjoying this sort of thing, but the habit of useless activity was too
strong to be snapped by change of scene. And in the evening, as she looked
at the map and marked where she had been with little bleeding footsteps of
red ink, she was enchanted afresh by the names and the bridle-paths, and,
forgetting the blistered heel and the dissatisfaction of that day’s walk,
planned a new walk for the morrow.
Nearly a week had gone by before she righted herself. She had made an
appointment with the sunset that she should see it from the top of a certain
hill. The hill was steep, and the road turned and twisted about its sides. It
was clear that the sunset would be at their meeting-place before she was,
nor would it be likely to kick its heels and wait about for her. She looked at
the sky and walked faster. The road took a new and unsuspected turn,
concealed behind the clump of trees by which she had been measuring her
progress up the hill. She was growing more and more flustered, and at this
prick she lost her temper entirely. She was tired, she was miles from Great
Mop, and she had made a fool of herself. An abrupt beam of light shot up
from behind the hedge as though the sun in vanishing below the horizon
had winked at her. ‘This sort of thing,’ she said aloud, ‘has got to be put a
stop to.’ She sat down in the extremely comfortable ditch to think.
The shades that had dogged her steps up the hill closed in upon her as
she sat in the ditch, but when she took out her map there was enough light
to enable her to see where the nearest inn lay. It was close at hand; when
she got there she could just read its name on the sign. Its name was The
Reason Why. Entering The Reason Why, she ordered tea and a conveyance
to drive her back to Great Mop. When she left the inn it was a brilliant night
of stars. Outside stood a wagonette drawn by a large white horse. Piled on
the seat of the wagonette were a number of waterproof rugs with finger-
rings on them, and these she wrapped round her with elaborate care.
The drive back to Great Mop was more filled with glory than anything
she had ever experienced. The wagonette creaked over bare hill-tops and
plunged downwards into the chequered darknesses of unknown winter
woods. All the stars shook their glittering spears overhead. Turning this way
and that to look at them, the frost pinched her cheeks.
That evening she asked Mrs. Leak if she would lend her some books.
From Mrs. Leak’s library she chose Mehalah, by the Rev. Sabine Baring-
Gould, and an anonymous work of information called Enquire Within Upon
Everything. The next morning was fine and sunny. She spent it by the
parlour fire, reading. When she read bits of Mehalah she thought how
romantic it would be to live in the Essex Marshes. From Enquire Within
Upon Everything she learned how gentlemen’s hats if plunged in a bath of
logwood will come out with a dash of respectability, and that ruins are best
constructed of cork. During the afternoon she learned other valuable facts
like these, and fell asleep. On the following morning she fell asleep again,
in a beech-wood, curled up in a heap of dead leaves. After that she had no
more trouble. Life becomes simple if one does nothing about it. Laura did
nothing about anything for days and days till Mrs. Leak said: ‘We shall
soon be having Christmas, miss.’
Christmas! So it had caught them all again. By now the provident
Caroline herself was suffering the eleventh hour in Oxford Street. But here
even Christmas was made easy.