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The Readable Darwin
The Readable Darwin
The Origin of Species Edited for Modern
Readers
Second Edition

Jan A. Pechenik
 Oxford University Press is a department of the University of Oxford. It furthers
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Library of Congress Cataloging-in-Publication Data

Names: Darwin, Charles, 1809–1882, author. | Pechenik, Jan A., editor.
Title: The readable Darwin : the origin of species /
edited for modern readers by Jan Pechenik.
Other titles: On the origin of species
Description: New York, NY : Oxford University Press, [2023] | Includes index.
Identifiers: LCCN 2022040692 (print) | LCCN 2022040693 (ebook) |
ISBN 9780197575260 (hardback) | ISBN 9780197575291 (epub)
Subjects: LCSH: Evolution (Biology) | Natural selection.
Classification: LCC QH 365 . O 25 P 4 34 2023 (print) |
LCC QH 365 . O 25 (ebook) | DDC 576.8/2—dc23/eng/20220920
LC record available at https://lccn.loc.gov/2022040692
LC ebook record available at https://lccn.loc.gov/2022040693

DOI: 10.1093/oso/9780197575260.001.0001

1 3 5 7 9 8 6 4 2
Printed by Sheridan Books, Inc., United States of America
Contents

Preface by Jan A. Pechenik ix

Acknowledgments xv

Introduction 1
1. Variation Under Domestication 5
Variability 5
Effects of Habit and of the Use or Disuse of Parts; Correlated Variation;
Inheritance 5
Character of Domestic Varieties; Difficulties of Distinguishing Between
Varieties and Species; Origin of Domestic Varieties from One or More
Species 7
Breeds of the Domestic Pigeon, Their Differences and Origin 9
Principles of Selection Anciently Followed, and Their Effects 18
Unconscious Selection 21
Circumstances Favorable to Man’s Power of Selection 25
Summary 27
2. Variation in Nature 30
Individual Differences 31
Questionable Species 32
Wide-​Ranging, Much Diffused, and Common Species Vary the Most 37
Species in Larger Genera Vary More Frequently Than Those in Smaller
Genera 38
Many Species Included in the Larger Genera Resemble Varieties: They Are
Closely Related and Have Restricted Ranges 40
Summary 41
3. The Struggle for Existence 43
The Term “Struggle for Existence” Used in a Larger Sense 45
Exponential Rates of Increase 46
Nature of the Checks to Population Growth Increase 49
Complex Relations of All Animals and Plants to Each Other in the
Struggle for Existence 52
The Struggle for Life Is Generally Most Severe Between Individuals and
Varieties of the Same Species 57
4. Natural Selection, or the Survival of the Fittest 62
Sexual Selection 70
Examples of Natural Selection, or the Survival of the Fittest, in Action 73
vi Contents

On the Outbreeding of Individuals 79

Circumstances Favoring the Production of New Forms Through
Natural Selection 82
Extinction Caused by Natural Selection 87
Divergence of Character 88
Effects of Natural Selection on the Descendants of a Common
Ancestor, Through Divergence of Character and Extinction 92
On the Degree to Which Organisms Tend to Advance in Complexity 100
Convergence of Character 104
Summary 106
5. Laws of Variation 110
Effects of Increased Use and Disuse of Parts as Controlled by Natural
Selection 111
Acclimatization 117
Correlated Variation 119
Compensation and the Economy of Growth 122
Multiple, Rudimentary, and Lowly Organized Structures Are
Especially Variable 124
A Part Developed in Any Species to an Extraordinary Degree
Tends to Be Highly Variable, in Comparison with the Same
Part in Related (Allied) Species 124
Specific Characters Vary More Than Generic Characters 128
Secondary Sexual Characteristics Are Highly Variable 129
Distinct Species Present Analogous Variations, So That a Variety of
One Species Often Presents a Trait Typical of a Related Species,
or Reverts Back to Some Trait Possessed by an Early Ancestor 131
Summary 138
6. Difficulties with the Theory 142
On the Absence or Rarity of Transitional Varieties 143
On the Origin and Transitions of Organic Beings with Peculiar
Habits and Structure 147
Organs of Extreme Perfection and Complication 158
Modes of Transition 161
Special Difficulties of the Theory of Natural Selection 166
Organs of Little Apparent Importance, as Affected by Natural Selection 174
Utilitarian Doctrine, How Far True? Beauty, How to Explain It? 177
Summary: The Law of Unity of Type and of the Conditions of Existence
Embraced by the Theory of Natural Selection 184
7. Miscellaneous Objections to the Theory of Natural Selection 189
Modifications Not Necessarily Simultaneous 190
Modifications That Are Apparently of No Direct Service 191
Supposed Incompetence of Natural Selection to Account for
the Incipient Stages of Useful Structures 195
 Contents vii

Widely Different Organs in Members of the Same Class,

Developed from One and the Same Source 213
Summary 221
Reasons for Disbelieving in Great and Abrupt Modifications 223
8. Instinct 229
Inherited Changes of Habit or Instinct in Domesticated Animals 233
Special Instincts 236
Objections to the Theory of Natural Selection as Applied to Instincts:
Neuter and Sterile Insects 251
Summary 259
9. Hybridism 262
Degrees of Sterility 263
Laws Governing the Sterility of First Crosses and Hybrids 270
The Origin and Causes of the Sterility of First Crosses and Hybrids 276
Reciprocal Dimorphism and Trimorphism 278
The Fertility of Varieties When Crossed, and of Their Mongrel
Offspring, Is Not Universal 282
Hybrids and Mongrels Compared, Independently of Their Fertility 285
Summary 288
10. On the Imperfection of the Geological Record 290
On the Passage of Time, as Inferred from the Rate of Deposition and
Extent of Denudation of Sediments 293
On the Poorness of Paleontological Collections 297
On the Absence of Numerous Intermediate Varieties in Any Single
Formation 303
On the Sudden Appearance of Whole Groups of Allied Species 310
On the Sudden Appearance of Groups of Allied Species in the Lowest
Known Fossiliferous Strata 313
11. On the Geological Succession of Organic Beings 320
On Extinction 324
On the Forms of Life Changing Almost Simultaneously
Throughout the World 328
On the Affinities of Extinct Species to Each Other, and to Living Forms 332
On the State of Development of Ancient Compared with Living Forms 337
On the Succession of the Same Types Within the Same Areas,
During the Later Tertiary Periods 340
Summary of the Preceding and Present Chapters 342
12. Geographical Distribution 346
Single Centers of Supposed Creation 351
Means of Dispersal 353
Dispersal During the Glacial Period 360
Alternate Glacial Periods in the North and South 365
viii Contents

13. Geographical Distribution, Continued 373

Freshwater Plant and Animal Productions 373
On the Inhabitants of Oceanic Islands 377
Absence of Batrachians and Terrestrial Mammals on Oceanic Islands 380
On the Relations of the Inhabitants of Islands to Those of the Nearest
Mainland 383
Summary of Chapters 12 and 13 388
14. Evidence for Mutual Affinities Among Organic Beings:
Evidence from Morphology, Embryology, and
Rudimentary Organs 392
Analogical Resemblances 402
On the Nature of the Affinities Connecting Organic Beings 407
Morphology 412
Development and Embryology 416
Rudimentary, Atrophied, and Aborted Organs 426
Summary 431
15. Recapitulation and Conclusion 434

Appendix A: Other Books by Charles Darwin 457

Appendix B: People Referred to in These Chapters 461
Illustration Credits 471
Index 477
Preface by Jan A. Pechenik

You care for nothing but shooting, dogs, and rat-​catching, and you will be
a disgrace to yourself and all your family.
—​From Charles and Emma (2009, by Deborah Heiligman)

Charles Darwin received this rant from his father (Robert Darwin, a successful med-
ical doctor and investor) as a teenager while studying medicine at the university in
Edinburgh, in Scotland. He had just decided to abandon a career in medicine after
seeing several people being operated on without anesthesia.
The father’s outburst is perhaps understandable, but how wrong he was! Here are
some comments about the importance of Charles Darwin’s first edition of The Origin
of Species:

It’s safe to say that The Origin of Species . . . is one of the most influential books ever
written. (David Quammen, The Reluctant Mr. Darwin: An Intimate Portrait of Charles
Darwin and the Making of His Theory of Evolution; 2006, W. W. Norton & Company)

[The Origin] is a book that makes the whole world vibrate. (Adam Gopnik, Angels
and Ages: Lincoln, Darwin, and the Birth of the Modern Age; 2009, Vintage Press)

The Darwinian revolution is widely considered the most important event in the en-
tire history of the human intellect. (Michael Ghiselin, 2008)

Charles Darwin published The Origin of Species in 1859, one year after he and
Alfred Russel Wallace had their remarkably similar papers on natural selection
presented at a meeting of a major scientific group, the Linnean Society, in London.
I think it’s fair to say that the world hasn’t been the same since. Here is the crux of his
argument:

• Organisms vary in a great many characteristics (traits), both anatomical and

behavioral.
• These characteristics are typically passed on to offspring—​they are inherited.
• Organisms compete for space, food, and mates and are subject to predation, so
that there is a constant struggle to stay alive and successfully reproduce.
• Those pressures give an advantage to individuals with certain characteristics.
• Individuals with those beneficial traits are most likely to survive and leave off-
spring, which will then inherit those useful traits.
x Preface

• Over very long periods of time, those characteristics will come to be found in a
great many individuals, and a new species will be thereby created.
• Diversification and extinction will then, over incredibly long periods of time,
create species that differ more and more from each other, creating new classes,
families, genera, and even new phyla.

It’s a brilliant argument, and the argument is brilliantly and exhaustively made.
The first printing of The Origin sold out on the first day. Remarkably though, the
book is rarely read today, not even in high school or college biology courses. That’s a
shame. Not only is it the basis for so much of our modern biological research in so
many areas, but it’s also a wonderful reminder of the incredible diversity of life on
this planet and a wonderful example of a fair and honest argument based on evidence
and logical thinking. Darwin explains his supporting evidence thoroughly but is also
very up front about the things he doesn’t yet understand, even those that might pose
problems for his theory.
Reading Darwin’s book is a bit of a slog though. The style of writing is very much
of the 1800s, and although some of his prose was delightfully memorable, many of his
paragraphs are long and unwieldy, as are many of the sentences, making the reading
rather difficult for many modern readers. He also refers to many people without ever
saying who they are, and he mentions the names of a great many animals and plants
that most people today are not familiar with.
To make it easier for teachers and students to read this wonderful and incredibly
important book, and to make it more accessible as well for interested adults, I have
edited the entire book—​all 15 chapters—​into more readable prose. In doing so, I have
tried to keep as much of Darwin’s original wording as possible and to preserve all of
the sense of what he is saying.
I chose to work with Darwin’s final edition (the Sixth), which was published in
1872. The first edition was published in 1859, and it was revised as new information
was collected (by Darwin and by many others around the world) and as new argu-
ments were advanced against Darwin’s proposition. In the sixth edition we can see
how Darwin responded to all of his critics. For the sixth edition, Darwin also added
an entirely new and wonderful chapter (Chapter 7), which includes fascinating infor-
mation on the evolution of baleen whales from toothed ancestors, the evolution of
climbing in plants, and the evolution of breasts in mammals. And it was in the sixth
edition of The Origin that Darwin introduced the word “evolution” (he had first used
it in The Descent of Man, published the previous year).
One disadvantage of reading the sixth edition rather than the first, however, is that
because Darwin was under increasing pressure to develop a complete theory of nat-
ural selection, he did more grasping at straws than he did in the first edition in dealing
with the possible causes of variability among individuals and the inheritance of those
variations by offspring. Even by the time of Darwin’s death, in 1882, nobody knew
what caused variation among individuals or how those variations were transmitted
to offspring. Mendel published what eventually turned out to be the beginnings of
 Preface xi

modern genetics in 1866, but the paper was cited only three times in the next 35 years.
It was published in an obscure journal, and those who read the paper seem not to have
understood its implications, particularly that the findings applied to all sorts of traits,
not just the especially distinctive traits that Mendel had been working with in his pea
plant studies. Intriguingly, Mendel read a German translation of Darwin’s work, but,
as far as we know, he never tried to contact Darwin, even when he visited England in
1862. Apparently Mendel himself didn’t see the connection between Darwin’s ideas
and his own work. In any event, it was many decades later that the implications of
Mendel’s work were realized and several decades after that that the basic mechanisms
of inheritance, including the role of DNA, were finally worked out and the mech-
anism of evolution through the process of natural selection, as Darwin had proposed,
was finally well accepted. Indeed, some of that work is still ongoing.
Darwin didn’t get everything right. I have included a number of footnotes to indi-
cate where Darwin was correct and where he was wrong. But his ideas about the role
of variation in shaping morphological and behavioral diversity through natural selec-
tion were on target, as were his ideas about the role of natural selection in gradually
creating new species and new larger taxonomic categories. As a result, natural selec-
tion is now widely accepted as the major explanation for present and past organismal
diversity and for the appearance of major new groups of organisms over time.

My Edits
I have spent much time during the past 10 years or so editing all 15 of Darwin’s
chapters, making multiple passes over every paragraph. But this is not my book: it’s
Darwin’s, and I have tried to maintain his voice as much as possible. I have based
much of my editing on the various rules from my Short Guide to Writing About Biology
(ninth edition). Indeed, I like to think that this is how Darwin would have written The
Origin had he read my Short Guide first! Here are the main rules that I have followed
while editing this material, along with some examples:

1. Omit unnecessary words, especially unnecessary prepositions.

Original: I suspect that the chief use of the nutriment in the seed is to favor the
growth of the young seedling. . . . (Chapter 3)
My revision: I suspect that the seed’s nutrients are chiefly used to promote the
growth of the young seedling. . . .
2. Eliminate weak verbs (what I call “wimpy verb syndrome”), in part by making
organisms the agent of the action.
Original: Many plants are known which regularly produce. . . .
My revision: Many plants regularly produce. . . .
3. Warn readers of what lies ahead, using words like “For example,” “Similarly,” and
“On the other hand,” and remind readers from time to time of what they have
just read.
xii Preface

4. Incorporate definitions into sentences.

Original: One of the most serious is that of neuter insects, which are often
differently constructed from either the males or fertile females (Chapter 6,
p. 174)
My revision: One of the most serious is that of neuter, non-​reproductive
insects, which are often differently constructed from either the males or the
fertile females of the species.
5. Use repetition, summary, and appropriate punctuation to link thoughts and im-
prove the flow of ideas: Never make readers back up!
Original: Although many statements may be found in works on natural history
to this effect, I cannot find even one that seems to me of any weight.
My revision: Although many works on natural history claim that some struc-
tures in one species do indeed serve for the exclusive benefit of a different
species, I cannot find even one example that seems to me to hold any weight.
Here is one more example:
Original: Yet it cannot be said that small islands will not support at least some
small mammals, for they occur in many parts of the world on very small is-
lands . . .
My Revision: Yet it cannot be said that small islands will not support at least
some small mammals, for small mammals occur in many parts of the world
on very small islands . . .

I have made a variety of other changes as well. One would-​be reader commented
on Amazon.com that “Another thing which made the book a little harder for me is
that Darwin mentions a lot of people and animals I’ve never heard of.” I now clarify
what the various animals and plants are that Darwin refers to. For example, when
Darwin mentions cirripedes or Balanus, I help readers by including the term “barna-
cles” somewhere in the sentence. I also have added a little more information to some
sentences, to help make them more meaningful to modern readers. For example:

Original: Thus, we can hardly believe that the webbed feet of the upland goose or
of the frigate-​bird are of special use to these birds.
My revision: Thus, we can hardly believe that the webbed feet of the upland goose
of South American grasslands, or of the frigate-​bird, which cannot swim or
even walk well, and which takes most of its food in flight, are of special use to
these birds.

I have also broken up overly long sentences and overly long paragraphs and altered
sentence structure when necessary to make the sentences easier to read.

Original: A trailing palm tree in the Malay Archipelago climbs the loftiest trees by
the aid of exquisitely constructed hook clusters around the ends of its branches,
and this contrivance, no doubt, is of the highest service to the plant; but as we
 Preface xiii

see nearly similar hooks on many trees which are not climbers, and which, as
there is reason to believe from the distribution of the thorn-​bearing species in
Africa and South America, serve as a defense against browsing quadrupeds, so
the spikes on the palm may at first have been developed for this function, and
subsequently have been improved and taken advantage of by the plant, as it un-
derwent further modification and became a climber.
My revision: In my version, that single sentence has now become three sentences.
Original: Seeing how important an organ of locomotion the tail is in most aquatic
animals. . . .
My revision: Seeing how important the tail is as an organ of locomotion in most
aquatic animals. . . .

Finally, I have written short previews for the start of each chapter, have boldfaced
particularly important sentences, and added many illustrations. Darwin’s sixth edi-
tion originally had only a single illustration, a schematic drawing of evolutionary
branching patterns Figure 4.11, in Chapter 4. My version includes numerous draw-
ings and photographs of the various plants and animals that Darwin talks about,
along with some other figures that clarify some of his major points.
Acknowledgments

Preparing this “translation” gave me a wonderful opportunity to read a variety of

books about The Origin and about Darwin’s life. In particular, Janet Browne’s Charles
Darwin: A Biography, Volume 1: Voyaging (1996, Princeton University Press) and
Volume 2: The Power of Place (2003, Princeton University Press); James Costa’s The
Annotated Origin (2009, Harvard University Press) and David Reznick’s The Origin
Then and Now: An Interpretative Guide to the Origin of Species (2010, Princeton
University Press) were invaluable in helping me to fully understand Darwin’s
thinking.
I am very grateful to my geological colleagues Jack Ridge (Tufts University) and
Alan Young (Salem State University) for carefully reading my versions of Darwin’s
Chapters 10 and 11, about difficulties in interpreting the geological record. I also wish
to thank my friend and colleague Gordon Hendler at the Natural History Museum of
Los Angeles County for critical feedback on Chapter 3 and for moral support, advice,
and encouragement throughout the project. My Tufts colleague George Ellmore also
provided helpful information about plant biology.
Thanks to the following reviewers for their critiques of the original book proposal
and sample chapter:

Stevan Arnold, Oregon State University

John Avise, University of California, Irvine
Alex Badyev, University of Arizona
David Begun, University of California, Davis
Douglas Futuyma, The State University of New York at Stony Brook
Richard Harrison, Cornell University
Mark Kirkpatrick, University of Texas at Austin
Carol Lee, University of Wisconsin
Michael Lynch, Indiana University
Mohamed Noor, Duke University
Patrick Phillips, University of Oregon
Adam Porter, University of Massachusetts
David Rand, Brown University
Michael Wade, Indiana University
Bruce Walsh, University of Arizona

I am also grateful to the following biologists who read and commented on com-
plete texts of the first eight chapters of the manuscript:
xvi Acknowledgments

Greg Bole, The University of British Columbia

Becky Fuller, University of Illinois at Urbana-​Champaign
Frid-​a Jóhannesdóttir, Cornell University
Adi Livnat, Virginia Polytechnic Institute and State University
Joel W. McGlothlin, Virginia Polytechnic Institute and State
Benjamin Normark, University of Massachusetts Amherst
Jeremy Searle, Cornell University
Christopher S. Willett, University of North Carolina at Chapel Hill
Pamela Yeh, University of California, Los Angeles
Rebecca Zufall, University of Houston University

It’s also a pleasure to thank Andy Sinauer for his enthusiastic reception of this idea
and his willingness to see it through and his entire team for their professionalism
and expertise in bringing the project to completion so smoothly: Dean Scudder,
Christopher Small, Chelsea Holabird, Stephanie Bonner, David McIntyre, Joan
Gemme, Jefferson Johnson, and Tom Friedmann. I am also grateful to those at Oxford
University Press for seeing this project through to completion with the full volume of
Darwin’s wonderful work, The Origin of Species.
My son Oliver and his wife Ardea Thurston-​Shaine helped me find many appro-
priate photographs, answered many of my questions about birds, and offered helpful
suggestions on various drafts. Ardea also contributed an excellent drawing of var-
ious pigeon breeds for Chapter 1 and developed the concept that resulted in the fab-
ulous image that adorns the cover of this book. One of my graduate students (Casey
Diederich) and several undergraduates (Elizabeth Card and Chinami Michaels) also
contributed wonderful figures for the first part of the project, and another of my grad-
uate students, Daria Clark, helped find appropriate figures for the second part of the
project. And finally I thank my entire family—​especially my wife, Regina—​for their
love, support, and patience throughout the adventure. J. S. Bach, Franz Schubert,
Johannes Brahms, and Carl Nielsen have also played major roles in keeping me sane
throughout the project.
—​Jan A. Pechenik, 2022
Introduction

For nearly five years, from December 27, 1831, until October 2, 1836, I served as nat-
uralist aboard the HMS Beagle, exploring. During that voyage I was much amazed by
how the various types of organisms were distributed around South America, and how
the animals and plants presently living on that continent are related to those found
only as fossils in the geological record elsewhere. These facts, as will be seen in later
chapters, seemed to me to throw some light on the origin of species—​that “mystery of
mysteries,” as it has been called by one of our greatest scientists, John Herschel.1 After
I returned home, it occurred to me, in 1837, that I might be able to help address this
great question by patiently accumulating and reflecting on all sorts of facts that might
have any bearing on it. Finally, after five years of work, I allowed myself to speculate
on the subject and wrote up some brief notes. I enlarged these in 1844, into a sketch
of the conclusions that seemed to be most probable from the evidence I had collected.
Over the subsequent 15 years I have steadily pursued the same object: trying to un-
derstand how new species come about. I hope you will excuse me for entering these
personal details of my work, as I give them only to show that I have not been hasty in
coming to a decision.
Now, in 1859, my work is nearly finished.2 Still, it will take me many more years to
complete it, and as my health is not strong, I have been urged to publish this brief ver-
sion of my findings. I have more especially been urged to do this as Mr. Alfred Russel
Wallace, who is now studying the natural history of the Malay Archipelago north of
Australia, has reached almost exactly the same conclusions that I have reached con-
cerning the origin of species. In 1858, he sent me an account on the subject, requesting
that I pass it along to the geologist Sir Charles Lyell, who in turn sent it for presenta-
tion at a meeting of the Linnean Society. Mr. Wallace’s paper has now been published,
in the third volume of that society’s journal. Sir Lyell and my colleague the excellent
botanist Dr. Joseph Hooker, who both knew of my work—​Dr. Hooker having read my
sketch of it in 1844—​honored me by suggesting that I also publish a summary of my
ideas in that journal (some brief extracts from my own manuscripts) along with Mr.
Wallace’s excellent paper.

1 John Herschel was a chemist, astronomer, mathematician, botanist, inventor, and the son of the bril-

liant astronomer William Herschel.

2 The first edition of The Origin of Species was published in 1859. Subsequent editions were published

in 1860, 1861, 1866, 1869, and 1872, as Darwin collected more information and refined and expanded
his ideas.

The Readable Darwin. Second Edition. Jan A. Pechenik, Oxford University Press. © Oxford University Press 2023.
DOI: 10.1093/​oso/​9780197575260.003.0001
2 Introduction

The brief summary of my ideas that I record here must necessarily be imperfect.
I do not have sufficient space in this volume to give references and authorities for
many of my statements; I must trust the reader to have some confidence in my accu-
racy. No doubt errors will have crept in, though I hope I have always been cautious
in trusting to good authorities alone. I can here give only the general conclusions at
which I have arrived, with a few facts in illustration, which I hope, in most cases, will
suffice. No one can feel more clearly than I do the necessity of eventually publishing
in detail all the facts, with references, on which I base my conclusions, as I am well
aware that some evidence can be found apparently leading to conclusions directly op-
posite to those that I have arrived at, for nearly every point I make. A fair result can be
obtained only by fully stating and balancing the facts and arguments on both sides of
each question, and it is impossible to do so here.
I also much regret that lack of space prevents my having the satisfaction of acknow-
ledging the generous assistance I have received from many naturalists, some of
whom I have never actually met. I cannot, however, let this opportunity pass without
expressing my deep obligations to Dr. Hooker, who, for the past 15 years, has aided me
in every possible way with his large stores of knowledge and his excellent judgment.
In considering the origin of species, it is certainly conceivable that some naturalist,
just by reflecting on the mutual interactions of all living organisms, on their embry-
ological similarities and differences, their geographical distribution, their geological
succession over time, and other such facts, might reach the conclusion that individual
species had not been independently created but had in fact descended from other
species. But such a conclusion, even if well founded and well argued, would be unsat-
isfactory until it could be shown how the innumerable species inhabiting this world
had been modified so as to acquire that perfection of structure and coadaptation
that now justly excites our admiration. What is the mechanism by which this could
come about?
Naturalists continually refer to external conditions, such as climate and food, as
the only possible source of variation. In one limited sense, as I will discuss in detail
later, this may be true; but it is preposterous to attribute to mere external conditions
the structure of any organism—​for instance that of the woodpecker, with its feet, tail,
beak, and tongue so wonderfully adapted to catch insects under the bark of trees.
And consider, too, the case of the mistletoe plant, which draws its nourishment from
particular trees, and which has seeds that must be transported by certain birds, and
which has flowers with separate sexes absolutely requiring the help of certain insects
to bring pollen from one flower to the other. It is equally preposterous to account for
the structure of this parasitic plant, and its intimate and essential relationship with
such a range of other organisms, by the effects of external conditions, or of habit, or of
the wishes and desires of the plant itself.
Thus it is of the highest importance to gain a clear insight into the means through
which organisms become modified and coadapted to interact with other organisms.
When I began my observations it seemed to me that a careful study of domesticated
animals and of cultivated plants would offer me the best chance of resolving this
 Introduction 3

difficult problem. Nor have I been disappointed: in this and in all other perplexing
cases, I have invariably found that our knowledge of variation under domestication,
imperfect as it is, provided the best and most satisfying clues to how the process works
in the wild. I am fully convinced of the value of such studies, even though they have so
far been typically neglected by naturalists.
For this reason I devote the first chapter of this summary of my ideas and find-
ings to Variation under Domestication. We shall see that a large amount of hereditary
modification is at least possible, and perhaps even more importantly, we shall see how
great is our power to accumulate slight variations in traits over time by simply choos-
ing which animals to breed together.
In Chapter 2, I will talk about the variability of species in the wild, although I’ll
not have space here to present the long catalogs of facts that the topic really requires.
I will, however, be able to discuss the circumstances that are most favorable to causing
variation. In the next chapter (Chapter 3), I will consider the struggle for existence
that takes place among all living things throughout the world and show how it is an
inevitable consequence of the exponential rates3 at which the populations of all living
beings tend to grow (see Figure 3.34). This is the basic doctrine of the English cleric
and demographer Robert Malthus, applied now to the whole animal and vegetable
kingdoms: As many more individuals of each species are born than can possibly
survive, and as, consequently, there is a frequently recurring struggle for existence,
it follows that any organism will have a better chance of surviving if it varies even
just slightly in any way that is helpful to itself under the complex and sometimes
varying conditions of life. Thus, that individual will be naturally selected for. As so
many variations are transmitted through inheritance, any selected variety will then
tend to propagate its new and modified form among its offspring.
I will discuss this fundamental topic of “natural selection” in some detail in
Chapter 4 and will show how natural selection almost always causes extinction of
the less capable forms of life and leads to what I have called “divergence of character”
within a population. In the next chapter (Chapter 5), I will discuss the complex and
little-​known laws of variation. In the succeeding three chapters, I will carefully con-
sider the most apparent and gravest difficulties presented by my theory. These include
(1) the difficulty in understanding how a simple organism or a simple organ can even-
tually be changed and perfected into a highly developed being or into an elaborately
constructed organ (Chapters 6 and 7) and (2) the difficulty in understanding how
complex animal behaviors and mental capacity may be shaped by the same forces
that shape morphology (Chapter 8).5 Then, in Chapter 9, I use examples of suc-
cessful crosses between what are considered to be separate species (i.e., hybridism)
to tighten the link between varieties and species, and, in the following two chapters

3 With exponential growth, a population continues to grow by the same percentage each year, which

means that the population size will increase faster and faster over time.
4 Figure 3.3 was not included in Darwin’s The Origin of Species.
5 The Origin of Species, sixth edition, contains 15 chapters; The Readable Darwin covers Chapters 1–​8.
4 Introduction

(Chapters 10 and 11), I discuss the imperfection of the fossil record and how that
imperfection explains why we don’t see all of the intermediate stages of evolutionary
change preserved as fossils. Chapters 12 and 13 explain the fascinating patterns of
how various species are distributed around the world and why, for example, we see
some of the same species currently living in very different, often widely separated
locations. In Chapter 14, I discuss how similarities in particular developmental char-
acteristics and the presence of rudimentary organs in adults provide evidence that all
of today’s living species have descended from ancient, common ancestors. In the final
chapter (Chapter 15), I give a brief recapitulation of the entire subject and a few con-
cluding remarks.
No one should be surprised that there is as yet much that is unexplained regarding
the origin of species and varieties considering how profoundly ignorant we are about
the mutual interrelations of all the animals and plants that live around us. Who can
explain why one species ranges widely and is abundant, while a related species has a
narrow range and is rare? Yet these relationships are of the greatest importance for
they determine the present welfare and, I believe, the future success and modifica-
tion of every inhabitant of this world. We know even less of the mutual relations of
the innumerable inhabitants of the world during the many past geological epochs
in Earth’s history. Although much remains obscure, and will long remain obscure,
I can entertain no doubt, after the most careful study and dispassionate judgment
of which I am capable, that the view which most naturalists entertain, and which
I previously entertained myself—​that each species was independently created—​is
erroneous. I am fully convinced that species are not unchangeable, but rather that all
those species that now belong to what are called the same “genera” are direct descen-
dants of some other species, a species that is probably now extinct. In the same way,
the acknowledged varieties of any one species are the direct descendants of that spe-
cies. Furthermore, I am convinced that natural selection has been the main engine,
although not the exclusive one, of modification over long periods of time.
1
Variation Under Domestication

In this chapter, Darwin talks about how domesticated animals and plants have grad-
ually come to look as they do through our powers of selecting for the particular traits
that we value, over the course of many generations. The keys to this process are indi-
vidual variability and the passing of particular traits to offspring.

Variability
For any given species, when we compare members belonging to the same variety or
subvariety of our older cultivated animals and plants—​there are at least 7,500 var-
ieties of apples, for example, all members of the species Malus domestica—​one of the
first things we notice is that they generally differ more from each other than do the
individuals of any one species or variety in nature. It seems clear that organisms must
be exposed during several generations to new conditions to cause any great amount
of variation, but, once the characteristics have begun to vary, they generally continue
to vary for many generations. We know of no case in which any variable organism
has ceased to vary under domestic cultivation. Our oldest cultivated plants, such as
wheat, still yield new varieties, and our oldest domesticated animals can still be rap-
idly modified or improved because of the variations that they continue to exhibit.1

Effects of Habit and of the Use or Disuse

of Parts; Correlated Variation; Inheritance
Changed habits produce an inherited effect, as, for example, the time during the year
at which a plant begins to flower after being transported from one climate to another.
With animals, the increased use or disuse of parts has had a more marked influence.
Thus I find that in the domestic duck, the bones of the wing weigh less in proportion
to the weight of the entire skeleton than do the same bones in the wild duck, while

1 I have shortened this section considerably. As you will see throughout this book, Darwin, having no

knowledge of chromosomes, DNA, or even the basics of Mendelian genetics (first put forward by Gregor
Mendel, in an obscure paper published in 1866), was at a loss to explain the causes of variation, or how
those variations were transmitted to offspring. He returns to this topic in Chapters 2 and 5.

The Readable Darwin. Second Edition. Jan A. Pechenik, Oxford University Press. © Oxford University Press 2023.
DOI: 10.1093/​oso/​9780197575260.003.0002
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