Darwin and the

Nature of Species

David N. Stamos
Darwin and the Nature of Species

i
SUNY series in Philosophy and Biology

David Edward Shaner, editor

Darwin and the Nature of Species

David N. Stamos

State University of New York Press

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Library of Congress Cataloging-in-Publication Data

Stamos, David N., 1957-

Darwin and the nature of species / David N. Stamos.
p. cm. –
(SUNY series in philosophy and biology)
Includes bibliographical references (p. ) and index.
ISBN-13: 978-0-7914-6937-8 (hardcover : alk. paper)
ISBN-10: 0-7914-6937-9 (hardcover : alk. paper
ISBN-13: 978-0-7914-6938-5 (pbk. : alk paper)
ISBN-10: 0-7914-6938-7 (pbk. : alk. paper)
1. Species–Philosophy.
2. Darwin, Charles, 1809-1882. I. Title. II. Series.

QH83.S748 2007
578'.012–dc22

2005036225

10 9 8 7 6 5 4 3 2 1
In memory of my mentor and friend the late Robert H. Haynes,
who enjoyed to the last what he called “the opiate of Darwinism.”
This page intentionally left blank.
 Contents


Preface ix
Acknowledgments xix
1. A History of Nominalist Interpretation 1
2. Taxon, Category, and Laws of Nature 21
3. The Horizontal/Vertical Distinction and the
Language Analogy 37
4. Common Descent and Natural Classification 65
5. Natural Selection and the Unity of Science 81
6. Not Sterility, Fertility, or Niches 107
7. The Varieties Problem 131
8. Darwin’s Strategy 153
9. Concept Change in Scientific Revolutions 187
10. Darwin and the New Historiography 207

Notes 231
References 249
Index 267

vii
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 Preface


Looking back, I think it was more difficult to
see what the problems were than to solve them.
—Charles Darwin (letter to Charles Lyell, September 30, 1859)

The year 1859 marks the beginning of an enormous earthquake, an earth-

quake that shook the world and continues to shake it to this very day. The
earthquake and the consequent tremors were not caused by the gradual
shift and strain of conflicting ideas, but by a sudden impact, the publica-
tion of On the Origin of Species by Charles Darwin. It started a revolution
in thinking, an enormous paradigm shift, the implications of which are still
being worked out. Interestingly, at the very core of that revolution is the
concept of species. It is important, then, to know exactly what Darwin did
with that concept. The problem, however, is that for a variety of reasons
scholars (biologists, philosophers of biology, and professional historians of
biology) have provided interpretations that just don’t fit the facts. A large
part of the reason, as we shall see, was caused by Darwin himself. At any
rate, the problem of Darwin on the nature of species, what was the prevail-
ing view and how he tried to change that view, has yet to be adequately
understood and appreciated. The time is definitely overdue for a detailed
historical reconstruction. This becomes even more important because the
concept of species in biology, from the time of Darwin right up to today,
is still far from settled.
The purpose of this book is basically fourfold: First and foremost, to
provide a full and detailed reconstruction of Darwin’s species concept fo-
cusing mainly on his mature evolutionary period, to get it right inasmuch
as that is possible. In fact the present work breaks entirely new ground and
constitutes a major reinterpretation of Darwin on the nature of species, in
stark contrast to the literature on this topic, which stretches back over 140

ix
x PREFACE

years. Second, to apply Darwin’s insights on the ontology of species to the

modern species problem. Third, to take my reconstruction work on Dar-
win and apply it as a case study to a core issue in philosophy of science,
namely, the problem of concept change in scientific revolutions. Fourth
and finally, to use Darwin’s species concept as an indictment against a now
dominant trend in professional history of science.
I shall expand on these purposes later in this Preface, but first I want
to deal with some preliminary matters, beginning with the identity of the
specific audiences for which this book was written. Obviously it should be
of great interest to Darwin scholars. They alone will be able to fully appre-
ciate and enjoy the detailed historical work (even though it is mainly inter-
nalist) and the new direction that it takes. In fact anyone who is interested
in things Darwinian should find this book worth their while. Historians of
science, of course, should be especially interested not only for the work on
Darwin but also for my application of it to what I call in the final chapter
“the new historiography.” The second major audience is biologists and
philosophers who are interested in the modern species problem. In fact this
book serves actually as a prequel to my previous monograph, aptly titled
The Species Problem (2003), which focuses mainly on the modern species
problem (the problem of determining whether species are real, and if real
the nature of their reality; hence the problem of defining the species cate-
gory). Darwin has much to say that is both interesting and important on
this matter, although it has been almost entirely lost on subsequent
scholars. Philosophers of science should also be interested, for the recon-
struction work in the present book proves to be an enlightening case study
for the topic of concept change in scientific revolutions, so much so that it
presents a serious challenge to what many consider the received view.
The problem begins primarily with Darwin’s most famous book, the
full title of which is On the Origin of Species by Means of Natural Selection,
Or the Preservation of Favoured Races in the Struggle for Life (Darwin 1859).
In spite of the realist tone of its main title, Darwin repeatedly defines
species, both individually and as a category, nominalistically, as arbitrary
groupings and therefore as extramentally unreal. This is possibly the great-
est enigma in the history of biology, even of science. Could it be that one
of the greatest scientific minds of all time, and the main force behind what
is arguably the most important scientific revolution of all time, was simply
muddled on so basic an issue? The sheer irony is that for over a hundred
years virtually everyone took him at his word, as believing that species are
not real. Then in 1969 a major breakthrough was made by the biologist
Michael Ghiselin, in his book The Triumph of the Darwinian Method
 PREFACE xi

(1969). Ghiselin argued that species taxa for Darwin are real, such as Canis
lupus and Homo sapiens, but not the species category, the class of species taxa
and the object of a species definition. Sixteen years later John Beatty (1985)
added to Ghiselin’s thesis a strategy theory to explain why Darwin would
define species nominalistically and yet hold that species taxa are real. For
Beatty, Darwin simply followed the species designations of his fellow nat-
uralists, but denied that the species category could be defined, simply to
better communicate his evolutionary views, given that his audience had a
theory-laden definition of “species.”
Beatty’s theory has enjoyed the status of being the received view ever
since. The present book, on the other hand, is the first major-length study
of Darwin on the nature of species, and one of its themes is that the re-
ceived view should be received no more. Darwin did not simply follow the
species designations of his fellow naturalists. Moreover the places where he
declined to do so provide major evidence (in addition to other evidence) for
reconstructing his implicit species concept.
Granted, there have been a few who have attributed a species concept
to Darwin (e.g., a morphological species concept, or one involving steril-
ity to some partial degree), implying a poverty of thought on Darwin’s part,
but in each case they succeeded only in revealing the poverty of their own
research. The time is long overdue for a thorough and detailed analysis of
Darwin’s writings to bring out not only his actual species concept but also
the richness and fruitfulness of that concept.
Although I make no claim to providing the last word on the subject, I
do claim that this book marks a substantial advance. And it was not pro-
duced lightly. Rather it is the culmination of a period of research spanning
roughly twelve years, parts of which have been presented in a number of pub-
lications (Stamos 1996, 1998, 1999, 2000, 2002, 2003, 2005), but most of
which is new, the remainder being either completely rethought or refined.
When doing research on Darwin, I look at his writings much as a
paleontologist looks at strata. Stephen Jay Gould (1989) put the case of
the paleontologist best: “We search for repeated pattern, shown by evidence
so abundant and so diverse that no other coordinating interpretation could
stand, even though any item, taken separately, would not provide conclu-
sive proof ” (282). To perceive patterns that everyone else has missed, or to
provide new and revolutionary interpretations for already perceived pat-
terns, is the glory of the paleontologist (aside, of course, from discovering
new bones). Although himself not a paleontologist, one has to think of the
discovery by the physicist and Nobel laureate Luis Alvarez and his team
and their explanatory theory, namely, the discovery of high levels of
xii PREFACE

iridium at the K/T boundary and their theory of extraterrestrial impact to

explain both it and the K/T extinction. The discovery of shocked quartz a
few years later provided further considerable evidence in support. Interest-
ingly, all of this had been missed by professional paleontologists, but it has
now become the dominant theory in explanation of the mass extinction
that leveled the dinosaurs and many other species 65 million years ago.
The same can happen in professional history. In the case of Darwin, the
strata is the enormous amount of writings he left behind: his published
books and articles, his manuscripts and notebooks, his correspondence and
marginalia. Here there are still new patterns to be discovered and room for
better theories to explain already discovered patterns. And one need not be
a professional historian to do this. In fact, expertise in a different discipline,
as with the Alvarez example, might be just what is needed to see what every-
one else has missed and to thereby effect a paradigm shift. I make my case
in the following chapters.
I shall also, as I’ve stated above, apply Darwin’s insights to the mod-
ern species problem. But just what is that problem? Quite simply, it is the
problem of determining the ontological status of species taxa, whether they
are arbitrary mental constructs or real entities existing outside of the mind,
whether they are something we make or something we discover; and if the
latter is the case, it is the further problem of determining their precise
nature and of formulating it in a definition.
The modern species problem has both purely theoretical and eminently
practical dimensions. Beginning with the former, the species problem can
be seen as the central problem of the Modern Synthesis. Begun in the 1920s
with the marriage of Darwinian natural selection to Mendelian genetics, the
union of the various subdisciplines in biology, ostensibly completed in the
1950s, has so far been without a unified species concept. Within each sub-
discipline there are various contenders, and between them there has yet to
be a clear winner. In fact, in the past three decades species concepts have
proliferated in a Darwinian bush pattern, as evidenced by the recent an-
thologies devoted to the topics of species concepts and mechanisms of spe-
ciation (Otte and Endler 1989; Ereshefsky 1992a; Claridge et al. 1997;
Howard and Berlocher 1998; Wilson 1999; Wheeler and Meier 2000).
The reason for this proliferation is not only theoretical. In addition to
being the basal units of taxonomy and (as most think) the main units of evo-
lution, species are also the main units of biodiversity. Unfortunately our world
is in the midst of a major crisis in biodiversity, mass extinction #6 (mass ex-
tinction #5 being the one that occurred roughly 65 million years ago). The
main cause of the current mass extinction, of course, is not extraterrestrial, but
 PREFACE xiii

rather the rapid overpopulation of Homo sapiens and the corresponding

destruction of the environment. According to the best estimate of Edward
Wilson (1992, 278–280), we are losing 50–100 species per day and at the
present rate shall reach roughly a 50% loss in biodiversity by the year 2050.
In response to this crisis, there has risen in recent decades a noticeable
conservation movement, involving many different countries and levels of
society, from grassroots to the United Nations. The main problem with
laws and treaties is that (aside from the need for much greater funds) they
need a unified species concept if they are to be uniformly applied. The sit-
uation is the same in other areas of law. Without an agreeable definition of
pornography, for example, pornography laws cannot help but be vague or
ambiguous and will accordingly suffer in their application.
The official species concept of the U.S. Endangered Species Act of
1973 explicitly employs in its definition of “species” the biological species
concept, which is based on reproductive isolation and which was named by
its most vociferous advocate, Ernst Mayr. Unfortunately this Act was made
at a time near the end of the hegemony (at least in zoology) of the biolog-
ical species concept. The current situation in biology is clearly that of plu-
ralism, in that there are many species concepts actually in use in biology,
and many more vying for contention. Some biologists, as we shall see in the
next chapter, are species nominalists. Seemingly more are pluralists out-
right, believing that modern biology positively needs a variety of different
species concepts to suit the needs of different biologists. Many have de-
spaired of the species problem altogether and along with Robert O’Hara
(1993) think that, like a dissolved marriage, we should try to “get over”
(232) the species problem and simply get on with doing biology. Unfortu-
nately this will not make the species problem go away.
Part of the problem is that different species concepts divide up the bi-
ological world in different ways. For example, Joel Cracraft (1997, 331)
estimates that his phylogenetic species concept roughly doubles the 9,000
or so species of birds currently recognized by the biological species con-
cept. More specifically a similar problem surrounds the case of the red wolf
(Wayne and Gittleman 1995), the flagship of the U.S. conservation pro-
gram. Millions of dollars were spent by the government, capturing, breed-
ing, and reintroducing this species into the wild. Although an ambiguous
species from the viewpoint of the biological species concept, it is a good
species from the viewpoint of the morphological species concept. Recent
DNA studies, however, have confirmed that the red wolf is a hybrid of the
gray wolf and the coyote, and thus not a good species from the viewpoint
of either the biological or phylogenetic species concepts. And yet from the
xiv PREFACE

viewpoint of an ecological species concept the government’s efforts have

been well spent.
Because of the biological species concept’s lack of finer discriminations
and other problems (including hybridization and its inapplicability to
asexual forms), more and more biologists have been arguing that biology
needs a better species concept, one that is universal in scope and that fits
the needs of conservation biology. Indeed many still hold out hope for a
universal species concept, one that will complete the Modern Synthesis and
satisfy the various needs of biology, including conservation.
Now what has all of this to do with Darwin? I am certainly not under
the illusion of thinking that whatever insights can be gleaned from Darwin’s
writings will be sufficient to solve the modern species problem. But I do
think that his insights are sufficient to put us on the right track (and we are
not on the right track!). As James Mallet (1995) put it,

by 1859 he was an experienced systematist, having just finished his bar-

nacle monograph, and had accumulated an encyclopaedic knowledge
about species, both from his own travels and researches, and through
prodigious correspondence with other zoologists and botanists. His private
income left him free of bureaucracy and teaching; he had the time, the
facts at his disposal, and the intellect to solve the problem of the nature of
species. It is at least worthwhile reexamining Darwin’s arguments. [294]

Mallet characterizes Darwin’s species concept as “materialistic, morpho-

logical” (294). We shall see in later chapters that this is not at all close, and
indeed that thus far no one else has come close either. But Mallet’s point about
Darwin’s unique position and superior competence remains. In fact what we
shall see in chapter after chapter is that Darwin had a wealth of insights highly
relevant to the modern debate on species, insights that for the most part have
gone unrecognized by virtually everyone who has written on the topic.
But surely, one might reply, even if we grant Darwin’s unique position
and superior competence, the situation was far different in Darwin’s day com-
pared to today. To a large extent, of course, this is true. Although there were
many different species concepts bandied about in Darwin’s day, the species
problem was quite different from that of today. For a start, the main species
concepts back then were at bottom creationist species concepts and the main
issue was whether species are fixed or evolve. As Darwin put it in a letter to
Leonard Jenyns (October 12, 1844) with regard to “the question of what are
species,” “The general conclusion at which I have slowly been driven from a
directly opposite conviction is that species are mutable & that allied species are
co-descendants of common stocks” (Burkhardt and Smith 1987, 67).
 PREFACE xv

Today, of course, evolution is taken for granted, as a fact, so much so that

“Nothing in biology makes sense except in the light of evolution” (Dobzhan-
sky 1973). Accordingly the species problem has taken on quite a different
meaning since Darwin’s day. Although there are many issues that define the
modern species problem, there are six in particular that I shall focus on in this
book: (i) whether species are extramentally real or unreal (nominalism), (ii)
whether species are abstract classes or concrete individuals, (iii) whether species
are primarily horizontal or vertical entities, (iv) whether species can have mul-
tiple origins (polyphyletic) or must have single origins (monophyletic), (v)
whether species are primarily process or pattern entities, and (vi) whether
species must be consistent with history reconstruction. According to the
botanist Melissa Luckow (1995), “the species problem will be solved by the
continued collection and analysis of data, the clarification of issues and terms,
and the application of new ideas” (600). Darwin, I shall argue, has something
vitally interesting and important to say on each of the six issues just outlined,
so much so that although his insights do not provide a final solution to the
modern species problem they certainly help show us the way.
In chapter 1 I do not begin with reconstructing Darwin’s species con-
cept but instead spend most of the chapter providing a short history of
nominalist interpretation of Darwin on species. Part of the problem, as we
shall see right from the start, was created by Darwin himself. We shall also
see, however, that Darwin, throughout the entirety of his career as an evo-
lutionist, did indeed think that species are real.
In chapter 2 the reconstruction begins. What we shall see is that Dar-
win did have a distinction between species as a taxon and species as a cat-
egory. Moreover, he provided a number of laws of nature for the species
category, not for any particular species taxon but for the class of species taxa
as a whole. Given that a number of thinkers on the modern species problem
argue that the species category is objectively real because there are laws of
nature that apply to it, I similarly argue that the species category (not just
species taxa) was likewise objectively real for Darwin.
In chapter 3 I argue that the evidence is overwhelming that Darwin, to
use a modern distinction, conceived of species as primarily (though not ex-
clusively) horizontal entities in the Tree of Life. This is the first major step
in understanding Darwin’s view on the nature of species taxa. We shall also
see that Darwin early in his career as an evolutionist toyed with but then
rejected the idea that species are or are like individual organisms (which
are temporally vertical entities), and that he did this in favor of the many
analogies between species and languages, the latter of which for Darwin, as
today, were thought of mainly as horizontal entities.
xvi PREFACE

In chapter 4 I focus on Darwin’s emphasis on common descent for

natural classification. I show that in spite of his emphasis, Darwin did not
subscribe to a concept of monophyly for species taxa (as is common today).
Darwin’s comments on extinction are relevant here, and my conclusion
that Darwin did not insist on monophyly for species proves consistent not
only with chapter 3 on the temporal dimension of species but also with
what we shall see in chapter 5.
Chapter 5 presents the key to understanding Darwin’s species concept,
the key that everyone else has missed. What we shall find is that the key was
not morphological discreteness, or characters constant and distinct, but
adaptations. We shall see this in example after example in Darwin’s writ-
ings, and it is the sole reason for why he went against his fellow naturalists
(when he did) in their species designations. What we shall also see is that
for Darwin adaptations were the key for distinguishing species not only
because adaptations were the most amazing features of species, but because
they were produced by a natural law, namely, what Darwin called “natural
selection.” Moreover, it will be shown that by bringing species under nat-
ural law, and also by using natural law to distinguish species, Darwin in
one stroke was attempting to bring both species classification into the realm
of scientific classification (which at that time put a high premium on nat-
ural law) and biology into the unity of science.
In chapter 6 I examine what was not part of the nature of species on
Darwin’s view, namely, reproductive isolation between species, fertility
within a species, and the occupation of an ecological niche. Of particular
interest are the reasons Darwin gave for rejecting these criteria, the first two
of which enjoyed common currency in his own day and all three of which
enjoy widespread currency today. Darwin’s rejection of these criteria will
also be seen to fit exactly with the criteria shown in previous chapters that
he did accept. In short, Darwin clearly thought of species as pattern enti-
ties, not as process entities, and accordingly it is at the end of this chapter
that a formal definition of Darwin’s implicit species concept is given.
In chapter 7 I turn to a related issue, namely, Darwin’s concept of
variety. If Darwin thought that species are real and varieties are incipient
species, then he must also have thought that varieties are in some sense real
as well. Darwin’s concept of variety has been even less explored than his
concept of species, and this chapter attempts to make up for that glaring
omission. For reasons given in the chapter, the various concepts of variety
of Darwin’s fellow naturalists are also explored. This is an area of research
that has received pathetically little attention in the literature, and this
chapter attempts to make up for it.
 PREFACE xvii

In chapter 8 I develop my own theory for why Darwin in the Origin

and elsewhere explicitly denied that species (both category and taxa) are
real and yet gave numerous indications elsewhere that he thought they are
real. The theories of Ghiselin (1969), Mayr (1982), Beatty (1985), Hodge
(1987), McOuat (1996, 2001), and my former self (Stamos 1996) are
examined in detail and rejected, before presenting in detail what I believe
to be the true pattern of Darwin’s modus operandi.
In chapter 9 I broaden my focus and examine one of the basic issues
in history and philosophy of science, namely, the problem of correctly mod-
eling the nature of concept change in scientific revolutions. Beatty’s (1985)
interpretation of Darwin on species, which quickly became the received
view, was, we shall see, influenced by a preconceived model of such change.
Since his interpretation of Darwin on species does not fit the evidence, his
failure raises anew the problem of a historically correct model. John
Dewey’s (1910) famous get over it model, we shall see, fails to refer, as well
as the highly influential incommensurability thesis of Thomas Kuhn (1970,
1977), while Philip Kitcher’s (1978, 1993) model of reference potential
receives surprising corroboration (for both “species” and “variety”), even
though he did not recognize it because (like so many others) he followed
Beatty’s (1985) strategy theory.
Chapter 10 shifts focus to an even more basic issue in history of science,
namely, the issue of historiography. The current trend in professional his-
tory of science is externalist (the sociology of ideas), embedding scientists
like Darwin in their time and place and keeping them there, in opposition
to (even so much as trying to replace) the previous trend which was inter-
nalist (the history of ideas). The book presented here, of course, is not
meant to be a competitor to biographies of Darwin, such as Desmond and
Moore’s (1992) controversial though highly influential biography. Never-
theless it is definitely meant to be a counterbalance to the modern trend in
historiography, typified by Desmond and Moore’s book. Indeed this final
chapter, which builds on the chapters that precede it, serves as an indict-
ment against that trend. To accomplish that task, I keep my eye on Dar-
win on the nature of species and respond not only to Desmond and Moore
but to a number of other professional historians of biology who either try
to denigrate the Darwinian revolution or Darwin’s role in that revolution.
Although science is a human activity and all scientists work in a social
context, when viewed from a bird’s-eye view it should be clear that science
does, to varying degrees, transcend its social milieu. Science is not merely
a social construction. There are genuine revolutions in science and genuine
progress in science, in an objective, epistemic sense, unlike, for example, the
xviii PREFACE

history of music. Moreover there are individual scientists who transcend

not only their social milieu but also the science of their time. There is no
finer example of this than Darwin. This is the grand theme of Michael
Ghiselin’s The Triumph of the Darwinian Method (1969), which I highly
recommend. Although dated from the viewpoint of modern historiogra-
phy, his book is still essential reading and is, on my view, basically correct.
The present book extends Ghiselin’s thesis in the one area in which it is
truly outdated, namely, the issue of Darwin on the nature of species.
Although Ghiselin did not get it right, he did nevertheless take great strides
in the right direction. My ultimate thesis is that with his species concept
Darwin belongs in the present time much more than his own, so much so
that he still has plenty to teach us.
 Acknowledgments


Special thanks to Editor in Chief Jane Bunker, Series Editor David Shaner,
the two anonymous readers for SUNY, Michael Ghiselin, Polly Winsor,
Jon Hodge, Gordon McOuat, David Johnson, Bernie Lightman, and last
but not least Sharon Weltman for helping with the initial proofreading of
the manuscript.

xix
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 Chapter 1

A History of Nominalist Interpretation

Ever since the publication of Darwin’s Origin, biologists, historians, and

philosophers have interpreted Darwin as being a species nominalist. Species
nominalism is the view that species are not real, that they are not out there
in nature, existing irrespective of observation, but rather that they are man-
made, like monetary currency or constellations, so that, from an objective,
naturalistic point of view, they are real in name only.
This “received view” is based mainly on a literal reading of a number
of passages in the Origin. In this chapter I shall begin by examining those
passages. Following that I shall go back and examine Darwin’s species con-
cept(s) in his early period as an evolutionist, the period of his transmuta-
tion notebooks. I shall then proceed briefly up through the strata of his
writings, trying to find where his supposed species nominalism began. I
shall then take a brief excursion through the secondary literature, begin-
ning with reviews of Darwin’s Origin and proceeding right up to today. It
will be interesting to see how the perception of Darwin as a species nomi-
nalist has been employed by a number of authors. Finally, I shall then ex-
amine how Darwin himself replied to the charge of species nominalism, as
well as examine some other evidence which, together with what we shall see
in subsequent chapters, should lead one to conclude that Darwin was in fact
a species realist. In the very least, the end of this chapter along with the
next four should bring to a close the easy days of finding in Darwin an ally
for species nominalism.

1
2 DARWIN AND THE NATURE OF SPECIES

Beginning with the Origin (1859),1 in the concluding chapter Darwin

proclaims that as a result of his investigations “we shall have to treat species
in the same manner as those naturalists treat genera, who admit that gen-
era are merely artificial combinations made for convenience. This may not
be a cheering prospect; but we shall at least be freed from the vain search
for the undiscovered and undiscoverable essence of the term species” (485).
This passage relates to both halves of a modern distinction that partly de-
fines the modern species problem, namely, the distinction between species
as a taxon and species as a category, a distinction not always recognized but
made much of by, for example, Ernst Mayr (e.g., 1987, 146). Again, briefly,
species taxa are particular species, each of which is given a binomial, such
as Tyrannosaurus rex or Homo sapiens. The species category, on the other
hand, is the class of all species taxa. Among realists, the species category is
captured in their respective definitions of the species concept. Thus, what
is a genuine species according to one definition might not be counted as a
genuine species according to another definition. A species nominalist, of
course, would say that species definitions are ultimately arbitrary, because
species taxa are ultimately arbitrary.
In the passage from Darwin’s Origin quoted above, he seems quite
clearly in the first part to assert that species taxa are unreal. He says that we
shall have to treat species in the same way as genera nominalists treat gen-
era, as not real but man-made, made simply for the sake of convenience.2
In the second part of the passage, by referring to the “term species,” Dar-
win seems clearly to be referring to the species category. There are other pas-
sages in the Origin that seem to second this view. For example, he says
“From these remarks it will be seen that I look at the term species, as one
arbitrarily given for the sake of convenience to a set of individuals closely
resembling each other” (52). This passage is often quoted as supporting the
interpretation of Darwin as a species nominalist, but it has to be remarked
that the context of the passage makes it clear that Darwin is drawing his
conclusion not from nature or from his own theory of evolution but from
the taxonomic behavior of other naturalists. For in the previous paragraph
he states that “If a variety were to flourish so as to exceed in numbers the
parent species, it would then rank as the species, and the species as the va-
riety” (52). This was not Darwin’s view. Instead it was a practice common
to his fellow naturalists.
Indeed, part of Darwin’s overall argument for evolution was that in
many cases expert naturalists could not themselves agree on whether a par-
ticular form was a variety or a species. For example, in the Origin he says
“wherever many closely-allied species occur, there will be found many forms
 A History of Nominalist Interpretation 3

which some naturalists rank as distinct species, and some as varieties; these
doubtful forms showing us the steps in the process of modification” (404;
cf. 47, 49, 111, 248, 296–297). It was essential for Darwin that there be
no clear distinction between species and varieties, otherwise varieties could
not be what he called “incipient species” (52, 111, 114, 128), and the fact
that expert naturalists could not agree in many cases on what is a species and
what is a variety added a further prong in his attack on the fixity of species
(in addition to his arguments from the fossil record, from biogeography,
from embryology, from artificial breeding, etc.). And so again and again in
the Origin we see Darwin assert that there is no essential or fundamental
distinction between species and varieties. For example, he says “neither
sterility nor fertility affords any clear distinction between species and vari-
eties; but that the evidence from this source graduates away, and is doubt-
ful in the same degree as is the evidence derived from other constitutional
and structural differences” (248; cf. 51–52, 268, 272, 484–485).
A further part of Darwin’s argument was that not only did naturalists
in many cases disagree on what is a species and what is a variety, but they
themselves could not agree on a definition of the species category. Even
though, as Darwin early in the Origin recognized, “most naturalists” viewed
species as “independently created” (6)—one might call this the common
denominator3—they nevertheless gave “various definitions . . . of the term
species” (44), definitions that concerned mainly the diagnostic criteria. This
created a problem in itself, for as Darwin later in the Origin pointed out,
“to discuss whether they [‘many forms’] are rightly called species or vari-
eties, before any definition of these terms has been generally accepted, is
vainly to beat the air” (49).
And yet Darwin clearly recognized in the Origin the need for species
talk. Consequently, on the issue of whether a particular form should be
ranked as a species or a variety, he took the position that “the opinion of
naturalists having sound judgment and wide experience seems the only
guide to follow,” and where they disagree the problem is to be settled sim-
ply by appealing to “a majority of naturalists” (47). This, of course, has an
arbitrary ring to it. And indeed Darwin in the Origin, as we have seen ear-
lier in this chapter, in apparent reference to his contemporaries, stated that
he looks “at the term species, as one arbitrarily given for the sake of conve-
nience” (52). Furthermore, again as we have seen earlier, in his concluding
chapter Darwin took his own position that there is in fact no essential and
fundamental distinction between species and varieties as a liberating one,
since systematists “will not be incessantly haunted by the shadowy doubt
whether this or that form be in essence a species” (484) and “shall at last be
4 DARWIN AND THE NATURE OF SPECIES

freed from the vain search for the undiscovered and undiscoverable essence
of the term species” (485).
Small wonder, then, given all of the above, that scholars have com-
monly interpreted Darwin as a species nominalist, as we shall see later in
this chapter. And yet how utterly odd, if those scholars are right, that Dar-
win would title his book On the Origin of Species, let alone with the addi-
tion by Means of Natural Selection! That the received view is wrong, that it
is based on a superficial reading of Darwin, is something I shall argue later.
For now, we need to ask when such apparently nominalist talk on Dar-
win’s part began.
Certainly it did not start when Darwin began developing his evolution-
ary views. In his transmutation notebooks (Barrett et al. 1987) Darwin pro-
vides a number of definitions of “species,” all realist in tone. Sometimes his
definition is in terms of constant characters, as in Notebook B, begun in
July 1837: “Definition of Species: one that remains at large with constant
characters, together with other beings of very near structure” (213).
In other definitions Darwin focused on interbreeding, as in Notebook
C, begun in March 1838 and finished in July of the same year: “A species
is only fixed thing with reference to other living being—one species May
have passed through a thousand changes, keeping distinct from other & if
a first & last individual were put together, they would not according to all
analogy breed together” (152). Darwin at some time later added an anno-
tation to this page, writing “As species is real thing with regard to contem-
poraries—fertility must settle it.” This page, both the original passage and
the annotation, is interesting for its relation to the modern biological species
concept made famous by Dobzhansky (1937) and especially Mayr (1942,
1970), which is based on interbreeding populations and genetic reproduc-
tive isolation mechanisms. What makes it interesting is not the emphasis on
the fertility test. This was common in Darwin’s time and before, having
been made famous by Buffon (Lovejoy 1959). Instead, what is interesting
about Darwin’s passage is that as a species evolves radically over time, so
that vertically in the Tree of Life it would in principle be incapable of in-
terbreeding with its originals if they could be brought together, Darwin
still insists on the reality of species at any given horizontal dimension, at any
given cross-section in time. Like the modern biological species concept,
Darwin, the evolutionist, provided a horizontal species concept and fixed
the reality of species in the horizontal dimension, unlike a number of mod-
ern species concepts that insist on the vertical reality of species. What we
shall see in Chapter 3 is that Darwin maintained this view in the Origin and
that his main analogy for species (namely, languages) provides a powerful
 A History of Nominalist Interpretation 5

reason for believing that we today should also conceive of the reality of
species primarily as horizontal rather than as vertical entities.
Interestingly, a little later in Notebook C Darwin seems to slightly
change his mind, insisting now that “My definition of species. has nothing
to do with hybridity,, is simply, an instinctive impulse to keep separate,
which no doubt be overcome, but until it is the animals are distinct species”
(161). However, earlier in Notebook B he had already used this criterion,
when with regard to speciation he wrote that “repugnance to intermar-
riage—settles it” (24).
Indeed there can be little doubt that in his transmutation notebooks
Darwin waffled between fertility/sterility and instinct. For example, in
Notebook E, in an entry dated between October 16 and 19 of 1838, he
states that “If they give up infertility in largest sense, as test of species.—they
must deny species which is absurd.—their only escape is that rule applies
to wild animals only. from which plain inference might be drawn that
whole infertility was consequent on mind or instinct, now this is directly
incorrect” (25). Similarly in his abstract of John Macculloch’s Proofs and Il-
lustrations of the Attributes of God (also in Barrett et al. 1987), which Dar-
win probably wrote in late 1838, he wrote “With respect to whether
Galapagos beings are species, . . . it his highly unphilosophical to assert,
that they are not species, until their breeding together has been tried” (167).
Except for his evolutionary perspective with his emphasis on the horizon-
tal reality of species, Darwin in the above was doing nothing new. Indeed,
years earlier James Prichard (1813, 3–15) provided a fairly detailed summary
of the various criteria by which naturalists characterized species, which in-
cluded not only constant character differences and the sterility test but also
instinctual repugnance, immunological differences, and parasitological differ-
ences. It is not known whether Darwin had, by the time of the transmuta-
tion notebooks, read Prichard or a later edition, but it would not seem to
matter, since he could have gotten the same ideas from other sources.
Turning now to Darwin’s Sketch of 1842 and his Essay of 1844 (Dar-
win 1909), although they contain many ideas that are to be found later
elaborated in the Origin, such as the idea that sterility is not an unfailing
test, or that there are many forms about which expert naturalists cannot
agree on whether they are species or varieties, there is not, unlike the Ori-
gin, any clear hint of species nominalism. Beginning with the Sketch, we
find, actually, just like the transmutation notebooks, statements to the con-
trary. For example, Darwin says “Looking now to the affinities of organ-
isms, without relation to their distribution, and taking all fossil and recent,
we see the degrees of relationship are of different degrees and arbitrary—
6 DARWIN AND THE NATURE OF SPECIES

sub-genera—genera—sub-families, families, orders and classes and king-

doms” (35). Granted, Darwin is referring to his contemporaries, for he fol-
lows this passage with the sentence “The kind of classification which
everyone feels is most correct is called the natural system, but no one can
define this.” Nevertheless, what is interesting is that with other higher taxa
nominalists of his time (and most were higher taxa nominalists), Darwin
does not include species in his list of arbitrary categories. Moreover, a lit-
tle later in the Sketch he writes of “undoubted species” (48) and of “real
species” (49). There is, however, immediately following, a passage that hints
of species nominalism. In reference to “real species,” which are distinct by
every criterion, but admitting common descent, he writes “Can genera re-
strain us; many of the same arguments, which made us give up species, in-
exorably demand genera and families and orders to fall, and classes
tottering. We ought to stop only when clear unity of type, independent of
use and adaptation, ceases” (49). But here there is no reason to suppose
that by the phrase “giving up species” Darwin means giving up the reality
of species. A much more natural reading, given the basic presupposition of
the majority of his antagonists, is “giving up the independent creation of
species,” or “giving up the fixity of species,” which amounts to the same
thing. Indeed we shall see in this and in later chapters that the phrase “giv-
ing up species” was a common one with Darwin, even though, again as we
shall see in this and in later chapters, he did not give up their reality.
Turning now to the much longer Essay, beginning with the second
chapter where he first uses the word “breed,” Darwin added a note in the
manuscript, writing “Here discuss what a species is” (81). However, unlike
the Origin, Darwin did not follow through. Instead, much like the Sketch,
even though he raises numerous problems for the independent creation of
species and their fixity, and argues for evolution, he still continues to write
of species as real. In fact, much like the Sketch, even though he denounces
the categories of his contemporaries above the species level as “quite arbi-
trary” (202, 204–205), he continues to write of “true species” (204, 212,
241, 243, 246). And so unlike the Origin, with its many implied references
to species as arbitrary, and with its concluding chapter which states that we
shall have to treat species taxa as artificial and made for convenience, there
is absolutely none of this in the Essay, neither in its nine argumentative
chapters nor in its tenth concluding chapter.
Turning next to Darwin’s correspondence, we do not find any clear
signs of species nominalism until well into the 1850s. In fact, until that
time, the impression we get from Darwin’s correspondents is that most ex-
perienced naturalists believed in the reality of species, and Darwin, in turn,
 A History of Nominalist Interpretation 7

does not indicate that his view was otherwise. For example, in a letter from
his closest friend and main correspondent, the botanist Joseph Dalton
Hooker (September 4–9, 1845), Hooker wrote that “Those who have had
most species pass under their hands as Bentham, Brown, Linneaus, De-
caisne & Miquel, all I believe argue for the validity of species in nature”
(Burkhardt and Smith 1987, 250). In his reply letter (September 10, 1845),
Darwin recognized that “Lamarck is the only exception, that I can think of,
of an accurate describer of species at least in the invertebrate kingdom, who
has disbelieved in permanent species” (253). (As we shall see later in this
chapter, Lamarck did more, and disbelieved in the reality of species.) In-
stead, the main effect that Hooker’s letter had on Darwin was to doubt his
own competence to theorize about species. Remarkably, Darwin wrote
“How painfully (to me) true is your remark that no one has hardly a right
to examine the question of species who has not minutely described many”
(253). Darwin was especially taken by Hooker’s extended criticism of the
French writer Frédéric Gérard, who argued for species nominalism caused
by his poor understanding of messy situations in nature and his lack of ex-
perience. (Indeed in an earlier letter to Darwin, written in late February
1845, Hooker states Gérard’s species nominalism explicitly, and offers to
send Darwin a copy of Gérard’s tract; Burkhardt and Smith 1987, 149.)
And even though Hooker would reply that he by no means meant to imply
that Darwin was not in a position to theorize about species, Darwin did not
attempt to procure species nominalism from his evolutionary theories. In-
stead, toward the end of 1846 he embarked on an eight-year taxonomic
study of barnacles, which he completed in September of 1854.
Much of what is interesting about Darwin’s work on barnacles is that
he was struck by the variability of organisms. What we shall see in chapter
3, when we focus on his published works on barnacles, is that even though
he stressed that variability, he did not talk of species as if they were arbitrary.
Instead he argued that most species of barnacles, even when minutely stud-
ied, turn out to be taxonomically good species. Equally revealing is what
Darwin wrote in his correspondence. However problematic was the vari-
ability of barnacles taxonomically, Darwin still did not espouse species
nominalism. The following reply letter to Hooker (June 13, 1850) per-
fectly captures Darwin’s thinking throughout this period:

You ask what effect studying species has had on my variation theories; I
do not think much; I have felt some difficulties more; on the other hand
I have been struck (& probably unfairly from the class) with the variabil-
ity of every part in some slight degree of every species: when the same
8 DARWIN AND THE NATURE OF SPECIES

organ is rigorously compared in many individuals I always find some slight

variability, & consequently that the diagnosis of species from minute dif-
ferences is always dangerous. I had thought the same parts, of the same
species more resembled than they do anyhow in Cirripedia, objects cast
in the same mould. Systematic work wd be easy were it not for this con-
founded variation, which, however, is pleasant to me as a speculatist
though odious to me as a systematist. [Burkhardt and Smith 1988, 344]

Other letters confirm this view. For example, earlier in the same year, in a
letter to J.J. Steenstrup (January 25, 1850), Darwin wrote “I much dislike
giving specific names to each separate valve, & thereby almost certainly
making three or four nominal species for each true species” (Burkhardt and
Smith 1988, 306).
Granted, toward the end of his work on barnacles, Darwin had be-
come quite tired of detailed species work, so much so that he started to
sound like he might be swaying to species nominalism. In an often-quoted
letter to Hooker (September 25, 1853), Darwin wrote

In my own cirripedial work (by the way, thank you for the dose of soft
solder [i.e., flattery—OED], it does one, (or at least me) a great deal of
good,—in my own work, I have not felt conscious that disbelieving in the
permanence of species has made much difference one way or the other; in
some few cases (if publishing avowedly on doctrine of non-permanence)
I shd. not have affixed names, & in some few cases shd. have affixed names
to remarkable varieties. Certainly I have felt it humiliating, discussing &
doubting & examining over & over again, when in my own mind, the
only doubt has been, whether the form varied today or yesterday (to put a
fine point on it, as Snagsby would say). After describing a set of forms,
as distinct species, tearing up my M.S., & making them one species; tear-
ing that up and making them separate, & then making them one again
(which has happened to me) I have gnashed my teeth, cursed species, &
asked what sin I had committed to be so punished: But I must confess,
that perhaps nearly the same thing wd. have happened to me on any
scheme of work. [Burkhardt and Smith 1989, 155–156]

What is typically overlooked, however, is what Darwin says to Hooker at the

very end of his letter: “whether you make the species hold up their heads or
hang them down, as long as you don’t quite annihilate them or make them
quite permanent; it will all be nuts to me [i.e., a source of pleasure or delight—
OED].” Darwin was not yet talking the language of species nominalism.
The fact is, we don’t first start to find species nominalism talk in Dar-
win until we turn to his long though unfinished book on species evolution,
 A History of Nominalist Interpretation 9

titled Natural Selection, which was begun in mid-1856 and stopped on June
18, 1858, when Darwin received the letter from Alfred Russel Wallace ba-
sically anticipating Darwin’s views (which was of course to spark the writ-
ing of his Abstract, later to be titled On the Origin of Species). As Stauffer
(1975, 7–9) points out, Darwin waited roughly 20 years to publish his evo-
lutionary views because he wanted to present a strong scientific case for evo-
lution (more particularly, evolution by natural selection and divergence)
and thus avoid the scientific ridicule heaped upon earlier writers on evolu-
tion, in particular, Lamarck and Chambers. Before we turn to Natural Se-
lection, though, we have to wonder why Darwin would wait so long to start
sounding like a species nominalist.
One theory that might suggest itself follows from the important work of
Dov Ospovat (though I know of no one who has used Ospovat to develop
it). According to Ospovat (1981), from the time Darwin hit upon natural se-
lection in his transmutation notebooks, through the Sketch and Essay, and
until he had finished his barnacle work, Darwin shared with his contempo-
raries the belief (which was theologically based) in harmony and perfect adap-
tation in nature, with variation being minor, so that in his view natural
selection worked only intermittently, in those periods when changes in con-
ditions meant that a species was no longer perfectly adapted to its environ-
ment. Between September 1854, however, and June 1858, when he received
the shocking letter from Wallace, Darwin’s view on variation and adaptation
gradually though radically changed, from perfect adaptation with intermit-
tent natural selection to imperfect adaptation with continuous natural selec-
tion. One might think that this new view would have occurred to him early
on in his barnacle work. But Ospovat (ch. 7) argues that it was not until after
Darwin finished his barnacle work that he sat down to seriously rethink his
theory of evolution. The main problem was to explain the treelike, group
nested in group, hierarchical classification schemes of his fellow naturalists.
Darwin’s solution was his principle of divergence, which he developed in the
period from 1854 to 1858 and which, in a letter to Hooker (June 8, 1858),
he called (along with natural selection) “the key-stone of my Book”
(Burkhardt and Smith 1991, 102). According to this principle, wide-ranging
species will typically be exposed within their range to a variety of conditions,
most importantly to empty niches (to use modern terminology) which they
will tend to fill, and hence evolve in a branchlike fashion.
Based on Ospovat’s work, then, one might conjecture that prior to
1854—prior to when Darwin started rethinking his theory and still believed
in perfect adaptation with only intermittent natural selection—Darwin
would naturally think that species are real so long as natural selection is not
10 DARWIN AND THE NATURE OF SPECIES

working upon them (possibly there was an influence from Lamarck here;
cf. note 6), so that once his view changed to imperfect adaptation with con-
tinuous natural selection he consequently became a species nominalist.
This is an interesting conjecture. Unfortunately it fails for the fact, as
we shall see in this and in later chapters, that under his skin Darwin was not
really a species nominalist, not in his post-barnacle period nor in the Ori-
gin or beyond.
There is something else, however, which went on in the period be-
tween 1854 and 1858, which does help to fully explain Darwin’s species
nominalism talk, begun in Natural Selection. The evidence is in his corre-
spondence. As pointed out earlier, Darwin began his big species book, what
was to be his heavily detailed case for evolution to the world, in mid-1856,
and the problem was to avoid the scientific ridicule heaped upon the mainly
speculative attempts of earlier writers, in particular Lamarck and Cham-
bers. The problem, in short, was to convince expert naturalists more than
anyone else. What Darwin got from botanist correspondents such as
Hooker, but mainly from Hewett Cottrell Watson, was that the very con-
cept of species itself was a major impediment to convincing the scientific
world that species are not fixed but evolve.
Interesting in this regard is a letter from Hooker (July 8, 1855), in which
Hooker comments on a Himalayan thistle intermediate between two com-
mon species of English thistles. Hooker writes “The more I study the more
vague my conception of a species grows, & I have given up caring whether
they are all pups of one generic type or not” (Burkhardt and Smith 1989,
372). Hooker goes on to say that not caring anymore whether this or that is
a real species forms no impediment to tracing character distribution and dis-
covering the laws of distribution, which he thinks “is certainly all we can ex-
pect to prove in our day” (372). Here Darwin may have begun to realize that
the species concept, when trying to get his evolutionary views across, pre-
sented more of an impediment than anything else, and so was best bypassed.
And yet, interestingly, when we turn to chapter 8, we shall see that in his
correspondence, when Darwin is trying to convince an important naturalist
of his views on evolution, he uses the language of species nominalism, but
only until he is convinced that he had a convert, after which time he reverted
to the language of species realism. (Indeed, as I argue later in this chapter and
in chapters 2 through 6, Darwin had an implicit but fairly clear species
concept that was both realist and evolutionary.)
If Hooker’s letter did not make Darwin think of the value of not getting
bogged down on the topic of what a species really is when presenting his sci-
entific case to the world for evolution, one of the letters from Watson almost
 A History of Nominalist Interpretation 11

certainly did. Watson, who was converted to evolution (Watson 1845a)

shortly after reading Chambers’ Vestiges, anonymously published in 1844,
wrote to Darwin (August 13, 1855) that “The grand difficulty for natural-
ists or botanists of our turn of thought, is, that the use of the word ‘species’
by technical describers is indefinite & variable. . . . Practically, it means only
an idea of the mind, with no more real restriction in its application to objects,
than have the words ‘genus’ or ‘order.’” Watson then cites Hooker and the
French botanist Alexis Jordan as examples of lumpers and splitters respec-
tively (the former grouping varieties into species, the latter making a species
out of the smallest variety). Watson goes on to say “In all my attempts to ad-
vance geographical botany, I am stopt by the application & signification of
the word ‘Species.’ Where I seek to effect precise comparisons of objects &
numbers & proportions,—that word constantly frustrates & makes vague &
indefinite” (Burkhardt and Smith 1989, 406).
Indeed, turning now to Natural Selection, we can see the influence of
Watson, on both the “grand difficulty” presented by the variability of
species concepts in Darwin’s contemporaries, as well as the implicit sugges-
tion that it is better to bypass the concept altogether.4 First, in a choice of
words echoed shortly after in the Origin, he says “In the following pages I
mean by species, those collections of individuals, which have commonly
been so designated by naturalists” (Stauffer 1975, 98).
What is equally interesting is what Darwin wrote immediately before this:

. . . how various are the ideas, that enter into the minds of naturalists
when speaking of species. With some, resemblance is the reigning idea &
descent goes for little; with others descent is the infallible criterion; with
others resemblance goes for almost nothing, & Creation is everything;
with others sterility in crossed forms is an unfailing test, whilst with oth-
ers it is regarded of no value. At the end of this chapter, it will be seen that
according to the views, which we have to discuss in this volume, it is no
wonder that there should be difficulty in defining the difference between
a species & a variety;—there being no essential, only an arbitrary differ-
ence. [Stauffer 1975, 98]

This passage compares, interestingly, with a letter Darwin wrote to

Hooker (December 24, 1856) at roughly the same time:

I have just been comparing definitions of species, & stating briefly how
systematic naturalists work out their subject: . . . It is really laughable to
see what different ideas are prominent in various naturalists minds, when
they speak of “species” in some resemblance is everything & descent of
12 DARWIN AND THE NATURE OF SPECIES

little weight—in some resemblance seems to go for nothing & Creation

the reigning idea—in some descent the key—in some sterility an unfail-
ing test, with others not worth a farthing. It all comes, I believe, from try-
ing to define the undefinable. [Burkhardt and Smith 1990, 309]

In later chapters, after examining what I believe to be Darwin’s objec-

tive set of criteria for delimiting species taxa, only then will the disingenu-
ous nature of these passages become apparent, especially when put in their
context, and only then will it make sense to develop in detail a strategy the-
ory to explain them (chapter 8).
For the present, it will be useful to examine how reviewers of the Ori-
gin responded to the apparent species nominalism of that book. The first
point to notice, using late 1859 and 1860 as typical, is that many if not
most of the reviewers simply bypassed the issue of Darwin’s apparent
species nominalism. They didn’t so much as even mention it. Instead they
focused on Darwin’s argument for evolution, in the main rejecting it (e.g.,
Anon. 1859; Crawfurd 1859; Leifchild 1859; Murray 1859; Anon. 1860a;
Anon. 1860b; Bowen 1860; Haughton 1860; Sedgwick 1860; Simpson
1860; Wilberforce 1860).
Even among Darwin’s supporters, his apparent species nominalism was
typically ignored (e.g., Chambers 1859; Hooker 1859; Huxley 1859b,
1859c, 1860a, 1860b; Carpenter 1860; Gray 1860b).
Returning to his critics, there were some, however, who did indeed
take Darwin’s apparent species nominalism to be in fact his position. For
example, Louis Agassiz (1860b) raised what seemed to him a perfectly log-
ical point: “If species do not exist at all, as the supporters of the transmu-
tation theory maintain, how can they vary? and if individuals alone exist,
how can the differences which may be observed among them prove the
variability of species?” (143). Richard Owen (1860) claimed that “on the
hypothesis of ‘natural selection’ . . . . the species, like every other group, is
a mere creature of the brain; it is no longer from nature” (532), which he
rejects on what he calls “present evidence from form, structure, and procre-
ative phenomena.” Instead he agrees with the Linnean axiom that species
are the work of nature, which he quotes as “Classis et Ordo est sapientiæ,
Species naturæ opus” (532). Thomas Vernon Wollaston (1860) referred ex-
plicitly to a page of the Origin where we find apparent species nominalism
and wrote that “it is no sign of metaphysical clearness when our author
(p. 51) refuses to acknowledge any kind of difference between ‘genera,’
‘species,’ and ‘varieties,’ except one of degree” (133), which was, he contin-
ued, “to throw doubt on a distinction between essentially different ideas”
 A History of Nominalist Interpretation 13

(134). Others, without claiming or implying that their interpretation of

Darwin’s species nominalism came explicitly from Darwin himself in the
Origin, claimed that species nominalism followed from his theory of evo-
lution. John Dawson (1860), for example, claimed that Darwin’s book
“seeks . . . to reduce all species to mere varieties of ancient and perhaps per-
ished prototypes” (101) and that with his doctrine we “break down the dis-
tinction between species and varieties as to deprive our classifications of
any real value” (119). Similarly, William Hopkins (1860) wrote that “all
theories—like those of Lamarck and Mr. Darwin—which assert the de-
rivation of all classes of animals from one origin, do, in fact, deny the exis-
tence of natural species at all,” where by “natural species” he means “the
grouping is formed by nature,” whereas with “artificial species” the group-
ing is “arbitrary” (747).
Among Darwin’s supporters, so too did some recognize his species
nominalism, although often they did not actually quote Darwin as such
but inferred it from his views. Asa Gray (1860a), for example, argued that
it follows from Darwin’s theory that whether the human races constitute
one species or more is to be settled “according to the notions of each nat-
uralist as to what differences are specific” (158). Interestingly, against Agas-
siz, who in his species concept “discards the idea of a common descent as
the real bond of union among the individuals of a species, and also the idea
of a local origin,—supposing, instead, that each species originated simul-
taneously, generally speaking over the whole geographical area it now oc-
cupies or has occupied” (155), Gray claims that his (Agassiz’s) theory
equally makes species “subjective and ideal” (158)! This is an interesting use
of Darwin’s species nominalism. Henry Fawcett (1860) too, although he
did not quote anything from the Origin as espousing species nominalism,
implied that it also followed from Darwin’s view. Repeating (though mag-
nifying) the radical disagreement between Babington and Bentham on the
number of species of English plants (cf. Origin, 48), Fawcett writes “The
question of species may thus, at the first sight, appear to be a dispute about
an arbitrary classification, and it may naturally be asked, Why, therefore,
does the problem of the Origin of Species assume an aspect of supreme sci-
entific interest?” (82). Similarly George Henry Lewes (1860), likewise feed-
ing off the disagreement between naturalists over whether a particular form
is a species or a variety (which of course Darwin himself made much of in
the Origin), writes that “The reason of this uncertainty is that the thing
Species does not exist: the term expresses an abstraction, like Virtue or
Whiteness; not a definite concrete reality, which can be separated from
other things, and always be found the same” (443).5
14 DARWIN AND THE NATURE OF SPECIES

What we have to keep in mind in all of this is that in Darwin’s time,

so unlike today, the equation of evolution with species nominalism was
deeply entrenched. And arguably it was Lamarck who began this equation.
In the first chapter of his book on evolution (Lamarck 1809), he states that
all divisions of nature into classes, orders, families, genera, and species are
“artificial devices” (20), that “they appear to derive from certain apparently
isolated portions of the natural series with which we are acquainted,” and
that nature has produced “only individuals who succeed one another and
resemble those from which they sprung.” The relation of individual organ-
isms to the natural series, he immediately goes on to say, is that “these in-
dividuals belong to infinitely diversified races; which blend together every
variety of form and degree of organisation; and this is maintained by each
without variation, so long as no cause of change acts upon them” (21).6
Consequently we find Charles Lyell (1832), as he begins his long cri-
tique of Lamarck’s evolutionism, state the issue as “whether species have a
real and permanent existence in nature; or whether they are capable . . . of
being indefinitely modified in the course of a long series of gradations?”
(1; cf. 23), which, following his critique, he concludes that “it appears that
species have a real existence in nature, and that each was endowed, at the
time of its creation, with the attributes and organization by which it is now
distinguished” (65). This dichotomy—either species are permanent and
therefore real or impermanent and therefore unreal—is repeated again and
again in the literature of Darwin’s time. For example, William Whewell
(1837 III) wrote that “in short, species have a real existence in nature, and a
transmutation from one to another does not exist” (576). A further exam-
ple is Watson (1845b), who after arguing empirically about the mutability
of primroses and cowslips, wrote that “If we allow the cowslip and prim-
rose to be two species, and yet allow that one can pass into the other, either
directly or through the intermediate oxlip, we abandon the definition of
species, as usually given, and fall into the transition-of-species theory. . . .
Let a few other cases be adduced, between reputed species equally similar,
and we shall be forced to recast our ideas and definition of the term
‘species.’ It would unavoidably become arbitrary and conventional; with
no more exactness or constancy of application, than we can give to the
terms ‘genus’ or ‘order’” (219). As one final example, Wollaston (1860)
claimed that either species are permanent and real (the traditional species
concept) or else we are left with “the otherwise hopeless task of understand-
ing what a species really is” (133), which may be taken as an epistemolog-
ical assertion only, but possibly also as an ontological one.
 A History of Nominalist Interpretation 15

So it was easy and natural for reviewers to read species nominalism in

Darwin’s Origin and to see no need to scratch beneath the surface. In later
commentators on Darwin’s Origin, however, living in a different scientific
milieu, what we often find is that those who interpret Darwin as a species
nominalist do so to use Darwin as an imprimatur for their own nominal-
ist arguments. We shall also find, of course, that they just plain overlooked
the evidence for Darwin’s species realism.
A good example to begin with is E.B. Poulton (1903), a naturalist and
selectionist whom Mayr (1982, 272) took to be a “pioneer” of the biolog-
ical species concept. In reply to Max Müller, who claimed that in spite of
the title of the Origin Darwin never gave us a species concept, Poulton (78)
replies that Darwin did and that it is given at the end of the Origin where
he says “Systematists will have only to decide (not that this will be easy)
whether any form be sufficiently constant and distinct from other forms,
to be capable of definition; and if definable, whether the differences be suf-
ficiently important to deserve a specific name” (484). Throughout his paper
Poulton gives the impression that Darwin was a precursor of the syngamic
species concept which he himself prefers, “syngamic” meaning “free inter-
breeding under natural conditions” (90), an “inter-breeding community”
(94). But Poulton does nothing to elaborate on Darwin’s species concept,
for example, whether it includes sterility between forms or even whether in-
deed Darwin himself thought that species are fully syngamic. Instead, he re-
peatedly emphasizes the “subjective character” (89), the “subjective
element” (92), the “subjective criterion” (93) in Darwin’s species concept.
This, however, is not to attribute to Darwin species realism. Indeed natu-
ralists at this time tended to read Darwin as a species nominalist (e.g.,
Arthur 1908, 244, who quotes Darwin approvingly; Cowles 1908, 267), in
conformity with the species nominalism of the time (e.g., Morgan 1903,
33; Bessey 1908, 218; Coulter 1908, 272).
On the other side of the coin, keeping to the pre-Modern Synthesis
era, we have the geneticists, who were principally saltationists and tended
to be species nominalists (Mayr 1957a, 4–5, 1982, 540–550). I have found
it impossible, however, to find any of them quote Darwin as a species nom-
inalist, which makes sense since they were anti-selectionists and so therefore
would be unlikely to appeal to Darwin as an authority on the matter.
Turning now to the post-Synthesis period, it is remarkable to find bi-
ologists, philosophers, and historians repeatedly ascribe to Darwin species
nominalism. A good example to begin with is the geneticist J.B.S. Hal-
dane, together with Fisher and Wright one of the three main founders of
16 DARWIN AND THE NATURE OF SPECIES

the Modern Synthesis. In his contribution to a symposium on the species

concept in paleontology, Haldane (1956) states at the outset that “I share
the views of Darwin” (95), which he goes on to elaborate as being that “A
species . . . is a name given to a group of organisms for convenience, and
indeed of necessity” (95), and moreover that “the concept of a species is a
concession to our linguistic habits and neurological mechanisms” (96).
Seeing species in both space and time, he adds that “in a complete pale-
ontology all taxonomic distinctions would be as arbitrary as the division
of a road by milestones” (96). As we shall see in subsequent chapters, how-
ever, this view fails to recognize that Darwin thought of species as primar-
ily horizontal entities and as being delimited in the main by natural
selection, which is a far cry from the subjectivity that Haldane ascribes to
Darwin’s view.
In many ways a more important example is the ornithologist Ernst
Mayr (1957a), according to whom “In Darwin, as the idea of evolution
became firmly fixed in his mind, so grew his conviction that this should
make it impossible to delimit species. He finally regarded species as some-
thing purely arbitrary and subjective” (4; cf. Grant 1957, 58–59, for the
same view expressed in the same volume, and also Mayr 1970, 13, 1976,
259, 1991, 30). What is interesting about Mayr is not that he was using
Darwin as the imprimatur for his own view (Mayr was, after all, a hardcore
species realist), but that he would later blame Darwin’s species nominalism
on Darwin’s association with botanists. In explanation of Darwin’s mature
view of species as “purely arbitrary designations” (269), as opposed to Dar-
win’s earlier view in the 1830s which “was very close to the modern biolog-
ical species concept” (266), Mayr wrote that “His reading as well as his
correspondence indicate that after 1840, and particularly from the 1850s
on, Darwin was increasingly influenced by the botanical literature” (267),
and he goes on to quote William Herbert (a leading English authority on
plant hybridization), for whom he says “the genus was the only ‘natural’ cat-
egory” and of whom he says “Perhaps no other botanist influenced Dar-
win’s thinking more” (268).
There are at least two problems with this view, however. The first one
concerns Herbert in particular. Darwin had indeed read Herbert (his
Amaryllidaceæ is frequently cited in Darwin’s Notebook E), had exchanged
a number of letters with him in mid-1839, and had even visited him once
in September 1845 (Herbert died in 1847). Equally important, in the Ori-
gin Darwin favorably refers to Herbert on the topic of the struggle for ex-
istence among plants (62), and even more favorably on the topic of perfect
fertility in interspecific hybrids in the genera Crinum and Hippeastrum
 A History of Nominalist Interpretation 17

(249–251). For Herbert, hybrids are sometimes very fertile, so that the dis-
tinction between species and varieties has “no real or natural line of differ-
ence” (Burkhardt and Smith 1986, xvii–xviii, 182 n. 1). His species
nominalism, however, if indeed it was such, followed apparently from tak-
ing sterility as the defining criterion of species. It was like Lyell, who
thought that if evolution is true then species must be unreal. But as we shall
see, in spite of Darwin’s acceptance of evolution and of the non-universal-
ity of the sterility of hybrids, he nevertheless thought that species were real
(a view shared, of course, with most biologists today). Moreover, in his cor-
respondence Darwin does not seem particularly impressed by Herbert’s ex-
pertise. For example, in a letter to Hooker (October 28, 1845) written
shortly after visiting Herbert, Darwin remarks that Herbert “knows sur-
prisingly little what others have done on same subjects” (Burkhardt and
Smith 1987, 261).
But even more importantly against Mayr, Darwin repeatedly tells us
that most of his contemporary naturalists were species realists. For exam-
ple, near the beginning of the Origin Darwin tells us that “the view which
most naturalists entertain” is that “each species has been independently cre-
ated” (6). Later in the Origin he gives specific names in the fields of pale-
ontology and geology, stating that “all the most eminent paleontologists”
and “all of our greatest geologists . . . have unanimously, often vehemently,
maintained the immutability of species” (310). But we should not take this
to mean that Darwin did not think the same was true of botanists. As we
have seen earlier in this chapter, Bentham, Hooker, Gray, and even Wat-
son (each of them eminent botanists, with the latter three being Darwin’s
main botanist correspondents) were species realists. Moreover, that the vast
majority of eminent botanists were species realists had been driven home
to Darwin a number of times. For example, Watson (1843) states that there
is a consensus among British botanists that although “genera are allowed to
be purely conventional groups, . . . species are commonly believed to have
a distinct and permanent existence in nature” (613). Moreover there is
Hooker’s letter to Darwin (September 4–9, 1845) which we have seen ear-
lier, in which he wrote “Those who have had most species pass under their
hands as Bentham, Brown, Linnaeus, Decaisne & Miquel, all I believe
argue for the validity of species in nature” (Burkhardt and Smith 1987, 250).
Each member of this list was a first-rate botanist. In sum, all of this adds
credence to Darwin’s remark in his autobiography (1876a), when looking
back at his pre-Origin days, that “I occasionally sounded not a few natural-
ists, and never happened to come across a single one who seemed to doubt
about the permanence of species. Even Lyell and Hooker, . . .” (124).
18 DARWIN AND THE NATURE OF SPECIES

Given the above evidence, it is quite possible that Mayr, then, in blam-
ing the influence of Darwin’s botanist correspondents, was actually pro-
jecting onto history his own problems with botanists, for many modern
botanists have argued that the biological species concept (endorsed more
strongly by Mayr than by anyone else) applies poorly to the world of plants,
a claim that Mayr was long eager to discount and that he attempted to re-
fute by studying a local flora (Mayr 1992). In chapter 6, I shall examine the
views of some of these modern botanist critics of a reproductive criterion
for species.
What is interesting for our purposes here is that one of them, Donald
Levin (1979), in arguing that the biological species concept does not apply
well to plants, argues consequently for species nominalism—“plant species are
utilitarian mental constructs” (381)—and quotes Darwin in support. As he
puts it, “Darwin concurs with Locke” (382; cf. Cowan 1962, 434–435, for
the same equation). John Locke, of course, is famous for arguing in his Essay
Concerning Human Understanding, first published in 1689, that our species
designations are not made by nature but by ourselves, that species words sim-
ply refer to our abstract ideas produced by abstracting what is common from
a number of individuals. Thus, for Locke, “this is a Man, that a Drill [ba-
boon]: And in this, I think, consists the whole business of Genus and Species”
(cf. Stamos 2003, 40–47). In an earlier work (Stamos 1996, 128–129), in
reply to Antony Flew who believed that Darwin never read Locke, I not only
cited a source to the contrary, but quoted an interesting passage from Dar-
win’s Notebook M (84), in which he wrote, “Origin of man now proved.—
Metaphysic must flourish.—He who understands baboon would do more
towards metaphysics than Locke” (Barrett et al. 1987, 285). Although Note-
book M was devoted to the metaphysics of mind, it is quite possible that in
this passage Darwin was referring to Locke’s species nominalism as well as to
his own rejection of that view. What we have to keep in mind is that Dar-
win in his transmutation notebooks, as we have seen, was a species realist.
What we shall see in subsequent chapters is that he never, not even in his
mature period, concurred with Locke.
In the above we have looked at three biologists who read Darwin as a
species nominalist. There are, of course, many more (e.g., Gould 1980,
205–206; Wiley 1981, 41; Howard 1982, 17, 37; Rieppel 1986, 304, 307;
Eldredge 1989, 109–110; Luckow 1995, 590). And among philosophers
the same view naturally persists. For example, the philosopher Elliott Sober
(1993) wrote that Darwin’s book should have been titled “On the Unreal-
ity of Species as Shown by Natural Selection” (143). (For other examples of
 A History of Nominalist Interpretation 19

philosophers who share this view, cf. Hull 1965, 203; Thompson 1989, 8;
Ereshefsky 1992a, 190).
Historians are interesting here in a slightly different way. Alvar Ellegård
(1958, 200), for example, repeats the same view. In fact, the winds of
change did not begin to blow until the biologist Michael Ghiselin (1969)
argued that for Darwin species taxa are real but not the species category,
that Darwin was in one sense a species realist but in another sense a nom-
inalist, so that Darwin did not have a species concept/definition. A num-
ber of years later the philosopher John Beatty (1985), following a suggestion
by Frank Sulloway (1979), added a strategy theory to Ghiselin’s thesis to
explain why Darwin in the Origin would repeatedly define species nomi-
nalistically and yet in fact hold that species taxa are real. Historians have
seemed to simply follow this lead. Jon Hodge (1987), for example, as well
as Gordon McOuatt (1996, 2001), both subscribe to the Ghiselin/Beatty
thesis, while attempting to provide their own twists. I shall return to these
authors in chapter 8, where I develop my own strategy theory. What is in-
teresting to note at this point is that among professional historians, and in-
creasingly among philosophers (e.g., Kitcher 1993, 32 n. 45; Laporte 2004,
192 n. 13; Grene and Depew 2004, 213), the Ghiselin/Beatty thesis has be-
come the received view (cf. chapter 8).7
What I shall attempt to do in the following chapters is to take the now
received view—that Darwin was a species taxa realist but not a species cat-
egory realist—to the next level, that is, to show that he was in fact a species
category realist, that when he looked at taxa he had an implicit species con-
cept that he applied again and again. But that is not all that I shall do.
Before we begin, however, it is important to finish off this chapter with
some strong evidence, direct and indirect, that Darwin’s view in the Origin
and beyond was not that of a species nominalist, in other words that he did
not think of species as akin to constellations, the standard example of nom-
inalism (e.g., Lyell 1832, 19; Darwin 1859, 411), where the individuals are
real but the groupings of them are subjective and arbitrary. A good place to
begin is with Darwin’s reply to Agassiz’s quip that if species are not real then
it makes no sense to say they vary. In a letter to Asa Gray (August 11, 1860)
Darwin wrote “I am surprised that Agassiz did not succeed in writing some-
thing better. How absurd that logical quibble;—‘if species do not exist how
can they vary?’ As if anyone doubted their temporary existence” (Burkhardt
et al. 1993, 317, italics mine). Moreover, in the margin of his copy of Agas-
siz’s review, where Agassiz’s quip is to be found, Darwin wrote “exist only
temporarily” (Burkhardt et al. 1993, 318 n. 4). Temporary existence is, of
20 DARWIN AND THE NATURE OF SPECIES

course, nonetheless real existence, not nominal existence, and in chapter 3

I shall attempt to determine exactly what Darwin meant.8
There is other evidence as well. In the Origin itself, Darwin wrote “I
believe that species come to be tolerably well-defined objects, and do not
at any one period present an inextricable chaos of varying and intermedi-
ate links” (177). This passage, along with many others, will help to estab-
lish an important part of what Darwin thought on the ontology of species,
again as well shall see in chapter 3. Another piece of good evidence is to be
found in Darwin’s letter to Hooker (October 22, 1864), in which he wrote
“The power of remaining for a good long period constant, I look at as the
essence of a species, combined with an appreciable amount of difference;
& no one can say there is not this amount of difference between Primrose
& Cowslip” (Burkhardt et al. 2001, 376). Without the last clause, this pas-
sage has the power to mislead, as, for example, it did Poulton (1903, 91).
Both parts together, however, help to determine a further important feature
of Darwin’s mature species concept, as we shall see in Chapter 5. As a final
piece of evidence that should suffice for the present, Darwin in one of his
articles (1863b) calls it a “great truth,” regardless of evolutionary mecha-
nisms, “that species have descended from other species and have not been
created immutable” (81).
It remains now to determine exactly—inasmuch as that is possible—
what Darwin meant when he wrote of “species.”
 Chapter 2

Taxon, Category, and Laws of Nature

As pointed out earlier, in the literature on the modern species problem

arguably the most fundamental distinction is between species as a taxon
and species as a category. Species taxa are particular species and are given
binomial names such as Tyrannosaurus rex and Homo sapiens. It is species
taxa that evolve, that speciate, that have ranges, that are broad-niched or
narrow-niched, and that become endangered or extinct. The species cate-
gory, on the other hand, is the class of all species taxa.
The species problem is typically phrased in terms of determining the on-
tology of species taxa. But the species category is also a problem. Some con-
ceive of it as an abstract class, a class captured by a definition, the definition
determining (or capturing) membership in the class, such that the class (along
with its definition) stays the same through time as particular species taxa come
and go in terms of existence. Others, however, conceive of the species cate-
gory not as an abstract class but as a concrete set, simply the set of all species
taxa (for examples of both views, cf. Stamos 2003, 150, 258).
This latter view, however, that the species category is a set rather than
a class, has serious problems typically overlooked by its advocates. Granted,
it has greater parsimony, since the species category simply is its member taxa
and nothing more. But this view comes at a great cost. For one thing, it can-
not possibly capture the sense of the species category remaining the same
while its member taxa come and go. If the species category is simply a set,
then as a set it changes every time a species taxon comes into existence or

21
22 DARWIN AND THE NATURE OF SPECIES

goes out of existence. If the species category is viewed as an abstract class,

on the other hand, it does not have this problem. Just as with the category
gold, which stays the same even though particular atoms of gold come and
go in terms of existence, so too, on this view, the species category stays the
same even though particular species taxa come and go in terms of existence.
But there is a further problem for the view that the species category is
a set, namely, that it precludes laws of nature for the species category. It has
become a staple of modern philosophy of biology that there are no laws of
nature for particular species taxa. There are no laws, for example, specifi-
cally for Tyrannosaurus rex. And this absence is exactly what one finds in
modern biological literature. Moreover, this observation makes sense from
a number of key points. Specifically, a species taxon cannot be the subject
of a law of nature because a species taxon gradually evolves. Laws, how-
ever, on the usual view of laws, do not change or evolve. The law of grav-
ity, arguably, has not been changing over time, and the same holds true for
the speed of light in a vacuum (cf. Nagel 1961, 378–380; Armstrong 1983;
Weinert 1995; Stamos 2003, 215–220). But is the species category the sub-
ject of laws of nature? This is indeed a very live issue in modern biology.
The problem is that if the species category is conceived to be a set, then de-
bate over whether the species category is the subject of laws of nature is im-
mediately shut off. If the species category is a set, then it cannot possibly
be the subject of laws of nature because the species category, as a set, is
gradually changing over time, whereas laws of nature do not change over
time. If the species category is conceived to be an abstract class, on the other
hand, then it could possibly be the subject of laws of nature. It need not be
the subject of laws of nature, but the possibility is not precluded, given the
common view of laws as unchanging.
Perhaps the main attraction of conceiving of categories as sets, as noth-
ing more than their members, is that it conforms with a materialist philos-
ophy of nature, according to which what is real is, or is fundamentally,
material. My impression is that many scientists (and not just scientists) are
typically too materialistic to take seriously the view that abstract categories
are part of the fabric of nature. But this should not be considered such a far-
out view (cf. Weinberg 1992, 46). In cosmology, it is debated among physi-
cists whether the so-called cosmic constants (e.g., the law of gravity, or the
strong nuclear force, which holds protons and neutrons together in the nu-
cleus of an atom) are truly universal or are different in different pockets of
the universe (sometimes called “baby universes”). One idea is that the Big
Bang could have produced a different set of cosmic constants. The set that
it did produce obviously allowed for the formation of stars and planets,
 Taxon, Category, and Laws of Nature 23

with life eventually evolving on at least one of the planets. This view can
easily be interpreted in terms of abstract categories built into the fabric of
the universe. The Big Bang, as it actually happened some 15 billion years
ago, resulted in the categories of the chemical elements, each element being
a category. Many of these elements, such as gold, have naturally occurring
physical members, namely, each and every atom that has 79 protons in its
nucleus. Some of the other categories of chemical elements, on the other
hand, have no naturally occurring physical members in the universe. They
are empty categories, their membership being nothing more than a perma-
nent possibility. With some of these categories, of course, humans in recent
years have supplied membership, the highest so far being (I believe) ele-
ment 114. But humans did not create this chemical element/category. The
Big Bang did. Humans, instead, created an atom that simply fit the cate-
gory (even if it was only for a split second). Presumably, because of the
strong nuclear force and the mass of protons and neutrons, there is a phys-
ical limit as to how many protons a nucleus can have, even for a split sec-
ond. Whatever that limit is, it exists, as an abstract reality, out there in
nature, even though humans have not yet discovered it.
Similarly one can think of the species category, as an abstract perma-
nent possibility of nature, coming into existence at the time of the Big
Bang. Richard Dawkins (1983) alludes to this idea when he refers to what
he calls “Universal Darwinism,” the idea that natural selection must be the
main mode of species change wherever in the universe there are species.
Had the Big Bang happened differently, had it produced a different set
of cosmic constants, it is quite possible, even probable, that the species cat-
egory would not exist, that it would not be part of the fabric of the universe,
its having members not a possibility at all. This would occur if the set of
cosmic constants resulting from the Big Bang would not allow for the for-
mation of stars. Without the formation of stars, there could not be planets.
And without planets, life could not evolve and species would be impossible.
Fortunately that is not the universe we have. It is not only a physical
universe, but it also contains abstract categories built into it. Many of these
categories are the chemical elements. The chemical element categories are
strictly essentialistic. Any atom that has 79 protons in its nucleus, for ex-
ample, is an atom of gold, while any atom that does not is not an atom of
gold. Another category is the species category.
The idea that categories are objective, out there in nature, irrespective
of and independent of minds, is of course a controversial one in the history
of thought. There are basically two traditions. Kant thought of categories as
existing only in the mind. On the nature of nature in itself, the noumenal
24 DARWIN AND THE NATURE OF SPECIES

world, he was agnostic. Human perception is conditioned by the categories

of the mind, so that reason can go no further than the phenomenal world,
which is itself the joint product of the mind and nature in itself. This kind
of view, however, is alien to the modern scientific mind, which is typically
realist in orientation. Scientists discover the way things really are, not sim-
ply the ways in which the mind creates the phenomenal world. They dis-
cover real laws, laws in nature, as well as processes and histories and entities
and kinds in nature. In this sense modern scientists are much more in line
with Aristotle. On the topic of categories, Aristotle thought of them as ways
of being, and he specifically listed primary substances (concrete individu-
als) and secondary substances (eidos and genos) among them, examples of
the latter being man, horse, and animal (Categories 1b25–2a19). Moreover,
he explicitly claimed that of all the ways of being (such as property, rela-
tion, quantity, having, and doing) only substances can exist independently
(Physics 185a31–32). (For more on Aristotle on species, cf. Stamos 2003,
102–113; Grene and Depew 2004, 1–34.)
This idea, that there are objective categories in nature, in particular for
species and for other categories, was continued by Carolus Linnaeus. This
self-proclaimed Prince of Botanists, who received his education in Swedish
universities, which were themselves among the last strongholds of Aris-
totelianism in Europe, thought of categories as boxes. As Leikola (1987)
puts it,

His great idea was a rigid and homogeneous categorizing: everything in

nature should and could be fitted into the framework of four basic cate-
gories: class, order, genus and species. Every class consisted of at least one
order, every order of at least one genus, every genus of at least one species.
And vice versa: every species belonged to a definite genus, every genus to
a definite order, every order to a definite class. Nature, the treasury of
Lord, was seen as a cupboard full of departments, and every department
included boxes and these still smaller boxes. A hierarchical order in Na-
ture was not new as such—its idea goes back at least to Aristotle—but the
idea of a uniform hierarchy, up to the point of distorting Nature itself,
was peculiar to Linnaeus. And—natural or not, theoretically feasible or
not—this compartmentalisation of nature was most useful and welcome
for all those naturalists who were busily discovering and describing new
species; here they had an universal framework where to fit and attach
their findings. [46]

Darwin, of course, would come along and kill this view. No longer
could all categories be viewed as boxes, as containers with bridgeless gaps.
 Taxon, Category, and Laws of Nature 25

Because of evolution, categories such as the species category could not be

strictly essentialistic. And this is a theme repeated numerous times in phi-
losophy of biology. Darwin killed essentialism in biology. Species taxa are
not like chemical elements. A physical atom, of course, can change, by ra-
dioactive decay, into an atom of a different element. But one chemical el-
ement does not change or evolve into another. Each element, in fact each
chemical kind (including compounds such as H2O), is a static, essentialis-
tic kind. A species taxon, however, since it gradually evolves, or has a new
species gradually evolve from it, cannot have a strict essence. Of course
clones or monozygotic twins have the same DNA (barring mutations since
they individuated), but for the rest of the organisms that constitute a
species, variation is the norm, both phenotypic and genotypic (cf. Stamos
2003, 119–122, 143).
And indeed one might argue that in biology such non-Aristotelian cat-
egories are now the norm, with concepts such as individual and colony
(slime molds being equivalent to dusk and dawn), living and nonliving
(ditto for viruses), endo- and ectotherm, adult and infant, male and female,
sexual and asexual.
In like manner, that species cannot be essentialistic does not preclude
the possibility of an objective category for species. The species category will
have species in it and the taxa counted as species will reflect the relevant
species concept captured in the particular species definition. The species
category has more species taxa in it according to, for example, the species
concept of Joel Cracraft than the species concept of Ernst Mayr. The ques-
tion then becomes the matter of deciding which species concept deserves
to be the winner, in the sense of best capturing what a species is. Many
think that there should be only one winner, a view known as species
monism, the winner being the universal species concept, while others think
that biology needs a number of different species concepts, so that the species
category should be considered heterogeneous (disjunctive), which is the
view known as species pluralism (cf. Ereshefsky 1992b). This is the species
problem. If we grant that species are real in nature, then what are they?
What is their ontology?
Darwin, as we shall see in subsequent chapters, was not a species plu-
ralist. He did not employ a variety of species concepts. Instead, he repeat-
edly and consistently employed a set of objective criteria for delimiting
species in nature. Moreover, as we shall see in this chapter, Darwin, like
many biologists today, thought that there are laws for the species category.
He therefore would have to count as a species category realist. This helps
to build the case that Darwin was not only a species taxa realist but also a
26 DARWIN AND THE NATURE OF SPECIES

species category realist, with an implicit species concept and definition.

Even though, as we have seen in the previous chapter, many have thought,
and continue to think, that for Darwin species taxa are arbitrary, we shall
see in the following chapters that this view just does not fit the facts. He did
not think of species taxa as arbitrary constructs, like constellations, which
as pointed out in the previous chapter was the classic example of fictional
entities made for convenience and satisfying a convention. Had he done so,
then the species category would likewise in his view be a fiction, made sim-
ply for convenience. But again, as we shall see in later chapters, Darwin ap-
plied repeatedly and consistently a set of objective criteria for delimiting
particular species taxa. Moreover, he added that there are objective laws of
nature for species per se, which by implication means the species category.
It is true that Darwin did not explicitly make a distinction between species
as a taxon and species as a category. Arguably no one did at that time. The
distinction does not seem to have been made until after the Synthesis, one
of the earliest explicit uses being found in Simpson (1961, 19). But this
does not mean that Darwin did not have the distinction in mind. As we
have seen at the beginning of chapter 1, he repeatedly employed a distinc-
tion between species and “the term species.” What we shall see in this chap-
ter is strong evidence that Darwin did not think that the species category
is only in the mind.
Sometimes the distinction between taxon and category is phrased using
the distinction between group and rank. According to Peter Stevens (1994),
“the critical distinction between grouping (forming taxa) and ranking
(placement of those taxa in a hierarchy) is rarely made explicit or even im-
plicit in the historical literature” (10). Using George Bentham as an exam-
ple, Stevens thinks that Bentham was what he (Stevens) calls a “hierarchical
nominalist,” a group but not rank realist. In Bentham’s view, according to
Stevens, “groups from variety upwards were more or less discretely
bounded, . . . but ranking was arbitrary” (176). Interestingly, Stevens places
Darwin in the same category, as a group but not as a hierarchy realist (177).
Translated into the language of taxon and category, this is to say that Dar-
win was a taxon realist but not a category realist. Ghiselin (1969) and Beatty
(1985) also claim this for Darwin. I have my doubts that it is true of Ben-
tham (cf. chapter 8), but I am certain that it is not true of Darwin.
For a start, there is something that the group/rank distinction does not
capture that the taxon/category distinction does. As pointed out earlier, in
some modern views the species category is not just a rank in an existent hi-
erarchy. That view is conformable with the species category merely being
a set. Instead, for many, the species category is something more, namely, a
 Taxon, Category, and Laws of Nature 27

subject of laws of nature, which would make it an abstract category, or to

vary the terminology again, a natural kind.
To get this point across, it is useful to look at the views of Ernst Mayr.
More than anyone else, Mayr has stressed the importance of the distinction
between species as a taxon and species as a category (e.g., Mayr 1970,
13–14, 1982, 253–254, 1988, 321), and he has likewise stressed the impor-
tance of viewing the species category as an abstract class rather than as a set
(e.g., Mayr 1970, 13, 1987, 146, 148). But even he has failed to appreci-
ate the view that the species category should properly be thought of as a nat-
ural kind. In one of his characteristic discussions on what it is to be a
category realist in biology, Mayr (1982) provides two important criteria,
which he states as follows: “(1) Are (most of) the groups (taxa) which we
rank in the higher categories well delimited? and (2) Is it possible to give
an objective (nonarbitrary) definition of such higher categories as genus,
family, or order?” (208). Mayr immediately goes on to argue that the species
category is real but not taxonomic categories above that level. The impor-
tant point, however, is that even for species category realism his two crite-
ria, although necessary, are not really sufficient. To be sufficient, I would
add the following criterion: (3) Is the category spatiotemporally unre-
stricted? This criterion seems to me equally necessary, for surely if one holds
that there are species taxa but they are only to be found on earth, in spite
of holding that life also exists elsewhere in the universe and in an abun-
dance of quantity and complexity, then one could not properly be regarded
as a species category realist. To be such a realist, one would agree with Ed-
ward Wiley (1981)—though one need not agree with his species concept
definition—that “we might expect to find evolutionary species [as he de-
fines the term] anywhere in the universe where organic evolution has oc-
curred” (74). This is in conformity with Dawkins’ Universal Darwinism.
Anything less than this is not a category realism but merely only a group
(set) realism. The difference, logically, is profound. A category realist would
hold that the category (and hence also the definition of the category, pro-
viding that it is correct) remains the same even though members of the cat-
egory come and go in terms of existence. A group realist, on the other hand,
cannot make this claim. As members of a group come and go in terms of
existence, the definition of the group necessarily changes, since a group is
defined by its particular members.
This difference is only further highlighted when we take into account
laws of nature (at least on most positive accounts of laws of nature). Cate-
gories can have their own laws of nature, but groups qua groups cannot. To
think otherwise is to deprive laws of their universal character. Indeed it
28 DARWIN AND THE NATURE OF SPECIES

may be said that any positive claim about one or more laws of nature for a
particular category is an indubitable hallmark of a category realist, species
or otherwise.
As we shall see in the next chapter, Darwin definitely thought that
species taxa, at any one horizontal level, are generally well delimited, so he
meets Mayr’s first criterion. We shall also see, very shortly in this chapter,
that Darwin thought that there are genuine laws of nature for species per
se, so that by my third criterion he thought that the species category is a gen-
uine natural kind. And yet, as we have seen in the previous chapter, he re-
peatedly stated that “the term species” cannot be defined except arbitrarily,
so that, prima facie at least, he does not meet Mayr’s second criterion.
Given all of this, it would appear that Darwin was seriously muddled
on the species question. I do not, however, think that this was the case. In-
stead, in chapters 3 through 6 I shall attempt to prove that, in spite of his
negative claims about defining the term “species,” Darwin repeatedly and
consistently throughout his mature writings employed a set of criteria for
delimiting species. These criteria, moreover, are objective (nonarbitrary)
given—as Darwin, of course, was convinced to be the case—evolution pri-
marily by natural selection. What we shall see is that Darwin did in fact
have a universal species concept, with an implicit definition. Moreover, in
chapter 5 I shall attempt to show that by repeatedly employing this species
concept Darwin considered his practice to be in conformity not only with
the latest taxonomic advances in other areas of science such as crystallog-
raphy, but also with the philosophy of science of his day, which put a high
premium on laws of nature. Why his species concept and definition re-
mained by him implicit, and that he even repeatedly denied that there could
be a definition of the species category, will require an answer in the form
of a strategy theory, given in chapter 8, a strategy theory that not only com-
petes with other strategy theories, such as that of Beatty (1985), but that
employs to the highest degree the principle of charity as well as conformity
to evidence.
For the present, it remains to show that Darwin did indeed think that
there are laws of nature for species per se, and also to determine as best we
can his concept of a law of nature. This, without getting into the details of
his species concept (which will be undertaken in subsequent chapters), will
lay the groundwork for his species category realism.
For the first task, it will prove to be profitable to begin not with Dar-
win but with Michael Ghiselin, who has philosophized extensively on the
relation between species and laws of nature. Ghiselin is, of course, as
pointed out in the Introduction, the father of the modern species-as-
 Taxon, Category, and Laws of Nature 29

individuals view, the view that species taxa are not abstract classes but
concrete individuals, individuated both in space and time. Although part
of the rationale of this ontological shift was to secure the reality of species
taxa against the view of species nominalism (Ghiselin 1966)—since the
reality of abstract classes has long been an issue of debate and the claim of
their unreality a mainstay of nominalism—it would be a serious mistake to
think that for Ghiselin only concrete individuals are real. For Ghiselin the
species category is an abstract class and it is just as real. His justification for
this conclusion is that there are laws of nature for species, not for individ-
ual species taxa but for the species category. (Ghiselin shares the view with
many others that there are no laws of nature for individuals but only for
classes of individuals.) In other words, the species category in Ghiselin’s
view is a natural kind. Indeed he devoted a whole paper to this view and ar-
ticulated what some of these laws (albeit statistical) might be (Ghiselin
1989).1 And as if to quell any doubt on his species category realism, he
reiterated in his book on the metaphysics of species (Ghiselin 1997) that
“just in case anybody wonders, it does not seem reasonable to me to say that
classes exist only in the mind. If the laws of nature refer to classes, that
creates some very sticky problems for anybody committed to such a meta-
physical position” (127, cf. 219–230).
What makes this all the more remarkable is that Ghiselin thought both
that Darwin was a precursor of the species-as-individuals view and that he was
a species category nominalist. With regard to the former, Ghiselin (1969) says
“Darwin had proposed a radical solution to the traditional question of the
‘reality’ of taxonomic groups. What is ‘real’ is the genealogical nexus, and the
groups, or taxa, are chunks, so to speak, of this nexus. As a consequence, it be-
came possible for the nominalist (and Darwin was something of a nominalist)
to look upon taxa not as universals but as particulars, or individuals. . . . Thus
to Darwin, a taxon is real because it is a clade (‘cleft’) or genealogical unit”
(85). This conclusion, claims Ghiselin, “is demonstrable from his actual pro-
cedure in systematic work” (92). It is not important here to repeat or sum-
marize Ghiselin’s analysis of Darwin’s systematic work, for reasons that
should become apparent when I offer my own analysis. What is important
here is to notice that Ghiselin interprets Darwin as a precursor of his own
view, for which Ghiselin is justly famous (Ghiselin 1974, 1987), the view
that species taxa are concrete individuals rather than classes, having both a be-
ginning and an ending in space and time and a horizontal cohesion accom-
plished by sex. What is also important to notice is that much of Ghiselin’s
argument for his interpretation of Darwin as a species category nominalist is
based on his ascription to Darwin of an Aristotelian concept of definition,
30 DARWIN AND THE NATURE OF SPECIES

such that the reality or naturalness of a category requires a discrete essence.

Since in Darwin’s view evolution by natural selection precludes such discrete
essences, because of its gradualism by minute steps, Darwin was therefore, in
Ghiselin’s view, a species (and higher) category nominalist. As Ghiselin
(1969) puts it, Darwin “insisted on Aristotelian definition as a criterion of re-
ality or naturalness. To Darwin, as to many other taxonomists, an inability
to give rigorous definitions for the names of taxonomic groups led to a belief
that somehow such assemblages were artificial” (82).
In chapters 3, 4, and 6, I shall argue in detail against Ghiselin’s view
that Darwin was a precursor of the species-as-individuals view (cf. also Sta-
mos 1998, 1999, 2003, 51–74), also in c hapter 6 I shall reconstruct his
implicit definition of his mature species concept, while in chapter 8 I shall
reply to Ghiselin’s claim that Darwin insisted on an Aristotelian concept of
category. For the present it is important to point out a serious preliminary
problem with Ghiselin’s argument, and it has to do with the species cate-
gory and laws. What is so problematic is not the claim that Darwin was a
species taxa realist (although the claim that Darwin thought of them as in-
dividuals will be rejected), but that Darwin was also a realist about laws of
nature, and not for particular species but for species taxa per se. The prob-
lem is that if this is true, and we shall see shortly that it is, then the only con-
sistent conclusion based on Ghiselin’s own principles is that Darwin was
also a species category realist.
Before we look at those examples, however, it is important to examine
two laws in particular, accepted by the majority of Darwin’s fellow natu-
ralists, which Darwin did in fact reject. His rejection of these laws could eas-
ily lead one to conclude that for Darwin the species category is not real. The
first law is what might be called the law of limited variation. In the Origin
Darwin says “It has often been asserted . . . that the amount of variation
under nature is a strictly limited quantity” (468). Interestingly Lyell (1832)
had listed this in the form of two of his five “laws” for species: “first, that
the organization of individuals is capable of being modified to a limited
extent by the force of external causes; . . . thirdly, that there are fixed lim-
its beyond which the descendants from common parents can never deviate
from a certain type” (23; cf. Darwin 1909, 20 and n. 67). Darwin of course
rejected this law in the Origin, stating that “the assertion is quite incapable
of proof” (468). Of course for Darwin all the evidence he brought to bear
in the Origin for the fact of evolution seemed sufficient to prove that the
law of limited variation must be rejected as a real law.
The second law is closely related to the previous. It is the so-called law
of reversion. According to this law, as Darwin in the Origin briefly puts it,
 Taxon, Category, and Laws of Nature 31

“domestic varieties, when run wild, gradually but certainly revert in char-
acter to their aboriginal stocks” (14). Darwin refers to this law as “well-
known” (25), and Wallace (1859) tells us that it had “great weight with
naturalists, and has led to a very general and somewhat prejudiced belief in
the stability of species” (10). Wallace describes the law as follows: “varieties
produced in a state of domesticity are more or less unstable, and often have
a tendency, if left to themselves, to return to the normal form of the par-
ent species; and this instability is considered to be a distinctive peculiarity
of all varieties, even those occurring among wild animals in a state of na-
ture, and to constitute a provision for preserving unchanged the originally
created distinct species.” He then adds, with regard to the difficulty that the
existence of permanent or true varieties posed for these naturalists, that
“the difficulty is overcome by assuming that such varieties have strict lim-
its, and can never again vary further from the original type, although they
may return to it, which, from the analogy of the domesticated animals, is
considered to be highly probable, if not certainly proved” (10–11). This
law had an impressive pedigree. Linnaeus, for example, exhibited a belief
in the law of reversion, along with the law of limited variation, in his Crit-
ica Botanica, first published in 1737, in which he wrote “every day new
and different florists’ species arise from the true species so-called by
Botanists, and when they have arisen they finally revert to the original
forms. Accordingly to the former have been assigned by Nature fixed lim-
its, beyond which they cannot go: while the latter display without end the
infinite sport of nature” (Ramsbottom 1938, 200 n). Similarly Antoine-
Laurent de Jussieu, in the Introduction to his Genera Plantarum, first pub-
lished in 1789, wrote that a species can be “occasionally subverted for a
while by chance or human industry; that is to say, some individuals may
vary one from another on account of location or climate or disease or cul-
tivation; . . . But these varieties, obeying the law of nature, . . . return to the
primordial species, their character restored, if other factors do not inter-
fere” (Stevens 1994, 340–341). This law, it is to be noted, was thought to
apply only to the species category. Logically there could be no law of rever-
sion for genera and higher taxa, in the minds of those naturalists who held
it, not only because they were species fixists, but also because they were
higher taxa nominalists (cf. Stamos 2005, 83–84). Darwin in the Origin,
of course, readily admitted that “In both varieties and species reversions to
long-lost characters occur” (473), such as stripes on the shoulders and legs
of horses. But his admission only went as far as characters. Of the wide-
spread claim that varieties, if unperturbed, reverted to their primordial state,
Darwin in the Origin wrote “I have in vain endeavored to discover on what
32 DARWIN AND THE NATURE OF SPECIES

decisive facts the above statement has so often and so boldly been made”
(14), and “there is not a shadow of evidence in favour of this view” (15).
And again, the law of reversion for Darwin (as for Wallace and their follow-
ers) would have to be false if varieties are to be incipient species.
Having rejected two of the most commonly accepted laws for the
species category, Darwin in the Origin provides a number of his own laws
for species, although admittedly some of them are not stated explicitly as
laws. What makes them interesting and relevant, nevertheless, is not only
that they are implicitly if not explicitly treated as laws by Darwin, but that
they also figure as laws in modern evolutionary biology. For a start, we
might begin with an apparent anticipation of what is known as the Red
Queen hypothesis, named and enunciated by Leigh Van Valen in the early
1970s as a “new law” (Van Valen 1973), according to which a species must
evolve just to maintain its place in nature and to avoid extinction, since its
physical environment is continually changing as well as the species, due to
competition, with which it interacts. In the Origin Darwin states “Hence
we can see why all the species in the same region do at last, if we look to
wide enough intervals of time, become modified; for those which do not
change will become extinct” (315). The statistical nature of this law is ev-
ident in the fact that there exist “anomalous forms [which] may almost be
called living fossils; they have endured to the present day, from having in-
habited a confined area, and from having thus been exposed to less severe
competition” (107).
Darwin similarly appears to have anticipated what is known as Dollo’s
Law, named after the version of the law enunciated by Louis Dollo in the
late 1880s, according to which the statistical probability is extremely low
that evolution will ever precisely reverse or repeat itself (cf. Gould 1970).
In the Origin Darwin states “When a species has once disappeared from
the face of the earth, we have reason to believe that the same identical form
never reappears” (313), the reason being “for the link of generation has
been broken” (344), which he further elaborates as follows: “We can un-
derstand why a species when once lost should never reappear, even if the
very same conditions of life, organic and inorganic, should recur. For
though the offspring of one species might be adapted (and no doubt this
has occurred in innumerable instances) to fill the exact place of another
species in the economy of nature, and thus supplant it; yet the two forms—
the old and the new—would not be identically the same; for both would
almost certainly inherit different characters from their distant progenitors”
(315). Granted, Darwin in all of this is mainly talking about groups, for he
says, following the second quotation, “When a group has once wholly dis-
 Taxon, Category, and Laws of Nature 33

appeared, it does not reappear, for the link of generation has been broken
(344), and following the third quotation, “Groups of species, that is, gen-
era and families, follow the same general rules in their appearance and dis-
appearance as do single species, . . . A group does not reappear after it has
once disappeared” (316). Nevertheless, that Darwin applies the law to
species is highly significant given, as we have seen and shall see in subse-
quent chapters, that he thought of species as real.
To the above one might add that for Darwin the extinction of species
is also a law of nature. Although this does not come out explicitly in his dis-
cussion on extinction in the Origin (109–111, 127, 172, 317–322, 433,
475), it is nevertheless highly implicit and is made explicit in one of his let-
ters to Andrew Murray (April 28, 1860), in which he states “the extinction
of every form of life in the course of time is a law of nature” (Burkhardt et
al. 1993, 179).
Darwin also thought it a law of nature that each species has a single cen-
ter of origin in both space and time, as opposed to multiple origins, even for
situations (contra, e.g., Louis Agassiz, James Dana, and Alphonse de Can-
dolle) where you have “exceptional cases of the same species, now living at dis-
tant and separate points,” as Darwin put it in the Origin. Given the origin of
species by means of natural selection and the various means of geographic
distribution, the latter discussed by Darwin in Chapter XI of the Origin, Dar-
win thought that single origins constitute a “universal law” (354).
Of course the most important law for Darwin, of which all the above
may be said to be in large part derivative, is natural selection, the main
mechanism for Darwin of adaptation and species evolution. For a start,
right at the end of the Introduction to the Origin Darwin tells us that he
is “convinced that Natural Selection has been the main but not exclusive
means of modification” (6), to which we need to keep in mind the title of
his book: On the Origin of Species by Means of Natural Selection. However,
rather than proffer natural selection in the Origin as a law of nature, Dar-
win calls it a “principle” (61, 80, 95, 127, 475), a “theory” (237, 245, 320,
322, 472, 469), and a “doctrine” (95), as well as a “process” (109, 169,
467) and a “power” (61, 109, 205, 242, 454, 469). Of course the distinc-
tion needs to be made between the principle or theory or doctrine of natural
selection (which is man-made and revisable) and the process or power of
natural selection (which is in nature), the former being better or worse de-
pending on how well it captures (describes) the latter (assuming the latter
exists). The same applies to laws. There needs to be a distinction between
the laws of science (which are man-made and revisable) and the laws of na-
ture (which if they exist are irrespective of the former) (cf. Weinert 1995,
34 DARWIN AND THE NATURE OF SPECIES

4–5). Nevertheless, it remains a fact that nowhere in the Origin does Dar-
win explicitly call natural selection, whether the principle/theory/doctrine
or the process/power, a “law.” Instead, he leaves it implicit, as when, for ex-
ample, he refers to “one general law, leading to the advancement of all or-
ganic beings, namely, multiply, vary, let the strongest live and the weakest
die” (244), or when he refers to “the laws which have governed the produc-
tion of so-called specific forms” (472), or when he says “these elaborately
constructed forms, so different from each other in so complex a manner,
have all been produced by laws acting around us. These laws, taken in the
largest sense, being Growth with Reproduction; Inheritance which is al-
most implied by reproduction; Variability from the indirect and direct ac-
tion of the external conditions of life, and from use and disuse; a Ratio of
Increase so high as to lead to a Struggle for Life, and as a consequence to
Natural Selection, entailing Divergence of Character and the Extinction of
less-improved forms” (489–490). Achingly close, but still no cigar. Equally
tantalizing is that in his correspondence at the time he wrote the Origin
Darwin repeatedly stressed that natural selection is a bona fide vera causa
(true cause). For example, in a letter to Charles Bunbury (February 9, 1860)
he wrote “With respect to Nat. Selection not being a ‘vera causa’; . . . Nat-
ural selection seems to me in so far in itself not to be quite hypothetical, in
as much as if there be variability & a struggle for life, I cannot see how it
can fail to come into play to some extent” (Burkhardt et al. 1993, 76). In
Darwin’s day, the claim that natural selection is a vera causa would be taken
as equivalent to the claim that natural selection is a law, more specifically
a causal law such as gravity. Darwin himself indicates this in one of his let-
ters to Lyell (February 23, 1860), in which he wrote “With respect to
Bronn’s objection that it cannot be shown how life arises, & likewise to
certain extent Asa Gray’s remark that natural selection is not a vera causa.—
I was much interested by finding accidentally in Brewster’s life of Newton,
that Liebnitz objected to the law of gravity, because Newton could not
show what gravity itself is” (Burkhardt et al. 1993, 102; cf. also Darwin’s
letters to Gray, February 24, 1860, and Lyell, June 17, 1860, Burkhardt et
al. 1993, 106–107, 258). Elsewhere in Darwin’s writings, however, well
before and after this period, the label is explicit, as when in the Essay of
1844 (Darwin 1909) he says, when comparing artificial to natural selection,
“Very differently does the natural law of selection act” (95), or when in his
autobiography (1876a) he says “The old argument of design in nature, as
given by Paley, which formerly seemed to me so conclusive, fails, now that
the Law of Natural Selection has been discovered” (87).
 Taxon, Category, and Laws of Nature 35

In chapter 5 we shall examine more clearly in what sense Darwin

thought natural selection a law of nature, as well as what he thought of the
nature of laws in general. Moreover we shall look closely at how he con-
nected natural selection as a vera causa with the versa causa ideal in the phi-
losophy of science of his time. In arguing that natural selection is the vera
causa of species, Darwin was bringing evolutionary biology within the cir-
cle of genuine science, which at the time was circumscribed philosophi-
cally by an emphasis on laws of nature. What others did for crystallography,
for example, Darwin did for biology. At any rate, for the present it is suf-
ficient to conclude, before getting into the specifics of Darwin’s species
concept, that he was indeed a species category realist.
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 Chapter 3

The Horizontal/Vertical Distinction

and the Language Analogy

If one surveys the modern literature on the species problem, one will find
a number of distinctions that together define the debate. Two, of course,
are realism versus nominalism and species as a category versus species as a
taxon, both of which have been dealt with in the previous two chapters.
Two others are species as classes versus species as individuals and species as
process entities versus species as pattern entities, both of which we shall
deal with in this and in later chapters. But there is a further distinction,
which is just as common as the others although it is not always explicit.
This is the temporal distinction between species as horizontal entities ver-
sus species as vertical entities.
This distinction borrows from the now entrenched evolutionary
metaphor of the Tree of Life, instituted, of course, by Darwin (1859,
129–130). Basically, horizontal species exist as one cuts through the Tree
horizontally. The idea here is not necessarily one of species existing at an
instant or microsecond. Instead, the idea usually is that of a time frame
smaller than what is minimally needed for the completion of gradual spe-
ciation. This is the view of species, to name but two examples, contained
in the biological species concept of Mayr (1942, 273, 1970, 12, 1991, 186),
arguably still the dominant species concept in zoology, and in the morpho-
logical species concept, which arguably is the dominant species concept in

37
38 DARWIN AND THE NATURE OF SPECIES

botany (cf. Cronquist 1978; Luckow 1995; McDade 1995). What needs to
be added and emphasized is that the conception of species as primarily hor-
izontal entities need not preclude their existence over vast stretches of time.
Instead, it is only to affirm that their reality is primarily horizontal.
Vertical species, on the other hand, are species conceived either as min-
imal (branchless) branches on the Tree, as one finds in cladistics (e.g., Hen-
nig 1966, 58–66; Ridley 1989, 3), or as differentiated segments of a
minimal branch, as one finds in the theory of punctuated equilibria (e.g.,
Gould 1982, 109–110; Eldredge 1985, 115, 122). Both of these views are
subsumed under the ontological paradigm of species that has reigned in re-
cent decades, namely, the species-as-individuals view first argued by Ghis-
elin (1966, 1974, 1997) and then Hull (1978, 1988). In this view, a species
is not an abstract class but instead a spatiotemporally restricted physical in-
dividual, having a beginning in time (speciation), an ending in time (extinc-
tion), and a definite location in space (Ghiselin and Hull add integration
by sex, but some of their followers do not; cf. the references in Stamos
2003, ch. 4), with membership in a species being a part/whole relation-
ship. Moreover since species, in this view, are genuine individuals very
much like individual organisms, they cannot possibly exist again once they
have gone extinct. As with you or me, once we are dead, we are not just
dead but most sincerely dead, and later copies of us (no matter how perfect)
would not be you or me.
Although some have been reluctant to admit the horizontal/vertical
distinction (e.g., Laporte 1994, 158 n. 49; de Queiroz 1999, 53–54), most
who contribute to the literature on the species problem in some way rec-
ognize it. Interestingly, a number of biologists, although their terminolo-
gies differ, have used the distinction to not only distinguish their species
concepts from others but even to dichotomously classify competing species
concepts. Simpson (1961), for example, characterizes species according to
Mayr’s biological species concept as “contemporaneous” and “horizontal”
(164), while species according to his own evolutionary species concept are
“lineages” (153) with “a long time dimension” (154). Similar to Simpson,
who characterizes the debate as “species at any one time” versus “species
delimited in a certain span of time” (166), Mayr (1988) characterizes the
debate as “whether an evolutionary (vertical) concept is superior to the stan-
dard (horizontal) biological species” (314). In a similar vein, Salthe (1985)
divides modern species concepts into “diachronic” and “synchronic”
(225–226), Endler (1989) uses “contemporaneous versus clade” (627),
while Ridley (1993) uses “non-temporal” and “temporal” (385) and, syn-
onymously, “horizontal” and “vertical” (399).
 The Horizontal/Vertical Distinction and the Language Analogy 39

The distinction also relates to the issue of species realism versus nom-
inalism. Nothing makes this clearer than contrasting two quotations from
Mayr and Eldredge. According to Mayr (1988), “Modern biologists are al-
most unanimously agreed that there are real discontinuities in organic na-
ture, which delimit natural entities that are designated as species” (331).
According to Eldredge (1985), however, “Indeed, at any moment, seem-
ingly at least half the world’s evolutionary biologists are perfectly prepared
to deny that species—any species—even exist” (98). These quotations look
like they are mutually inconsistent, but they are not really. Mayr, the or-
nithologist, is a neontologist, meaning that he is used to dealing with liv-
ing (or recently living) organisms. Not surprisingly his biological species
concept is a horizontal species concept. When he says that most biologists
recognize the reality of species, then, he is referring to species as horizon-
tally conceived, and his claim makes sense given that most modern biolo-
gists are neontologists of one sort or another and that when viewed
horizontally it is fairly easy to argue that species are real (cf. Stamos 2003,
80–97). Eldredge, on the other hand, is a paleontologist, used to dealing
not only with fossilized organisms in horizontal strata but also with their
trends through strata. Not surprisingly his species concept, which is based
on his theory of punctuated equilibria, is a vertical species concept. When
he says that at least half of all biologists think that species are not real, he
is thinking of species as vertical entities. And indeed, although most biol-
ogists are neontologists, most do think that if viewed vertically, species are
unreal (cf. Stamos 2003, 76–79). Determining whether species are (or
should be conceived as) primarily horizontal or vertical entities, therefore,
is a matter of some importance.
At any rate, if Darwin was truly a species realist, and a thoughtful one
at that, we should expect to find evidence that he conceived of species taxa
either as primarily horizontal or as primarily vertical entities. And we do.
But which was it? Ghiselin (1969), as we have seen in the previous chapter,
claimed that Darwin was a precursor of the species-as-individuals view. But
only a precursor. In addition he claimed “Darwin seems not to have fully ap-
preciated the importance of discontinuity, and in this sense he did not em-
brace what we would call a biological species concept. However, he did
recognize that there are species, and he did conceive of them as units or
stages in the evolutionary process. Perhaps the evolutionary species concept
of Simpson is the closest modern parallel to Darwin’s” (101). Malcolm Kot-
tler (1978), on the other hand, while following Ghiselin’s view that Darwin
was a species taxa realist though not a species category realist, thought that
the closest modern parallel was Mayr’s biological species concept. He claims
40 DARWIN AND THE NATURE OF SPECIES

that in the Origin and later writings Darwin “did appreciate the importance
of biological criteria, including reproductive isolation, as opposed to purely
morphological criteria, in the characterization of species and did believe that
species were in some sense real” (292). Moreover he makes much of Darwin
on sibling species in willow wrens (279–280 and n. 9), though at the end of
his paper he waters it down by adding “One might say that for Darwin in
his later works this relationship between interbreeding and transition was not
so clear—closer to ‘less’ than ‘more’ acknowledged” (297).
In chapter 6 I shall argue against Kottler’s claim about Darwin on re-
productive isolation, although in the present chapter we shall see that he
was right if his view was indeed that Darwin (like Mayr) thought of species
as primarily horizontal entities. As a consequence we shall also see that
Ghiselin’s interpretation of Darwin does not withstand scrutiny. If Dar-
win was a precursor of the species-as-individuals view, if his view on species
was closer to Simpson’s view than anyone else, then we should be able to
find strong evidence that Darwin thought of species as primarily vertical en-
tities. But all the evidence, as we shall now see in this chapter, points to the
opposite conclusion, to the conclusion that Darwin thought of species as
primarily horizontal entities. We shall also explore some of the interesting
implications of this.
Before we get into Darwin’s mature writings, however, it is important
to notice that at the beginning of his career as an evolutionist he did seem
to toy with the idea of species as individuals. This was when he held what
today is called his “monadic theory of evolution” (cf. Gruber 1981, 103,
129–149). Three of the essential ideas here were a belief in spontaneous
generation (which of course was fairly widespread), a belief in a fixed du-
ration of time for the existence of each species, and a branching conception
of life. Accordingly in his early notebooks (Barrett et al. 1987) we find en-
tries such as “Tempted to believe animals created for a definite time:—not
extinguished by change of circumstances” (Red Notebook, 129). This was
written in about March 1837 (Barrett et al. 1987, 18), while contemplat-
ing what he believed to be an extinct species of llama. Again in the same
notebook he says “There is no more wonder in extinction of species than
of individuals” (133). Similarly in Notebook B, begun about July 1837,
he wrote “There is nothing stranger in death of species, than individuals.
If we suppose monad definite existence [⫽ definite duration], as we may
suppose is the case, their creation being dependent on definite laws, then
those which have changed most must in each state of existence have short-
est life; Hence shortness of life of Mammalia” (22–23).
 The Horizontal/Vertical Distinction and the Language Analogy 41

The idea that each species has a predetermined lifespan would not be
new to Darwin. The Italian geologist Giovanni Brocchi, for example, sug-
gested in 1814 that every species is constituted in such a way as to have a
predetermined lifespan, so that a species must eventually go extinct even if
its environment remained favorable. Darwin would have learned of Broc-
chi’s theory from his reading of the second volume of Lyell’s Principles of
Geology (1832), which he read while onboard the Beagle. As Lyell put it,
“The death, he [Brocchi] suggested, of a species might depend, like that of
individuals, on certain peculiarities conferred upon them at their birth”
(128), a theory that Lyell immediately went on to reject although he held
Brocchi high in his estimation of him as a geologist.
At any rate, for reasons that are not entirely clear (cf. Gruber 1981,
140–145), Darwin quickly gave up the monadic theory of evolution (eject-
ing spontaneous generation and the aging thesis but retaining the branching
conception of life), which had a lifetime of maybe four months (having been
born just before the summer of 1837 and dying in September of that year).
During that brief period he began to believe that the extinction of a species
is not internally predetermined but contingent on external circumstances. As
he put it in Notebook B, “death of species is a consequence (contrary to what
would appear from America) of non adaptation to circumstances” (38–39).
There was possibly in this change of view some occasional backsliding (cf.
Notebook B, 135; Gruber 1981, 145), but if so they were death spasms. Dar-
win never again entertained the idea of species aging.1
And indeed it is possible, in spite of the analogies made in his monadic
theory of evolution, that Darwin never even thought of species at this time
as individuals, as having a beginning and ending in space and time. Inter-
estingly, however, Darwin’s son Francis has claimed otherwise. Quoting
passages from Notebook B—specifically, “Propagation explains why mod-
ern animals same type as extinct which is law almost proved” (14), “They
die; without they change; like Golden Pippens, it is a generation of species
like generation of individuals” (63), and “If species generate other species,
their race is not utterly cut off” (72)—Francis Darwin (1909) claims that
these quotations “show, I think, that he recognized the two things not
merely as similar but as identical” (xii).
Interestingly, the idea of species as individuals was a bit in the air at the
beginning of the twentieth century. We can also see the idea in the first
book by Francis Darwin’s friend and younger contemporary Julian Hux-
ley (grandson of T.H.). For a start, Huxley (1912) writes of “the species-
individuality of which we are the parts” (24), he makes a distinction between
42 DARWIN AND THE NATURE OF SPECIES

two kinds of individuality, one for organisms and one for species (25), he
claims that “If evolution has taken place, then species are no more constant
or permanent than individuals” (27), and he adds “As individual emerges
from individual along the line of species, so does species emerge from
species along the line of life” (27–28).
In spite of Francis Darwin, however, I suggest that if Darwin did in-
deed think of species as individuals, it was only during the brief stint of his
monadic theory of evolution. Even then, there is strong evidence that he
thought of species primarily as horizontal entities, not as vertical entities as
required by the species-as-individuals view.
In chapter 1 we have seen that in the transmutation notebooks Darwin
employed something very close to the modern biological species concept.
What is interesting is that, in accordance with this view, he thought of
species as primarily horizontal. As we have seen in chapter 1, in one of his
definitions of species he says, in Notebook C, “As species is real thing with
regard to contemporaries—fertility must settle it” (152). This is a later an-
notation to what he wrote on that page, specifically “A species is only fixed
thing with reference to other living being.—one species May have passed
through a thousand changes, keeping distinct from other & if a first & last
individual were put together, they would not according to all analogy breed
together.” Already here we can see, in league with his pre-modern version
of the biological species concept (the modern version of which is adamantly
a horizontal species concept), an emphasis on the primarily horizontal
reality of species (cf. Hodge 1987, 241). What adds to this is an increasing
reliance on a different analogy for species, a reliance away from that of the
individual organism and toward that of the individual language.
The shift to the language analogy is a matter that I shall take up later
in this chapter, including its important philosophical consequences. For
the present, I want to turn to the time of the Origin and show that Darwin
clearly conceived of species as primarily horizontal entities.
We have seen in chapter 1 that in reply to Agassiz’s quip, that if species
are not real they cannot vary, Darwin wrote that species have a “tempo-
rary” existence. But what did he mean by “temporary”? From an evolu-
tionary point of view, whether species are conceived of as primarily
horizontal or vertical, they are in each case “temporary.” I believe the evi-
dence is overwhelming, however, that by “temporary” Darwin had “hori-
zontal” in mind.
To begin, in the Origin Darwin says “To sum up, I believe that species
come to be tolerably well-defined objects, and do not at any one period
present an inextricable chaos of varying and intermediate links” (177;
 The Horizontal/Vertical Distinction and the Language Analogy 43

cf. 203). This is clearly a reference to the horizontal dimension, and equally
clearly its tone is that of species realism.
On the other hand, only two pages later, Darwin says “Lastly, looking
not at any one time, but to all time, if my theory be true, numberless in-
termediate varieties, linking most closely all the species of the same group
together, must assuredly have existed” (179). Here we see that even though
Darwin is dealing with a long time dimension, there is no talk of species
vertically conceived, only of species linked by intermediate varieties. In fact
Darwin used a number of phrases in addition to “intermediate varieties,”
namely, “transitional forms,” “transitional varieties,” and “intermediate
links,” all meaning the same thing. For example, in a passage about the fos-
sil record, Darwin claims that all throughout biological history there must
have been “an infinite number of those fine transitional forms, which on
my theory assuredly have connected all the past and present species of the
same group into one long and branching chain of life” (301). In a section
devoted to “the absence or rarity of transitional varieties,” Darwin says “ex-
tinction and Natural Selection will, as we have seen, go hand in hand.
Hence, if we look at each species as descended from some unknown form,
both the parent and all the transitional varieties will generally have been ex-
terminated by the very process of formation and perfection of the new
form” (172). In the passage following this, Darwin uses the phrase “innu-
merable transitional forms.” Finally, this time in his concluding chapter, on
the imperfection of the geological record, Darwin says “The number of
specimens in all our museums is absolutely as nothing compared with the
countless generations of countless species which certainly have existed. We
should not be able to recognise a species as the parent of any one or more
species if we were to examine them ever so closely, unless we likewise pos-
sessed many of the intermediate links between their past or parent and pre-
sent states; and these many links we could hardly ever expect to discover,
owing to the imperfection of the geological record” (464).
In all the above, species are clearly horizontal species. We do not find
the idea of species commonly extending through a long stretch of time, let
alone changing—and least of all changing radically—and yet remaining
the same species. In fact, nowhere in Darwin’s writings do we find this.
Darwin’s primary conception of species was horizontal.
Some comments are necessary, however, before we continue with the ev-
idence. First, by “well-defined . . . at any one period” Darwin did not mean
that he thought species are well-defined in the sense of essential characters
as captured in an Aristotelian definition. This meaning is eliminated by the
fact that Darwin conceived of species evolution as minutely gradual, which
44 DARWIN AND THE NATURE OF SPECIES

makes it impossible for a species to have an essence (cf. Stamos 2003, 122).
Instead, by “well-defined” Darwin was simply following common usage, in
which the phrase means sufficiently clear or well marked out and thereby al-
lowing for delineation and description. We can see this usage especially well
near the end of the Origin, wherein Darwin says “Systematists will only have
to decide (not that this will be easy) whether any form be sufficiently con-
stant and distinct from other forms, to be capable of definition; and if de-
finable, whether the differences be sufficiently important to deserve a specific
name” (484; cf. also 171, 174, 432, 1871 I, 226–227). Moreover the phrase
“well-defined” for species seems to be used synonymously by Darwin with
“good and distinct species” (259) and with “good and true species” (47).
This usage is common even today. For example, Mayr and Short (1970)
state that “The designation ‘good species’ refers to those that are clearly de-
limited from other species and whose recognition is not controversial among
specialists” (1). Moreover there was nothing idiosyncratic in Darwin’s use of
the term “definition” for “description” even in his own time. Hugh Strick-
land (1837), for example, in his preliminary suggestions for rules for zoolog-
ical nomenclature, wrote “A name may be expunged which has never been
clearly defined. . . . Many collectors of shells and fossils are in the habit of
labelling those species which they do not find described, with names of their
own invention; but, unless they publish descriptions of these new species,
they cannot expect these names to stand” (174). Whether by Darwin, by
some his contemporaries, or even by modern biologists, then, there is noth-
ing in the above phrases to imply essentialism.
Second, we should not suppose that by “at any one time” Darwin
meant an instant or microsecond for horizontal species. In fact in the pas-
sage above where Darwin affirms the horizontal reality for species, he uses
the words “at any one period.” Darwin clearly uses both phrases with the
same reference. Moreover when we look at Darwin’s comparison of species
with languages, which he conceives of both as primarily horizontal, the
only plausible interpretation is that Darwin did not conceive of the hori-
zontal dimension as an instant, but rather as a period of time which, at its
maximum, is too short, in the case of species, for the gradual evolution of
one species into another. Today this is thought to be roughly 3,000 years
(cf. Stamos 2002, 177). In the case of languages, of course, where language
is defined traditionally as a system of communication, the maximum time
period for the horizontal dimension is going to be much shorter.
Third and finally, we have to keep in mind that, among his few corre-
spondents who were evolutionists, one in particular seemed to conceive of
species as primarily horizontal. As we have seen in chapter 1 (note 4), the
 The Horizontal/Vertical Distinction and the Language Analogy 45

botanist H.C. Watson, in a letter to Darwin (November 8, 1855), wrote

“I must confess a pretty strong bias towards the view, that species are not
immutably distinct;—altho’ in our time-narrowed observation of the indi-
viduals they seem to be so” (Burkhardt and Smith 1989, 499). Similarly, a
little later in a letter to Darwin (December 20, 1857) he wrote “I cannot
find the proof of species being definite & immutable, whatever they may
seem to be at any one time & spot” (Burkhardt and Smith 1990, 511).
But why would Darwin think that “at any one period” species in gen-
eral are “tolerably well-defined objects”? Immediately following the passage
in the Origin in which this quotation is found, he gives us three reasons.
The first is that variation is a very slow process and natural selection requires
uninhabited niches, the second is that intermediate varieties will often be ex-
terminated by natural selection—this connects with his view that “extinc-
tion and natural selection . . . go hand in hand” (172)—and the third is that
intermediate varieties, because of their smaller numbers, will often be exter-
minated by accidents (presumably such as fire etc.) (177–178).
To these three reasons given by Darwin in the Origin, we also have to
add that he thought that interspecific hybridization is comparatively rare
(ch. 8). A further consideration is that Darwin thought that most species
have a fairly well-defined range. As Darwin put it, “In looking at species as
they are now distributed over a wide area, we generally find them tolerably
numerous over a large territory, then become somewhat abruptly rarer and
rarer on the confines, and finally disappearing. Hence the neutral territory
between two representative species is generally narrow in comparison with
the territory proper to each” (174). This view, incidentally, accords with
what modern biologists write (e.g., Mayr 1970, 301–302; Futuyma 1986,
ch. 13). Of further importance is Darwin’s pluralism with regard to speci-
ation. Darwin allowed for both allopatric (geographic isolation) and sym-
patric (principle of divergence) modes of speciation (more on this below).
Both modes entail branching speciation and his metaphor of the “great
Tree of Life” (130). As he says in the Origin, “In a tree we can specify this
or that branch, though at the actual fork the two unite and blend together”
(432). It is only the forks, then, at any one horizontal dimension, which
present messy situations, situations without good and distinct species. But
since the forks, at any one horizontal dimension, are not the norm, most
species at that dimension are going to be good and distinct species.
It is important here to further understand Darwin’s theory of specia-
tion. According to the influential theory of Mayr (e.g., 1982, 410–417), al-
though Darwin was an early adherent of speciation by geographic
(allopatric) isolation, by the time he wrote the Origin he no longer had a
46 DARWIN AND THE NATURE OF SPECIES

clear theory on speciation, so that his book was misnamed. A more likely
scenario, it seems to me, is that Darwin evolved into a limited speciation
pluralist (limited by his confinement to gradualism),2 accepting by the time
of the Origin both nonbranching speciation or anagenesis (cf. 119–120)
and branching speciation or cladogenesis, the latter including both sym-
patric speciation (in accordance with his principle of divergence; cf.
105–106, 112, 115–116) and allopatric speciation (cf. 399–404). Of
course, that he considered sympatric speciation the most important (cf.
105) does not take away from this pluralism, which, by the way, seems to
be a growing view in modern speciation studies (cf. Otte and Endler 1989;
Howard and Berlocher 1998).
Another objection might be based on Darwin’s work on barnacles. Dar-
win had spent eight years anatomizing and monographing barnacles from
1846 to 1854, both living and fossil. Did not their variability prove to him
that species (or rather what were called “species”) are too variable, even at any
one horizontal level, to be real, so that (under his breath at least while writ-
ing his monographs) he gave up species as real entities? The answer is no.
Granted, Darwin often complained of their variability. For example, in a
letter to Hooker (June 13, 1850) he wrote “I have been struck (& probably
unfairly from the class) with the variability of every part in some slight de-
gree of every species: . . . & consequently that the diagnosis of species from
minute differences is always dangerous. . . . Systematic work wd be easy were
it not for this confounded variation, which, however, is pleasant to me as a
speculatist though odious to me as a systematist” (Burkhardt and Smith
1988, 344). Moreover there is that often-quoted passage in another of Dar-
win’s letters to Hooker (September 25, 1853) in which he wrote, in refer-
ence to his barnacle work, “After describing a set of forms, as distinct species,
tearing up my M.S., & making them one species; tearing that up & mak-
ing them separate, & then making them one again (which has happened to
me) I have gnashed my teeth, cursed species, & asked what sin I had com-
mitted to be so punished” (Burkhardt and Smith 1989, 156).3
In spite of this “confounded variation,” however, Darwin nevertheless
found that barnacles on the whole, when studied anatomically, presented
good species. In a letter to Lyell (September 2, 1849), for example, Darwin
remarked “I sometimes after being a whole week employed & having de-
scribed, perhaps only 2 species agree mentally with Ld. Stanhope that it is all
fiddle-faddle” (Burkhardt and Smith 1988, 252, italics mine). Instead, the
“chaos” that Darwin so often complained about with regard to barnacle sys-
tematics referred only to the “book” species of his contemporaries, which
were based only on external characters and which often involved a nomen-
 The Horizontal/Vertical Distinction and the Language Analogy 47

clature with many synonyms. For example, in a letter to Hugh Strickland

(February 4, 1849), on the topic of the rules of nomenclature, Darwin wrote
“In not one large genus of Cirripedia has any one species been correctly de-
fined: . . . Literally not one species is properly defined: not one naturalist has
ever taken the trouble to open the shell of any species to describe it scientif-
ically, & yet all the genera have ½ a dozen synonyms. . . . The subject is
heart-breaking” (Burkhardt and Smith 1988, 206–207). Similarly in a let-
ter to Johannes Müller (February 10, 1849) he wrote “I find that the
anatomy of the Cirripedia has been most imperfectly done; nearly all the
most striking features in their organization having been overlooked.—Their
classification is likewise a perfect chaos, as must be the case until the whole
body of every species be examined, as I am now doing” (Burkhardt and
Smith 1988, 213).
And in fact when we look specifically at Darwin’s work on living bar-
nacle species we find that he claims that most, when properly anatomized,
are good species. For example, in his preface to his second volume, on the
Lepadidae (Darwin 1851b), he wrote that “no doubt they are subject to
considerable variation, and as long as the internal surfaces of the valves and
all the organs of the animal’s body, are passed over as unimportant, there
will occasionally be some difficulty in the identification of the several forms,
and still more in settling the limits of the variability of the species” (xi).
Even when he dealt with the most problematic cases of variation, Darwin
still maintained that, when properly anatomized, most species will turn out
to be good species. For example, in his second volume on living species, on
the Balanidae (Darwin 1854b), in his introductory discussion to the genus
Balanus, which for Darwin was by far the largest genus and which he con-
sidered to have “an especial amount of variation” (156), he remarked that
“Notwithstanding the difficulties now enumerated, I hope that, owing to
having examined a vast number of specimens of the most varying species,
I have not fallen into many errors” (190). Surely Darwin would not have
felt this way if he thought that most of the species he carefully anatomized
were not after all good species. Moreover one has only to read his descrip-
tions of his 45 species of Balanus (194–302), notwithstanding his addi-
tional 9 species of the subgenus Acasta (302–321), to see his confidence in
regarding most of the species as sufficiently good and distinct.
Of the latter subgenus, granted, Darwin says the first four species caused
him “much doubt and trouble” (308), while with species 5 and 6, though
very distinct from other species, he also expresses “some hesitation” (314) in
making them specifically distinct. With species 7 he doesn’t say, while
species 8 is “perfectly distinct” (318) and species 9 is “very distinct” (319).
48 DARWIN AND THE NATURE OF SPECIES

With Balanus the situation is much better, in spite of its variability.

Granted, species 1 and 19 are described as “the most difficult and variable
in the genus” (196–197) and as causing Darwin “utter despair” (243).
Species 18 is described as causing Darwin “much trouble” (235). With
species 34, 35, and 36, taken together, Darwin feels “somewhat doubtful”
(284). Similarly with species 43 and 44 Darwin has “some doubt whether
they ought to be specifically separated” (298). With species 4 and 8 he ex-
presses a little difficulty. Species 6, however, “can easily be distinguished”
(213). Species 10, 11, 12, and 13 are described as “quite distinct species”
(218). Species 17 is described as a “well-marked species” (232). Species 20
is described as having “amply diagnostic characters” (247). With species
21 and 22 Darwin says “I entertain no doubt whatever about the distinct-
ness of the two species” (250). Species 23 is described as a “very distinct
species” (253), as is species 30 (274) and 38 (288). Species 24 is described
as differing “distinctly” from species 27 (254). Species 25 is described as
being “very distinct from every other” (259). Species 33 is described as
being a “distinct and well-defined species” (281). Species 27 and 29, al-
though “confounded” in most collections (261), are found by Darwin,
when “disarticulated,” to be “at once distinguished,” which is only further
confirmed by Darwin’s observation that they inhabit different depths of
water (271). Similarly species 32, when “disarticulated,” “cannot be con-
founded with any other” (278). Similarly as well, when species 45 is “dis-
articulated” the result allows “of no mistake of the two species” (302), the
other species being species 27. Species 37 is described as being “so peculiar”
(285). Species 41 is described as being a “strongly characterized species”
(294). Species 42 presents “well defined distinctions” (295). Finally, with
species 2, 3, 5, 7, 9, 15, 16, 26, 28, 31, 39, and 40, it is implied that they
are good species.
To all of the above, one needs to keep in mind that Acasta and Balanus
were the most difficult groups.
So we can see that in the Origin Darwin was not disingenuous when
he claimed that horizontally most species are good species. Moreover it is
abundantly evident that the horizontal dimension was for him the primary
dimension for species reality. But this is not to deny that Darwin allowed
for a species, horizontally conceived, to exist vertically over a vast stretch of
time. For a start, he coined the term “living fossils.” These “anomalous
forms,” he says, which he later calls “species and groups of species” (486),
“have endured to the present day,” and the reason he gives for their exis-
tence, along with the name, implies that they have remained virtually the
same over vast stretches of time to the present, the reason being “from hav-
 The Horizontal/Vertical Distinction and the Language Analogy 49

ing inhabited a confined area, and from having thus been exposed to less
severe competition” (107). He also tells us that “species very rarely endure
for more than one geological period” (153). Moreover when we look at
Darwin’s work on fossil barnacles (Darwin 1851a, 1854a), in spite of the
problem that the soft parts were generally not preserved in the fossil record,
we find that in a number of cases Darwin classified fossil species as conspe-
cific with living species. In the case of Balanus (1854a, 13–33), which as we
saw was divided by Darwin into 45 living species, Darwin recognized
11 fossil species, each of which he identified with a living species, and in
each case because the diagnosis seemed to be the same.
All of this will make more sense when we examine what Darwin
thought was the closest analog of species, namely, natural languages. But
what needs to be reiterated at this point is that in allowing a species to
have a vertical extension this did not mean that Darwin thought that
species are primarily vertical. We have already seen much evidence that
Darwin thought of species as primarily horizontal. Later in this chapter I
shall argue that Darwin was on firm logical ground. What I want to do at
this point is provide a further piece of evidence that for Darwin the real-
ity of species is primarily horizontal. This further piece of evidence comes
from Darwin’s discussion on his one and only diagram in the Origin (the
diagram is reproduced on the next page). Interestingly, according to Kevin
de Queroz (1999)
An early version, or at least a precursor, of the general lineage concept can
be found in Darwin’s (1859) Origin of Species. In the only illustration in
that book, Darwin represented species as dashed and dotted lines, or col-
lections of such lines, forming the branches of what would now be called
a phylogenetic tree. In the accompanying text, he used the term species
more or less interchangeably with the term lines of descent. [76]

In this passage de Queroz reads Darwin as having a vertical species con-

cept, conceiving of species as branches in the phylogenetic tree of life. But
this reading does not at all survive close scrutiny of what Darwin actually
said in relation to his diagram. Perhaps de Queroz was misled by the con-
cluding section of the chapter containing the diagram, in which Darwin
states “The affinities of all the beings of the same class have sometimes been
represented by a great tree. I believe this simile largely speaks the truth.
The green and budding twigs may represent existing species; and those pro-
duced each former year may represent the long succession of extinct species”
(129). But even here species are compared to buds, not branches. Instead
a branch represents a “long succession of extinct species.”
 The Horizontal/Vertical Distinction and the Language Analogy 51

This interpretation only becomes stronger once we turn to Darwin’s

discussion on his diagram. It is interesting, and surely not insignificant,
that F14—descended from species F, says Darwin, 14,000 generations later,
or better yet 114,000 generations later, “either unaltered or altered only in
a slight degree”—is twice called by Darwin a “new species” (117, 124).
Certainly no one motivated by a primarily vertical species concept would
call F14 a new species in such a diagram. Simpson (1961) certainly did not
do this sort of thing in his own diagrams (164, 168). Neither would
cladists, since they conceive of species as branches in the phylogenetic tree
of life, and species conceived as such are allowed to undergo unlimited
change and remain numerically the same just so long as they don’t pro-
duce branches (cf. Hennig 1966, 58; Ridley 1989, 10). (I should add that
the line from species F to F14 in Darwin’s diagram is a straight line with no
branching; the same is true of the line from E to E10.) Nor is there any ev-
idence that Darwin thought of species in the way adherents of the modern
theory of punctuated equilibria do, as vertical entities characterized mainly
by stasis with rapid evolution (which equals for them speciation) delimit-
ing the temporal ends (cf. Eldredge 1985, 115, 122). Even though, as we
have already seen, Darwin did countenance stasis, he did not use it to con-
ceive of species, which would follow alone from his belief in the relative
infrequency of living fossils.
Although Darwin’s diagram indicates yet once again that for him the
horizontal dimension has priority over the vertical dimension, it should
not be taken to necessarily preclude the vertical dimension, as indicated
by what he says about living fossils. Moreover, it should be noticed that
the F in F14 is capitalized (unlike, e.g., a14 or f 14 or m14), which alone
suggests that F14 alone on that horizontal line may be thought of as a ver-
tical species numerically identical with its corresponding species on the
horizontal base line, namely, species F (and so also for E10 with E), un-
like the lines for the lowercase letters which seem to indicate only lin-
eages and not species.
To understand Darwin’s thinking further on this matter it will help to
turn to his main analogy for species, namely, the language analogy. We
have seen near the beginning of this chapter that Darwin held a monadic
theory of evolution very briefly at the beginning of his career as an evolu-
tionist. In this theory a species is a vertical entity highly analogous to an in-
dividual organism. Darwin quickly gave up that theory and the organism
analogy, however, replacing it with a theory of contingent evolution by
natural selection and the language analogy for species. This analogy is
highly significant and philosophically of the utmost importance.
52 DARWIN AND THE NATURE OF SPECIES

Beginning with Darwin’s early notebooks (Barrett et al. 1987), in

Notebook B he wrote “As man has had not time to form good species, so
cannot the domesticated animals with him!—Modern origin shown by only
one species, far more than by non-embedment of remains—?agrees with
non-blending of languages?” (244). In Notebook N he wrote “many
learned men seem to consider there is good evidence in the structure of
language, that it was progressively formed” (65). In Notebook OUN he
wrote “At least it appears all speculations of the origin of language.—must
presume it originates slowly—if these speculations are utterly valueless—
then argument fails—if they have, then language was progressive.—We
cannot doubt that language is an altering element, we see words invented—
we see their origin in names of People.—Sound of words—argument of
original formation.—declension &c often show traces of origin” (5). He
also wrote “H. Tooke has shown one chief object of language is prompt-
ness of consequence hence languages become corrupt, & whole classes of
words are abbreviations he thus derives from nouns & verbs—so that much
of EVERY language shows traces of anterior state??” (13). In these passages
it is not always entirely clear whether Darwin is commenting on the evo-
lution of the human language ability or on the evolution of languages. In-
deed as Stephen Alter (1999) puts it, “These passages start out as
speculations regarding the origin of speech, yet they quickly shade into re-
flections on the purely analogic similitude between languages and species”
(16). Although the two ideas are commingled, they nevertheless do show
that Darwin was starting to think of languages as analogous to species.
What is especially interesting is what he says in the last quotation above.
John Horne Tooke was a British linguistic philosopher who argued that
French, Italian, Anglo-Saxon, Dutch, German, Danish, Swedish, and
English “are little more than different dialects of one and the same lan-
guage” (Barrett et al. 1987, 603 n. 13–1). In capitalizing the word
“EVERY” in the quotation above, we can see that Darwin, against Tooke,
held a commonsense view of language, which may legitimately shed light
on what Darwin meant by the word “language” in the previous passages
(and possibly also in later passages) where he compares species to languages.
In the Sketch of 1842 there are allusions to the language analogy as used
by Lyell for geology, which I shall return to below. Of far more interest is
the Essay of 1844 (Darwin 1909), in which Darwin compares the impor-
tance of rudimentary organs for determining classification of species in the
natural system with rudimentary structures in languages. As Darwin put
it, “In the same manner as during changes of pronunciation certain letters
in a word may become useless in pronouncing it, but yet may aid us in
 The Horizontal/Vertical Distinction and the Language Analogy 53

searching for its derivation, so we can see that rudimentary organs, no

longer useful to the individual, may be of high importance in ascertaining
its descent, that is, its true classification in the natural system” (235).
This is an idea that Darwin would repeat in the Origin (455). There,
however, we find more comparisons between species and languages. We
are told, for example, that “a breed, like a dialect of language, can hardly
be said to have a definite origin” (40). Equally important, in arguing that
the only natural classification of species is genealogical, Darwin proceeds to
“illustrate” his view “by taking the case of languages,” of which he con-
cludes “the proper or even only possible arrangement would still be ge-
nealogical; and this would be strictly natural, as it would connect together
all languages, extinct and modern, by the closest affinities, and would give
the filiation and origin of each tongue” (422–423).
But it is in the Descent of Man (1871 I) that we find the most striking
passages. Therein Darwin writes:

The formation of different languages and of distinct species, and the

proofs that both have been developed through a gradual process, are cu-
riously the same. . . . The frequent presence of rudiments, both in lan-
guages and in species, is still more remarkable. . . . Languages, like organic
beings, can be classed in groups under groups; and they can be classed ei-
ther naturally according to descent, or artificially by other characters.
Dominant languages and dialects spread widely and lead to the extinction
of other tongues. . . . The same language never has two birth places. Dis-
tinct languages may be crossed or blended together. We see variability in
every tongue, and new words are continually cropping up; but as there is
a limit to the powers of memory, single words, like whole languages, grad-
ually become extinct. . . . The survival or preservation of certain favoured
words in the struggle for existence is natural selection. [59–61]

Much of what is interesting about Darwin’s language analogy is his

sources. In the final chapter of the first volume of his Principles of Geology,
Lyell (1830, 461–462) compares change in geologic epochs to change in a
written language over a period of 1,000 years. In both cases, he says, if one
knows only one-tenth of the objects of study as they exist today, one can-
not possibly hope to have a good understanding of the objects of study in
the distant past. In the third chapter of the third volume of his Principles,
Lyell (1833, 33–34) supposes that if an archaeologist were to find two
buried cities at the foot of Mount Vesuvius, one on top of the other, and
found the inscriptions at the lower city to be Greek and the inscriptions at
the higher city to be Roman, he would infer incorrectly if he thought the
54 DARWIN AND THE NATURE OF SPECIES

language change from Greek to Roman was abrupt. And if he later found
a third city in between the two, with inscriptions in Italian, he would then
see his error, and infer properly that the language change from the earlier
population to the later had been “very gradual.”
Darwin had read all three volumes of Lyell’s Principles assiduously dur-
ing his Beagle voyage (Burkhardt and Smith 1985, 562), and he would soon
after consider himself the heir to Lyell’s uniformitarianism, as applied to the
evolution of species. He was also familiar with a letter that John Herschel
had written to Lyell on February 20, 1836 (extracted in Charles Babbage’s
The Ninth Bridgewater Treatise, published in 1837), in which Herschel wrote
“Words are to the Anthropologist what rolled pebbles are to the Geologist—
Battered relics of past ages often containing within them indelible records
capable of intelligent interpretation,” and in which he called the origin of
species the “mystery of mysteries” (cf. Darwin 1859, 1; Burkhardt and Smith
1986, 8–9 and n. 5; Desmond and Moore 1992, 214–215).
Many years later, in his The Antiquity of Man, Lyell (1863) devoted a
whole chapter (ch. 23) to the comparison between language evolution and
the theory of species evolution and made a number of remarkable compar-
isons. Ontologically speaking, the most important of his comparisons are
that “the learned are not agreed as to what constitutes a language as distinct
from a dialect” (458), that practically speaking “two languages should be re-
garded as distinct whenever the speakers of them are unable to converse to-
gether” (458), which he compares to the sterility barrier, that there is a “real
question . . . whether there are any limits to . . . variability” (462), that an
important area of inquiry is over language change, in particular “what are
the laws which govern not only the invention, but also the ‘selection’ of
some of these words or idioms, giving them currency in preference to oth-
ers” (463); that “dialects . . . may be regarded as . . . ‘incipient languages’”
(464), that “we find in them some internal evidence of successive additions
by the invention of new words or the modification of old ones” (465), that
“No one of them can have had two birth places” (465), that “They may die
out very gradually in consequence of transmutation, or abruptly by the ex-
termination of the last surviving representatives” (467), and that “a lan-
guage which has once died out can never be revived, since the assemblage
of conditions can never be restored” (467). Some of these analogies are
straight from Darwin, as we have seen, but they surely prompted Darwin
to strengthen the analogy even further in his Descent, again as we have seen.
Interesting is what Darwin wrote in 1863 in a letter to Gray (February 23),
that “Lyell was pleased, when I told him lately that you thought that lan-
guage might be used as excellent illustration of derivation of species; you
 The Horizontal/Vertical Distinction and the Language Analogy 55

will see that he has admirable chapter on this” (Burkhardt et al. 1999, 166),
and what he wrote to Lyell (March 6) that “No praise can be too strong,
in my opinion, on that inimitable chapter on language in comparison with
species” (207).
Of greater interest is what was going on in the field of linguistics dur-
ing Darwin’s lifetime (the term “linguistics” was introduced to England by
Whewell 1837). While Darwin was developing his evolutionary views, and
most biologists thought of species as fixed (allowing at most for varieties),
virtually all linguists thought of languages as evolutionary. Interestingly,
what became prominent near the beginning of the ninteenth century, par-
ticularly in Germany, was the organism metaphor. This vertical conception
of languages occurred hand in hand with the concern for determining par-
ticular paths of language evolution and with reconstructing ancestral lan-
guages, most of all, Proto-Indo-European. Notable in this regard were
Friedrich von Schlegel and Jacob Grimm, both of whom in the first decade
of the ninteenth century claimed that the discipline closest to comparative
grammar was comparative anatomy. Of much greater importance was Franz
Bopp, highly influential as the author of the first comparative grammar of
the Indo-European languages. In 1827 Bopp wrote “Languages must be
taken as organic natural bodies which form themselves according to defi-
nite laws, develop carrying in themselves an internal life principle, and grad-
ually die” (Davies 1987, 84). Bopp, however, arguably did not take the
organism metaphor literally (cf. Wells 1987, 56). Nevertheless, he seems to
have thought, as Davies (1987) points out, that “a language can change its
grammatical ‘type’ while remaining in some sense the ‘same’ language” (91).
At any rate, the greatest exponent of the organism metaphor was August
Schleicher, regarded by many as the most influential figure in ninteenth cen-
tury linguistics (cf. Taub 1993, 175–176 and n. 13). As early as 1848 Schle-
icher began to argue that individual languages really are organisms, imbued
with life in a literal sense so that the scientific study of them is a biological
science in its own right, up there with zoology, botany, and paleontology.
In 1863, in finding further support for his views upon reading Heinrich
Bronn’s German translation of Darwin’s Origin, he wrote “Languages are
natural organisms that arose independently of human volition, grew in ac-
cordance with definite laws, and developed and in turn grew old and died.
They are also characterized by the set of phenomena we are accustomed to
understand by the term ‘life.’ Accordingly, glottics, the science of language,
is a natural science; its method is largely the same as that of the other nat-
ural sciences. This is why the study of Darwin’s book . . . was bound to seem
quite close to my area” (Percival 1987, 8). Equally important, it was
56 DARWIN AND THE NATURE OF SPECIES

Schleicher who made famous the use of branching tree diagrams to illustrate
genealogical relationships in language evolution. Moreover, he argued that
any study of language that was not evolutionary was not scientific (cf. Taub
1993, 188).
As Alter (1999) points out, “The ‘new philology’ of Bopp and Grimm
began seeping into England in the early 1830s, in part through the efforts
of Darwin’s cousin and brother-in-law Hensleigh Wedgwood” (11). One
of my main criticisms of Alter’s otherwise excellent book (Stamos 2000) is
that he doesn’t go far enough in telling us what Darwin got from Wedg-
wood. Granted, Darwin had many discussions with Wedgwood during his
stay in London following his Beagle voyage, Wedgwood’s main focus in his
own work was on etymology and the natural origin of language (which
would of course have been of interest to Darwin), Wedgwood had written
an extensive review (Wedgwood 1833) of Grimm’s Deutsche Grammatik,
and they kept up with correspondence and visits after Darwin moved to
Down. And yet we are not told anything about the organism metaphor.
Gillian Beer (1989) provides much more on Wedgwood’s review of
Grimm, and tells us that “It is likely that Darwin read his cousin’s only
published article, given his current interest in the subject and the close ties
between them” (158). And yet, in spite of the pages devoted by Beer to
Wedgwood’s review (158–160), we are not told anything about the organ-
ism metaphor. Beer (157) also claims that Darwin had probably read
Alexander Hamilton’s review (Hamilton 1820) of Bopp’s Conjugations Sys-
tems. And yet again we are told nothing of the organism metaphor. In fact,
upon examining Hamilton (1820) and Wedgwood (1833) myself, I found
not the slightest mention or even hint of the organism metaphor. Never-
theless, given Darwin’s close connection with his cousin Hensleigh, given
the scientific milieu of his time, and given Darwin’s wide reading, it is
highly unlikely that he would not have known about the organism
metaphor that had taken over linguistics.
Granting this, Darwin arguably resisted the organism conception of
languages, and like Lyell thought of languages in what might rightly be
called the commonsense view, as primarily horizontal entities delimited by
intercommunication. This follows from his comparison of species with lan-
guages, which, with regard to the former, we have seen was primarily hor-
izontal. Granted, although Darwin’s main analog for species became
languages after he relinquished his monadic theory of evolution, one can
still find him once in a while comparing species with organisms. But the
comparison is only with regard to the production by secondary laws. For
instance, in the Sketch of 1842 (Darwin 1909) he wrote “It accords with
 The Horizontal/Vertical Distinction and the Language Analogy 57

what we know of the law impressed on matter by the Creator, that the cre-
ation and extinction of forms, like the birth and death of individuals should
be the effect of secondary means” (51). Similarly in the Origin he wrote
“To my mind it accords better with what we know of the laws impressed
on matter by the Creator, that the production and extinction of the past
and present inhabitants of the world should have been due to secondary
causes, like those determining the birth and death of the individual” (488).
But the occasional uses of such comparisons does not mean that Darwin
thought of species as primarily vertical entities, in the manner of individ-
ual organisms. Darwin’s comparison was causal, not ontological.
Moreover there was a precedent for this in linguistics. Wilhelm von
Humboldt, friend and mentor of Bopp and brother of Alexander von Hum-
boldt (whose Personal Narrative of Travels had an enormous influence on
Darwin’s choice to become a scientist), arguably maintained a primarily hor-
izontal conception of languages while others were adopting the vertical con-
ception, although this did not prevent him from using organism metaphors
and analogies for languages (cf. Davies 1987, 101–103, nn. 38 and 45; Har-
ris 1993, 19). Moreover, Darwin had a strong reason for not making the
comparison between species and organisms a strong one, and that was prin-
cipally because in his view a species, unlike an individual organism, does not
have a developmental program. Indeed this is why Darwin refused to call his
theory of evolution a “developmental hypothesis,” which was the name often
given to the theories of Lamarck and Chambers.4 As Darwin put it in a let-
ter to Leonard Jenyns (February 14, 1845), “Thanks for your hints about
terms of ‘mutation’ &c; I had had some suspicions, that it was not quite cor-
rect, & yet I do not see my way to arrive at any better terms; it will be years
before I publish, so that I shall have plenty of time to think of better words—
Development wd. perhaps do, only it is applied to the changes of an individ-
ual during its growth” (Burkhardt and Smith 1987, 143). And indeed
Darwin had twice in the Origin made clear that he did not subscribe to what
would later be called orthogenesis (the theory, prevalent in the early 20th cen-
tury, that each species has a developmental program built into it carrying it
in a direction of change independent of selection imposed by the environ-
ment). In the Origin he says “I believe in no fixed law of development, caus-
ing all the inhabitants of a country to change abruptly, or simultaneously, or
to an equal degree. The process of modification must be extremely slow. The
variability of each species is quite independent of that of all others. Whether
such variability be taken advantage of by natural selection, and whether the
variations be accumulated to a greater or lesser amount, thus causing a greater
or lesser amount of modification in the varying species, depends on many
58 DARWIN AND THE NATURE OF SPECIES

complex contingencies” (314; cf. 351). Similarly, in a letter to Hugh Fal-

coner (October 1, 1862) on the topic of natural selection, Darwin wrote “I
suspect that you mean something further,—that there is some unknown law
of evolution by which species necessarily change; and if this be so, I cannot
agree” (Burkhardt et al. 1997, 441). For the same reason Darwin resisted the
word “evolution” in the Origin, because of its use for the development of an
individual organism (“ontogeny” today, the Latin evolutio and evolvere mean-
ing “to unfold,” as in unfolding a scroll), preferring the phrase “descent with
modification” instead. (I shall reply to Robert Richards on this matter in
Chapter 10.) And again, Darwin did not take up William Harvey’s sugges-
tion (October 8, 1860) that Darwin replace “Natural Selection” with “Nat-
ural Evolution” as expressing “the combination of all the powers of nature in
the production of species” (Burkhardt et al. 1993, 416).
Interestingly, linguistics eventually came around, following the revolu-
tion brought to a head by Ferdinand de Saussure roughly at the turn of the
century, switching from what he called diachronic linguistics (which more
often than not had previously involved the organism metaphor) to what he
called synchronic linguistics, in which the ontology of a language is consid-
ered as a system of communication and accordingly a primarily horizontal
entity. This revolution has stuck (for details, cf. Stamos 2002, 178–183).
There is today, following the Copernican revolution of Saussure, in spite of
robust health in the field of historical linguistics, no cladistic language con-
cept. There is no view of language in professional linguistics as expressed by
Ghiselin (1997), according to whom “English has never undergone the ana-
logue of speciation. The language of Beowulf, the Canterbury Tales, Hamlet,
and Huckleberry Finn has changed a great deal without ceasing to be one
and the same individual language” (141). Rather, the modern idea is that,
keeping with English and as Quine (1986) put it, “later English,” in con-
tradistinction to “earlier English,” is “another language” (14).
Darwin did not have the vertical conception of language, as indeed nor
did Lyell. Arguably his conception of language was the commonsense one,
that of a primarily horizontal entity, what Fodor (1975) in an entirely dif-
ferent context called “the good old way: viz., as a system of conventions for
the expression of communicative intentions” (106). All of this was in spite
of what was going on in the field of linguistics around him. And as we have
seen, Darwin thought the same of species, not as individuals, not as verti-
cal entities, but as strongly analogous to languages commonly conceived.
And if indeed I am correct in this, it further seems to me that Darwin
was on firm logical ground in placing the reality of species primarily in the
horizontal rather than in the vertical dimension. There are a number of
 The Horizontal/Vertical Distinction and the Language Analogy 59

arguments that drive home this point, each of them good. John Dupré
(1981), for example, argues that “the objective reality of the [phylogenetic]
branch can be no greater than the objective reality of the grouping of or-
ganisms that constitutes the beginning of the branch” (88). Since any of a
number of different relations between organisms, or between organisms
and their environment, could be used to define the group of organisms
which in turn constitutes the branching point, the vertical perspective alone
is dependent on the horizontal. As Dupré further puts it, “the phylogenetic
criterion must be parasitic on some other, synchronic, principle of taxon-
omy. It cannot generate privileged properties on its own” (89). Elsewhere
in my own writings (Stamos 1996, 138 n. 36, 1998, 462, 2003, 79), I ar-
gued that, for species as with languages, the horizontal dimension is logi-
cally and therefore ontologically prior to the vertical dimension for the
simple reason that a horizontal species can be real even though it lacks a ver-
tical reality (if it evolves gradually one cannot at all say, even roughly, when
it began and when it came to an end), while a vertical species cannot pos-
sibly be real without having successive horizontal realities. In my paper de-
voted specifically to the topic (Stamos 2002), I was required to provide a
further argument in order to satisfy an anonymous referee and the editor.
There I used an argument by analogy, arguing that because species are much
more analogous to languages than to individual organisms, and languages
according to the modern view (and rightly so) are primarily horizontal en-
tities, it follows that the ontology of species is probably also primarily hor-
izontal (probably because argument by analogy can never establish anything
more than a high probability). The nature of the argument can be schema-
tized as follows (l ⫽ any natural language, s ⫽ any species, o ⫽ any organ-
ism, H ⫽ primarily horizontal, V ⫽ primarily vertical):

l and s both have properties ABCDEFG

l has the property H
Therefore, s has the property H with a probability of p

s and o both have the properties TU

o has the property V
Therefore, s has the property V with a probability of q

p⬎q
Therefore, it is more probable that s has the property H than the property V

It remains to be argued that species and languages are similar in more

relevant respects than species and individual organisms. The following list
60 DARWIN AND THE NATURE OF SPECIES

(which is greatly simplified from Stamos 2002, 186–191), though not

exhaustive, will suffice: (i) Species and languages are plastic in a way that
organisms are not. Change in an organism is constrained by its genotype.
Species and languages have no such constraint. They have phylogeny, but
not ontogeny. (ii) With species as with languages, horizontally variation is
the norm. With a multicellular organism, on the other hand, although there
may be much variation in cell types, it is superficial given the unity of the
genome shared by all the cells. (iii) With both species and languages, vari-
ation does not present the same problems horizontally as it does vertically.
Horizontally, both species and languages, although they often have geo-
graphic variation, have in most cases fairly clear borders or ranges, beyond
which there are relatively few individuals, allowing for species maps and
language or dialect maps. Viewed vertically, however, the only borders one
can appeal to are branching points, which present their own kind of prob-
lems given that the history of life, and even more so the history of lan-
guages, is far from being perfectly hierarchical. These features and problems
have no analogs to change within an individual organism. (iv) Species and
languages exhibit both anagenesis (unidirectional) and cladogenesis
(branching) patterns of change, as well as both sympatric and allopatric
(including founder effect) modes of formation. The ontogeny of an indi-
vidual organism, on the other hand, is anagenetic only superficially, since
it retains basically the same genotype throughout its lifetime, while its
modes of reproduction only superficially resemble sympatric and allopatric
models of speciation and language formation (even with binary fission and
budding, the favorite analogs of species-as-individuals theorists, there is
nothing analogous to mitosis in amoeba reproduction or in the copying of
the large and small replicons in bacterial reproduction). (v) Species and lan-
guages are subject to similar forces and mechanisms driving their change,
namely, random variation, natural selection, and drift (especially in founder
populations and bottlenecks). Although these forces and mechanisms are
not completely the same in species and languages, they nevertheless have
much more in common than the forces or mechanisms that drive change
in an individual organism (vi). With individual organisms, it is unequivo-
cal that multiple births do not result in numerically the same individual. In-
stead it produces siblings. Even if the multiple births are qualitatively
identical, the result is not numerical identity. A copy of me, no matter how
similar, is not me (e.g., if it goes out and commits murder, I am not guilty).
For species and languages on the other hand, it is not at all unequivocal
that multiple births result in multiple offspring. In plants, for example, re-
peated allopolyploidy between two parental species is routinely treated by
 The Horizontal/Vertical Distinction and the Language Analogy 61

botanists as resulting in a third species, not a third, fourth, and fifth, etc.,
new species. In other words, the multiple origins, the resulting allopoly-
ploid plants or populations, are routinely treated as conspecific, as being all
of one species (e.g., Ashton and Abbott 1992). If the same were to happen
in linguistics, no modern linguist would treat the different groups as speak-
ing numerically different languages, if their languages were qualitatively
identical, because they in principle could communicate with each other.
In species biology, however, those committed to a cladistic or species-as-
individuals view are committed to identifying the multiple origins as nu-
merically distinct species, even though they are qualitatively identical (or
near identical). This odd consequence has been highly criticized, and rightly
it seems to me. At any rate, it is avoided once species are taken to be pri-
marily horizontal. (vii) In a closely related matter, while organisms typi-
cally undergo an enormous amount of change from birth to death, it is
unequivocal that such protean change results in numerically the same in-
dividual throughout. With species and language change, however, it is not
unequivocal. If the language of a population on an isolated island were to
undergo radical change over time, so that in principle one of the language
users at a later date could not possibly understand one of the language users
at an earlier date (and vice versa), no modern linguist would say they speak
one and the same language. The result is a lineage of language systems, not
a language system. In species biology, on the other hand, some biologists
(again cladists, proponents of species-as-individuals) want to maintain that
if a species undergoes “infinite evolution,” then it is still numerically the
same species throughout (e.g., Ridley 1989, 10). But many biologists do
not think this way (e.g., Mayr 1970, 248). (viii) With organisms, it is un-
equivocal that each organism ages. In linguistics, that view for languages
died over 100 years ago (cf. Fox 1995, 33 n. 3). In species biology, the idea
is similarly dead. Not only does a species lack a genetic program and con-
sequently the genetic mechanisms of aging, but as Raup (1991) also pointed
out, a species that stays above its minimum viable population number is
“virtually immune to extinction” (124, cf. 6). (ix) Finally, with individual
organisms, it is unequivocal that the death of an individual is necessarily
(logically) forever. Again, should a copy of me be made after I die, it is not
me. With species, however, while cladists and proponents of species-as-in-
dividuals claim that the extinction of a species is necessarily forever (they
balk at the thought of Jurassic Park), many other biologists entertain the
view that the extinction of a species is not necessarily forever—Richard
Dawkins (1986, 73) is one example, and Darwin, as we shall see in the next
chapter, was another. In modern linguistics, although the revival of a “dead”
62 DARWIN AND THE NATURE OF SPECIES

language is considered highly improbable, it is not precluded on logical

grounds (cf. Trask 1996, 329–330). And this is because languages are
viewed horizontally, as systems. Systems are not fixed and timeless entities,
but they are not spatiotemporally bounded either. Rather they are dynamic
entities that are logically free to re-evolve and re-exist whether at the same
time and in different places or at different times and places. They are gen-
uine historical entities, but of a very different sort than individuals.
This view of species has interesting implications for speciation. Of
course it is meaningless to speak of “speciation” unless one has first defined
what one means by “species.” We have not yet done that for Darwin, but
in determining that species for him are primarily horizontal, we can indeed
draw some implications for speciation just from the horizontal nature of
species alone.
First, speciation is coupled to evolution. A species that undergoes “in-
finite evolution” is not a species but something else, what might be called
a “species lineage” (cf. Stamos 2002, 190).
Second, it is sometimes thought that since the horizontal dimension is
static, speciation is impossible, so that speciation needs a vertical dimension.
But the horizontal dimension is not static. Instead it is dynamic, gradually
changing with every day, and as such provides (in the sense of time) every-
thing that is needed both for species reality and for speciation (cf. Stamos
2003, 324).
Third, the concept of species as primarily horizontal entities is not a
concept of chronospecies. This is a concept from paleontology, which refers
to chopping up a lineage in the fossil record into successive species.
Chronospecies are typically held to be arbitrary, ultimately nothing more
than subjectively delimited segments of a continuum, mere matters of
taxonomic convenience. And of course from a vertical perspective,
chronospecies are indeed arbitrary. From a horizontal perspective, how-
ever, vertical species appear just as arbitrary! Thus, given a phyletic lineage
in the fossil strata that spans, say, 10 million years, the question “How
many horizontal species, on your view, are there?” is an illegitimate ques-
tion, since it is asked from the vertical perspective. From that perspective,
one might just as well answer that there are infinitely many species as that
there are none. The answer makes no sense because the question was asked
from the wrong perspective, wrong because the vertical dimension is nei-
ther logically nor ontologically the primary dimension for species reality.
The concept of chronospecies is a product of that wrong perspective, and
that is what makes it an illegitimate, nonsensical concept. It is not the hor-
izontal dimension that makes it so. From that dimension, at any one hor-
 The Horizontal/Vertical Distinction and the Language Analogy 63

izontal level it makes perfect sense to ask how many species there are, and
from that perspective meaningful questions get meaningful answers (cf.
Stamos 2003, 316–317).
In arguing for species as primarily horizontal entities, I am definitely
going against a modern trend, namely, the trend in the past few decades to-
ward a cladistic or phylogenetic conception of species in biology and the
corresponding species-as-individuals view in philosophy of biology. How-
ever, as I also pointed out (Stamos 2002), one can see the beginnings of a
reverse trend in phylogenetic systematics, a trend away from a vertical and
back to a horizontal species concept.
Two examples should suffice. The first is the phylogenetic species con-
cept of Joel Cracraft. Originally conceiving of species as individuals (El-
dredge and Cracraft 1980, Cracraft 1987), he more recently (without
notice) changed his view, stating (Cracraft 1997) that species in his view are
not clades (branches) (326) but “terminal taxa” (332), a term in phyloge-
netic systematics that refers to the tips of clades. Another example is the
genealogical species concept of David Baum and Kerry Shaw (1995), which
is based on coalescing gene trees and according to which “exclusive groups
of organisms are only meaningfully delimited among groups of organisms
living at any one point of time” (300). (For more on these species concepts,
cf. Stamos 2002, 193–194, 2003, 270–276, 278 n. 68.)
I believe that Darwin, could he have known of this today, would have
embraced this reverse trend, and for the reasons I have given I would say
that in doing so he would be right.
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 Chapter 4

Common Descent
and Natural Classification

It might be thought that Darwin’s emphasis on genealogy for classifica-

tion, on common descent, completely undermines the analysis in the pre-
vious chapter. For example, in the Origin Darwin repeatedly emphasized
that the “Natural System” of classification, so debated and desirously sought
after by his fellow naturalists, is in fact a genealogical one. As he put it, “all
true classification is genealogical; that community of descent is the hidden
bond which naturalists have been unconsciously seeking, and not some un-
known plan of creation, or the enunciation of general propositions, and
the mere putting together and separating objects more or less alike” (420).
Does not Darwin’s emphasis on genealogy entail that he thought of taxa,
both species and higher, as vertical entities, as branches in the Tree of Life?
What only aids this supposition is that many biologists today, irregard-
less of Darwin, think that if species are to be evolutionary entities, real en-
tities in nature rather than timeless abstract entities, then they must be
thought of as vertical entities, indeed as individuals. Ghiselin (1974) and
Hull (1978), for example, have routinely argued that everything real is ei-
ther a class or an individual, classes are timeless entities, species evolve,
therefore species must be individuals. (I have inveighed against the class/in-
dividual distinction as a false dichotomy in Stamos 1998, 456–459, 2003,
214–215.) Following Ghiselin and Hull, a number of authors have claimed

65
66 DARWIN AND THE NATURE OF SPECIES

that if species are historical entities then they must be individuals. For
example, Eldredge (1985) claims that “punctuated equilibria puts the icing
on the cake in the argument that species are real historical entities, compa-
rable in a formal manner to individual organisms” (122). Moreover, one
can find the related claim that the importance of history reconstruction re-
quires a vertical species concept. Frost and Wright (1988), for example,
claim that “only individuals (⫽ entities) exist independently of definition
and have histories. Thus, only individuals should be included in biological
taxonomies” (201).
Returning to Darwin, in the Origin he seems to be saying the same
thing when he says “If we extend the use of this element of descent,—the
only certainly known cause of similarity in organic beings,—we shall un-
derstand what is meant by the natural system: it is genealogical in its at-
tempted arrangement, with the grades of acquired difference marked by
the terms varieties, species, genera, families, orders, and classes” (456). To
this we may add what he says near the very end, viz. “when we regard every
production of nature as one which has had a history . . . how far more in-
teresting, I speak from experience, will the study of natural history become!”
(485–486). Moreover, this was a view that Darwin long held. For example,
in a letter to George Waterhouse (July 26, 1843) he wrote “According to
my opinion . . . classification consists in grouping beings according to
their actual relationship, i.e. their consanguinity, or descent from common
stocks . . . . it is clear that neither number of species—nor grade of organi-
zation ought to come in, as an element” (Burkhardt and Smith 1986,
375–376). In Descent (1871 I) Darwin states, with regard to the natural
system, that “This system, it is now generally admitted, must be, as far as
possible, genealogical in arrangement. . . . The amount of difference
between the several groups—that is, the amount of modification which
each group has undergone—will be expressed by such terms as genera, fam-
ilies, orders, and classes” (188).
By focusing on common descent for natural classification, it might
seem that Darwin went so far as modern taxonomists of the now domi-
nant cladistic variety, who insist that taxa (both species and higher) must
be monophyletic. This latter term generally has two meanings. The more
general sense is that of single origin, meaning that a taxon cannot have mul-
tiple origins (as in repeated polyploidy). In phylogenetic systematics, poly-
phyletic groups are not considered natural groups. The other, more strict
sense of monophyly (which is not always applied to species but always to
higher taxa), is that of a taxon consisting of the common ancestral organ-
isms and all descendant organisms. If some of the descendant organisms
 Common Descent and Natural Classification 67

are not included in the group, then the group is called paraphyletic, and
most (but not all) of those who subscribe to phylogenetic systematics
eschew paraphyletic taxa since it brings in the criterion of similarity.
When we take a closer look at Darwin, we see that in stressing common
descent he did not at all come close to modern cladistics. In the case of
Darwin’s systematic work on barnacles, Ghiselin and Jaffe (1973) show
that Darwin went some way toward a phylogenetic classification but by no
means as far as cladism, since “he definitely did have paraphyletic taxa in
his system” (138), and also because he included amount of difference and
other practical considerations (134, 139). I suggest that when looking at
what Darwin did with other groups of organisms we get an even better pic-
ture of his view on genealogy and classification, including especially as it
pertains to the species category.
First, it is important to notice that, unlike species, Darwin did not
think that higher taxa are real. For example, early on, again in a letter to
Waterhouse (July 31, 1843), Darwin wrote “I believe . . . that if every or-
ganism, which ever had lived or does live, were collected together (which
is impossible as only a few can have been preserved in a fossil state) a per-
fect series would be presented, linking all, say the Mammals, into one great,
quite indivisible group—and I believe all the orders, families & genera
amongst the Mammals are merely artificial terms highly useful to show the
relationship of those members of the series, which have not become extinct”
(Burkhardt and Smith 1986, 378). To this he added that “classification . . . is
governed by the breaks or chasms in the series” (378). We can see the same
view implied in the Origin, where Darwin says “I believe that the arrange-
ment of the groups within each class, in due subordination and relation to
the other groups, must be strictly genealogical in order to be natural; but
that the amount of difference in the several branches or groups, though al-
lied in the same degree in blood to their common progenitor, may differ
greatly, being due to the different degrees of modification which they have
undergone; and this is expressed by the forms being ranked under differ-
ent genera, families, sections, or orders” (420).
But how strongly did Darwin insist on common descent when it came
to species? It would seem rather strongly but not totally.
A good example to begin with is the case of domestic dogs. Virtually
all naturalists from Buffon onward (as is the case with biologists today) cat-
egorized all the breeds of domestic dogs as members of a single species,
Canis familiaris, because of their interfertility. However, it is striking that
nowhere in his writings, either in his publications, his notebooks, or his
correspondence, did Darwin follow this practice. In other words, there is
68 DARWIN AND THE NATURE OF SPECIES

nowhere that Darwin calls domestic dogs a single species. The main reason,
quite apparently, is because he believed that the group as a whole was do-
mesticated not from a single wild species but from a number of wild species,
domestication eventually eliminating the sterility barriers between the do-
mesticated animals from the wild species. This issue often came up in Dar-
win’s correspondence, with Charles Lyell being almost always his
correspondent on this issue (cf. Burkhardt and Smith 1991, 357, 362–364,
384, 386, 392, Burkhardt et al. 1993, 170, 258, 261, 262–263, 320, 335,
366, 378, 383–384, 393, 397, 399), who by this time had come to think,
contrary to his earlier view (Lyell 1832, 27–28), that all domestic dogs are
descended from wolves.
In the Origin, Darwin’s refusal to classify domestic dogs as a single
species is even more evident, in particular alongside the related case of do-
mestic cattle, namely, European cattle and humped Indian cattle. Explic-
itly following (in part) the doctrine of Pyotr Pallas, Darwin wrote “I believe,
for instance, that our dogs have descended from several wild stocks; yet,
with perhaps the exception of certain indigenous domestic dogs of South
America, all are quite fertile together; and analogy makes me greatly doubt,
whether the several aboriginal species would at first have freely bred to-
gether and have produced quite fertile hybrids. So again there is reason to
believe that our European and the humped Indian cattle are quite fertile to-
gether; but from facts communicated to me by Mr. Blyth, I think they
must be considered as distinct species” (254). To add to this, early in the
Origin we find Darwin state that “I should think, from facts communi-
cated to me by Mr. Blyth, on the habits, voice, and constitution, &c., of
the humped Indian cattle, that these had descended from a different abo-
riginal stock from our European cattle; and several competent judges believe
that these latter have had more than one wild parent” (18).
Thus the same conclusion, on Darwin’s view, follows (though implicit)
for domestic dogs, which Darwin discusses, as analogous, immediately
above the cattle example, namely, that in spite of their perfect interfertility
they must be classified as more than one species because of their multiple
origin and differentiated characters (cf. also Variation 1868 I, 21, 26,
31–32, 34).
I will explore the significance of differentiated characters for Darwin’s
species concept in the next chapter. For the present, it is interesting to focus
more closely on Darwin’s source, namely Edward Blyth. In the Origin Dar-
win tells us that Blyth is one “whose opinion, from his large and varied
stores of knowledge, I should value more than that of almost any one” (18).
Given such high esteem for Blyth, we should trace the source from Blyth
 Common Descent and Natural Classification 69

for Darwin’s reference to humped Indian cattle. In a letter to Darwin (April

21, 1855), written in India where Blyth was residing from 1841 to 1862
as the curator of the museum of the Asiatic Society of Bengal, Blyth wrote
“It is easy to dogmatize & say that Bos indicus is but a humped variety of
B. taurus, but the stump is one of many differential characters, though seen
very early in foetal life!” (Burkhardt and Smith 1989, 312). The differen-
tial characters of Bos indicus, in spite of hybridizability with Bos taurus, in-
clude not only the muscular hump, but a great difference in voice, in habits
such as never seeking shelter from the sun, and in never standing knee and
belly deep in water. In a later letter of the same year (September 30 or Oc-
tober 7), Blyth adds a further number of morphological differences, such
as differences in height, ears, and horns. He then states “What more can be
required to characterize it as a peculiar species? Why the advocates of the
opposite opinion can adduce only the prolificacy of the hybrids” (451).
Aside from the significance of Blyth’s species concept (which quite appar-
ently is not based on fertility or sterility but on constant and distinct char-
acters), what is significant for our immediate purpose is that in these and
other letters Blyth nowhere suggests a multiple origin for the two cattle
forms. Instead it is Darwin who infers that and, at least in part, uses the lack
of common descent to keep them separate as species.1
Indeed the topic is only highlighted by Darwin’s view on convergence.
In a letter to Darwin (January 3?, 1860), H.C. Watson suggested, as a cor-
relate to Darwin’s principle of divergence, that Darwin allow “the hypoth-
esis or inference that individuals converge into orders, genera, species, as
well as diverge into species, genera, orders, through nepotal descent”
(Burkhardt and Smith 1989, 11–12). In reply to Watson (January 5–11),
Darwin stated “With respect to ‘convergence’ I daresay, it has occurred,
but I should think on a very limited scale . . . and only in case of closely re-
lated forms. . . . the same cause acting on two closely related forms (i.e.
those which closely resembled each other from inheritance from a com-
mon parent) might confound them together so closely that they would be
(falsely in my opinion) classed in same group” (18).
The possibility of convergent evolution, which we see here Darwin al-
lows, raises an interesting issue about similarity. We have seen earlier in this
chapter that Darwin claims that “propinquity of descent” is “the only known
cause of the similarity of organic beings” (413; cf. 456). And yet convergent
evolution is also a real cause of similarity, and Darwin recognizes this. In-
deed he distinguishes the two kinds of similarity by calling the former “real
affinities” and the latter “analogical or adaptive resemblances” (427). The lat-
ter are adaptive because animals “belonging to two most distinct lines of
70 DARWIN AND THE NATURE OF SPECIES

descent, may readily become adapted to similar conditions, and thus assume
a close external resemblance.” An obvious example is that of whales and fish.
The problem for Darwin, however, is that convergent similarity is “almost
valueless to the systematist,” the reason being that “such resemblances will
not reveal—will rather tend to conceal their blood-relationship to their
proper lines of descent” (427). Interestingly Lyell, in a letter to Darwin (Oc-
tober 28, 1859), raised much the same problem for Darwin’s theory of the
polyphyletic origin of domestic dogs, in that “if the three wild species men-
tioned above [‘wolf, fox & jackal’] could produce any or all these races of
dogs (or incipient species) by coalescing & interbreeding, I find it impossi-
ble to hope to trace any clue to the past transformations of species or their
probable birthplaces” (Burkhardt and Smith 1991, 363). At any rate, in De-
scent (1871 I) Darwin comments on the extreme unlikelihood, calling it “ut-
terly incredible,” of convergence at the species level “as to lead to a near
approach to identity throughout their whole organisation” (231).
Thus far it seems that Darwin’s species concept has a strong element
of common descent in it, strong enough perhaps to make it a primarily ver-
tical conception of species. However, in addition to what we have seen in
the previous chapter, there is further evidence that leads away from this
conclusion. Most important is that Darwin did not think that extinction
is necessarily forever. Granted, in the Origin he can be found saying in his
chapter on geology “When a group has once wholly disappeared, it does not
reappear” (344), and we have seen in chapter 2 that Darwin thought that
extinction is a law of nature. But we have to be extremely careful here. To
say that in the geological record extinct groups do not reappear is not nec-
essarily to say that it is logically impossible, or even physically impossible,
that they could reappear. Indeed elsewhere in the Origin Darwin says
“When a species has once disappeared from the face of the earth, we have
reason to believe that the same identical form never reappears” (313), the
reason being “for the link of generation has been broken” (344). Even if a
lineage should come to occupy a niche left vacant by an extinct species, the
new occupant of the niche is not the same species, “for both would almost
certainly inherit different characters from their distinct progenitors” (315).
Notice the key words in the first and third of these passages, namely, “we
have reason to believe” and “almost certainly.” Darwin is pretty clear here
that he does think it could happen, that an extinct species could reappear,
just that it is extremely unlikely.
Darwin is clearer on this in his correspondence. In a letter to Lyell (June
21, 1859) he wrote “You ask about specific centres, if you change terms into
specific areas, my theory quite requires them; i.e. it is, I think, next door to
 Common Descent and Natural Classification 71

an impossibility that the same species should have been formed identically
the same in any two areas” (Burkhardt and Smith 1991, 307–308). “Next
door to an impossibility,” of course, is nevertheless to affirm a logical possi-
bility. Thus for Darwin it is logically possible, albeit highly improbable, that
someday, to quote from Lyell (1830), “The huge iguanodon might reap-
pear in the woods, and the ichthyosaur in the sea, while the pterodactyle
might flit again through umbrageous groves of tree ferns” (123).
Darwin held the same view on languages. In Descent (1871 I) he says
“A language, like a species, when once extinct, never, as Sir C. Lyell re-
marks, reappears” (60). I suggest that the word “never” here should not be
taken very strongly. As we have seen with the Origin and from Darwin’s
correspondence, he thought that the reappearance of an extinct species is
highly unlikely. In the case of languages this interpretation is only strength-
ened if we turn to the passage in Lyell’s writings which Darwin refers to in
the Descent. Lyell’s remark is to be found in The Antiquity of Man (1863),
in which he says “a language which has once died out can never be revived,
since the same assemblage of conditions can never be restored” (467). It
may not be clear whether Lyell’s claim here is a claim of physical necessity
or merely only probability. However, it is surely significant that much ear-
lier Lyell (1838) stated only as a simple matter of geological observation
that no species “ever reappeared after once dying out” (275; cf. Darwin
Notebook E, Barrett et al. 1987, 105). Of much greater significance
is Lyell’s (1830) postulation of a vast, geologically recurring “great year”
(ch. 7), such that, as we have seen above, huge iguanodons, ichthyosaurs,
and pterodactyles will once again populate the earth.
Given, then, that Darwin thought that the extinction of a species, like
that of a language, is not necessarily forever, it follows that his concept of
common descent was not a concept of monophyly in any of the forms that
we encounter today (each of which insist on a single origin for each species).
Common descent was not for Darwin a matter of logic or methodology,
but rather only probability. The possibility that a particular species could
arise independently in more than one place and time was so improbable to
him that it was not worth worrying about.
This, of course, makes Darwin’s species concept radically different
from the species-as-individuals thesis of Ghiselin and Hull and from phy-
logenetic taxonomy, both of which hold that extinction is necessarily (log-
ically) forever and that species taxa must each have no more than a single
origin. Combined with what we have seen in the previous chapter, namely,
Darwin’s emphasis on the horizontal dimension for the reality of species,
we can only conclude that common descent is not part of the ontology of
72 DARWIN AND THE NATURE OF SPECIES

species for Darwin but instead serves as, or rather the evidence for it serves
as, an important criterion in picking out horizontal species (as well as higher
groups). To use a modern distinction, the difference is between constitu-
tive properties and diagnostic criteria (akin to the distinction between a
disease and its symptoms), common descent being diagnostic but not con-
stitutive of species.
It is also important to add here that although Darwin thought of com-
mon descent as a criterion of species delimitation, he did not think of it as
the most important criterion. What was most important for him is what we
shall examine in the next chapter. But for the remainder of the present
chapter I want to focus more closely on Darwin’s emphasis on common
descent as well as the emphasis on monophyly in modern systematics.
Part of Darwin’s argument was that naturalists were already using com-
mon descent in their classification of organisms into species. As he puts it
in the concluding chapter of the Origin, “every naturalist has in fact brought
descent into his classification; for he includes in his lowest grade, or that of
a species, the two sexes; and how enormously these sometimes differ in the
most important characters, is known to every naturalist” (424). Indeed then
as today, no naturalist, as Darwin says, “dreams of separating them.” Not
even extreme splitters would ever dream of this. And if the sexes were ever
separated by a taxonomist, it was only by accident, such that once the mis-
take became known it was quickly corrected. Such accidents are caused by
the fact that many species are remarkably sexually dimorphic. For example,
in the case of woodpeckers the sexes have evolved niche differentiation to
such a degree that there is an extreme dimorphism in bill morphology. In
the case of fur seals, the male is several times larger than the female. And
in the case of mallard ducks, the colors are so different that one would nat-
urally be inclined to think that they are different species if one did not ac-
tually see them copulate (indeed, according to Mayr and Short 1970, 88,
Linnaeus himself made the mistake of classifying male and female mallards
into separate species).
Darwin goes on to point out that naturalists also use descent for keep-
ing larval and adult stages in the same species. As well, he says, the natural-
ist “includes monsters; he includes varieties, not solely because they
resemble the parent-form, but because they are descended from it. He who
believes that the cowslip is descended from the primrose, or conversely,
ranks them together as a single species, and gives a single definition” (424).
Mayr (1957a) surmises that pre-Darwinian taxonomists emphasized
common descent below the species level because as essentialists it was the
only way they could deal with variation:
 Common Descent and Natural Classification 73

What is unexpected for this pre-Darwinian period . . . is the frequency

with which “common descent” is included in species definitions. When
such an emphatically anti-evolutionary author as v. Baer (1828) defines the
species as “the sum of the individuals that are united by common descent,”
it becomes evident that he does not refer to evolution. . . . Expressions
like “community of origin” or “individus descendants des parents com-
muns” (Cuvier) are frequent in the literature. These are actually attempts
at reconciling a typological species concept (with its stress of constancy)
with the observed morphological variation. [7]

This is an interesting suggestion and I won’t pursue it any further.

What is important for our present purpose is that in the Origin Darwin
goes on to point out that “may not this same element of descent have been
unconsciously used in grouping species under genera, and genera under
higher groups, though in these cases the modification has been greater in
degree, and has taken a longer time to complete? I believe it has thus been
unconsciously used; and only thus can I understand the several rules and
guides which have been followed by our best systematists” (425). Darwin,
however, was doing more than simply arguing that systematists, in classi-
fying groups within groups, were unconsciously employing common de-
scent. As we have seen at the beginning of this chapter, he also argued that
it ought to be so for any natural system. Indeed we can see Darwin using a
consistency argument. If you use common descent in classifying sexes, on-
togenetic stages, sports, and varieties into one species (and you all do), and
if branching evolution is true (and Darwin argued that it is), then you ought
also to use common descent when grouping higher taxa.2
What we have to keep in mind in all of this, however, is that Darwin
repeatedly claimed that common descent (he used the term “genealogy”)
was not enough for classification. The Origin itself caused some confusion
about this and in his correspondence Darwin attempts to clear it up. For
example, in a letter to Hooker (December 23, 1859) Darwin wrote “geneal-
ogy by itself does not give classification” (Burkhardt and Smith 1991, 444),
and in a letter to Andrew Murray (April 28, 1860) he wrote “In case of
classification, descent alone, as I believe I have shown, will not do; you
must combine principle of divergence of character & descent” (Burkhardt
et al. 1993, 179).
So Darwin was certainly not a cladist, but so what? What is instruc-
tive about Darwin’s emphasis on common descent, or rather his lack of ex-
clusive emphasis on common descent, is that it provides a much more
sensible approach to the topic of multiple origins than that found today
among cladists.
74 DARWIN AND THE NATURE OF SPECIES

Let’s see how this is. Darwin was arguably wrong in his claims against
multiple origins (including that an extinct species almost certainly never
reappears). What is now well-known is polyploidy, a chromosomal acci-
dent in a zygote resulting in at least three times the haploid (half) somatic
chromosome number of the parent(s). When only one species is involved
it is called autopolyploidy, when two species are involved (as in interspecific
hybridization) it is called allopolyploidy. Although quite rare in animals, it
is common in plants. The interesting thing about polyploidy is that it is
normally considered instantaneous speciation by biologists, since the poly-
ploids are reproductively isolated from their parental species. Indeed so
common is polyploidy in plants that it is estimated that 70–80% of all an-
giosperm (flowering plant) species in the wild were produced by polyploidy.
We therefore don’t need science fiction examples (or genetic engineering)
to discuss realistically the related issues of multiple origins and the reap-
pearance of extinct species. In the previous chapter I cited the example of
multiple origins given by Ashton and Abbott (1992), and there are many
more (cf. Stamos 2003, 320 n. 15). The problem is that repeated poly-
ploidy, if genealogy is taken strictly with similarity being completely irrel-
evant, arguably results in multiple species, not one species, no matter how
qualitively similar the resulting populations. Strict cladists are forced to this
conclusion, because for them only monophyletic taxa are natural taxa, so
that a species cannot have multiple origins. Thus for them, if species A and
species B repeatedly hybridize and produce 20 polyploid populations, each
of those polyploid populations (no matter how genetically and morpho-
logically identical, no matter how reproductively compatible) would have
to be counted as a distinct species with a distinct binomial. What is worse,
since each branching point is a speciation event for cladists—and only
branching points count for cladists as speciation events, such that at any
branching point (whether speciation by splitting or speciation by budding)
the parental species automatically goes extinct (this has been dubbed “Hen-
nigian extinction” by one of its critics)—both of the parental species auto-
matically go extinct at the point of polyploid hybridization and become
distinctly new species, even though they have not changed at all (cf. Rid-
ley 1989, 5; Stamos 2003, 265)!
Had Darwin known about polyploidy, I think it is safe to say he would
have thought the cladistic interpretation of it pure nonsense. It is as ridicu-
lous for species as it would be for languages. He would see the cladistic con-
cept of monophyly for what it is, viz. a convention, not a causal process, its
mistake being that it confuses methodology with ontology (cf. Stamos
2003, 309). In the Origin Darwin writes of “the vera causa of community
 Common Descent and Natural Classification 75

of descent” (159), but I think the case of polyploidy would have made him
agree that common descent is only really a subsidiary vera causa of species
ontology, it is not the main one (as we shall see in the next chapter). Logi-
cally speaking, Darwin had no objection to multiple origins. If species could
arise multiply from the same parental species, then, just like languages if the
same were to occur, he would rank all of the new productions as one. We
shall see from the next chapter that Darwin would be forced to this conclu-
sion because of his emphasis on adaptations. If the polyploids had new
adaptations, then the multiple polyploids would be conspecific. Whether
they were interfertile with each other and intersterile with their parental
species would have nothing to do with it, as we shall see in chapter 6. Since,
again as we shall see in the next chapter, Darwin thought it highly unlikely
that new adaptations are produced by anything except gradual, cumulative
natural selection, it is highly doubtful that he would give the polyploids
new specific status.3 At any rate, what Darwin’s species concept makes us
realize is that although common descent was very important for him as a
species criterion, it was nevertheless not absolute and in some cases would
not apply.
What the case of repeated polyploidy also relates to is the matter of
whether the extinction of a species is necessarily forever. Suppose our species
A and species B were to produce a polyploid population that subsequently
went extinct, and that, a season later, species A and species B were to do it
again. According to cladism, each of the two polyploid populations, the
former and the latter, would have to be distinct species, no matter how
qualitatively similar. Darwin, however, as we have seen in the previous para-
graph and shall see from the next two chapters, would probably not agree
that the first polyploid population is a new species, because it is a kind of
hybrid and almost certainly would not have any new adaptations. So like-
wise in the case of repeated polyploidy, whether in the above scenario or in
the previous scenario, he would not consider multiple origins multiple
species. Nevertheless, if the new polyploid population that went extinct
had a new adaptation, and if the second polyploid population was just like
it, then Darwin would have no problem (as with many biologists today)
classifying the two polyploid populations as conspecific. In other words,
just as in the case of languages if the same were to occur, he would agree that
an extinct species reappeared. This consequence, however, is logically pre-
cluded by cladistic principles and the species-as-individuals view.
Darwin, as we can see from the above, including his view on extinc-
tion, and as we shall see in the next chapter, was not against similarity in
the ontology of species, contrary to cladists and proponents of species as
76 DARWIN AND THE NATURE OF SPECIES

individuals. And again, guided by the language analogy, this makes good
sense. As we have seen in the previous chapter, according to the view of the
above philosophies a species that undergoes “infinite evolution,” to use
the phrase of Ridley (1989, 10), is numerically the same species through-
out. I don’t think Darwin would have agreed with this, again, because he
thought “genealogy by itself does not give classification” and he was not
completely against similarity. In fact, many cladists are not completely
against similarity either! I have found a significant number of cases where
cladists, contrary to the principles of Willi Hennig, refuse to allow that a
species that buds off a founder population (which in turn becomes a
species), but that itself remains unchanged, automatically goes extinct at
that branching point and becomes a new species (Stamos 2003, 262). In
such cases, as I put it, these cladists break with cladism proper and “allow
similarity to slip in through the back door” (263). Indeed arguably, if one
wants to have a sensible ontology for species, one cannot avoid similarity.
Of course one should not go to the other extreme either, involving only
similarity, as with the morphological and phenetic species concepts. This
is to invite a whole new set of problems and absurdities (cf. Stamos 2003,
81–83, 129–133, 311–312). Instead it would seem that the most viable
position, and I think Darwin would agree as we shall see in the next chap-
ter, is to find some sort of way of combining similarity with other consid-
erations important in the ontology of species, which is what I attempted
in my species book (Stamos 2003, ch. 5).
It is often argued that a species concept, to be viable, must be consistent
with history reconstruction. For example, this was Donn Rosen’s (1979)
complaint against the biological species concept of Mayr. Rosen’s paper is
considered the inauguration of so-called phylogenetic species concepts,
species concepts that attempt to make species consistent with phylogenetic
history but that are not necessarily cladistic, and it is important to look more
closely at his claims. According to Rosen, proponents of the biological
species concept such as Mayr sometimes unite into a single species two good
and distinct species that have a hybrid zone between them (276). The prob-
lem with this, for Rosen, is that reproductive compatibility is an ancestral
character (plesiomorphy) that is gradually lost in the descendant species dur-
ing geographic isolation. Moreover, he says, “It is to be expected that repro-
ductive compatibility, like other primitive traits, might be retained or altered
in a mosaic pattern during evolution, an inference which is entirely consis-
tent with the results of natural and laboratory mating patterns in Xiphopho-
rus” (277). The conclusion follows, then, that “the ‘biological species
concept’ will lead to inferences that are in direct conflict with the avowed
 Common Descent and Natural Classification 77

aims of systematics, viz., to reconstruct the genealogical history of lineages

by a process of estimating a hierarchy of relationships” (277). Rosen then
goes on to give the first version of what has later been called a phylogenetic
species concept, in his case one that defines a species as having “one or more
apomorphous features” (277) (an apomorphy is a derived character, not an
ancestral character) and having “a specifiable geographic integrity” (278).
Thus conceived, he says, “all populations [species by his definition] defined
by apomorphic traits can be incorporated into a cladistic hierarchy, and . . .
this cladistic hierarchy forms the only logical basis for discussions of the his-
tory of organic change in time and space” (277).
In my species book (Stamos 2003, 84–85, 273) I provide a critique of
Rosen’s phylogenetic species concept, but that is not what I want to bring
to focus here. Instead, it is the claim that a species concept should be con-
sistent with history reconstruction. The problem is that nature is not per-
fectly hierarchical as cladistics desires. As we have seen above, nature
includes rampant polyphyly, and allopolyploidy is not the only example
(cf. Stamos 2003, 240 n. 46). In focusing on the importance of genealogy
for classification, Darwin was surely right, but he was also surely right in not
focusing exclusively on genealogy. The case of multiple origins alone shows
that similarity must also be given some account. But to do so does not nec-
essarily preclude responsible history reconstruction. In fact, in some cases
it might be necessary. The case of repeated polyploidy speciation discovered
by Ashton and Abbott (1992) is a case of history reconstruction, but it was
not done using cladistic principles; indeed this would have been impossi-
ble. As McDade (1995) put it, “current phylogenetic methods are inappro-
priate for taxa with complex reticulating histories, and yet the phylogenetic
history of such groups is at least as interesting as those of divergently evolv-
ing groups” (616–617). At the species level especially, then, something
more is needed than cladistic principles. As the example of Ashton and Ab-
bott makes clear, good history reconstruction at the species level is not sim-
ply a matter of distinguishing the plesiomorphic from the apomorphic
characters. Instead one has to use all the evidence available (cf. Stamos
2003, 318–320 for a detailed discussion on Ashton and Abbott 1992). And
indeed if history reconstruction is really what one wants, if it is what really
matters, then why confine oneself to only one methodology? This seems to
me a misplaced loyalty.
Here again we can see the value of Darwin. In focusing on the hori-
zontal dimension for the ontology of species, in analogy with the ontology
of languages, he avoids in one stroke the absurdities involved with cladis-
tic species concepts and the species-as-individuals view. In focusing on
78 DARWIN AND THE NATURE OF SPECIES

genealogy, however, species must be consistent with history reconstruc-

tion. But this, for Darwin, did not preclude multiple origins or the reap-
pearance of an extinct species. In all of this, again, Darwin was guided
very sensibly by his language analogy. In historical linguistics, languages
are considered historical entities, and reconstructing language history is a
thriving exercise (cf., e.g., Fox 1995; Trask 1996; Lass 1997), and yet lan-
guages are viewed as primarily horizontal entities, not as individuals, and
language reconstruction, because of even greater reticulation in their his-
tories, is only partially guided by cladistics (cf. Hoenigswald and Wiener
1987, especially the paper by Wiener).
Edward Wiley (1981, 74–75) has made the useful suggestion that the
ontology of biology should admit of not two (as with Ghiselin and Hull)
but of three categories, namely, class, individual, and historical group. In
his view, the species category is a class, species are individuals, and higher
taxa are historical groups (because even though they are spatiotemporally
bounded they lack cohesion). What is needed, it seems to me, and Darwin
would seem to be in agreement, is the recognition not only that species are
not individuals, but that the category “historical group” should be changed
to “historical entity.” As primarily horizontal entities demarcated in part by
common descent, species are historical entities, not individuals or classes.
Individuals are historical entities too, but of a very different kind. As ver-
tical entities they contain their history within themselves (or rather a large
part of it). Historical entities such as species and languages, however, do not
as horizontal entities contain their history within themselves. But they are
nonetheless historical for it. As historical entities that are not individuals,
that are not even groups, they are not spatiotemporally bounded but are free
to have multiple origins and to go extinct and re-evolve, whether at the
same time and in different places or at different times and places. But as dy-
namic entities they are not the timeless and fixed entities of abstract classes
either. These categories (class and individual) just do not at all adequately
capture the ontology of species. At an intuitive level, guided by the lan-
guage analogy, Darwin knew this.
In sum, in conceiving of species as primarily horizontal, and yet in
maintaining that all true classification must be genealogical, Darwin was
not involving himself in a contradiction. In modern linguistics, particular
languages are thought of as primarily horizontal (synchronic) entities, even
though historical linguistics thrives. And this is in spite of getting over,
roughly 100 years ago, the view that languages are individuals. Biology,
however, seems to be over 100 years behind. In spite of agreeing that true
classification must be genealogical, that species (like languages) are histor-
 Common Descent and Natural Classification 79

ical entities, many biologists (and many philosophers) seem to think that
this means that species must be individuals. But it just doesn’t follow. Dar-
win knew this. Post-Synthesis biologists such as Mayr knew this. During
the last 30 or 40 years, however, mainly because of the influence of cladism
and the species-as-individuals view, biology and its philosophy seem to have
regressed. Fortunately, as pointed out at the end of the previous chapter,
there does seem to be a trend within the field of phylogenetic systematics
itself, back toward a primarily horizontal conception of species, while at
the same time retaining the view that species are historical entities. This is
a trend that Darwin would certainly have welcomed.
This page intentionally left blank.
 Chapter 5

Natural Selection
and the Unity of Science

In leading up to what is for Darwin the most important criterion in species

delimitation, it will be useful to look at the main feature of John Beatty’s
theory, which as we shall see in chapter 8 has become the received view.
Following Ghiselin’s (1969) view that Darwin was a species taxa though not
species category realist, so that Darwin’s nominalistic definitions of
“species” applied only to the species category, Beatty (1985) provided a
strategy theory to explain Darwin’s nominalistic definitions of “species.”
According to Beatty, Darwin distinguished

between what his fellow naturalists called “species” and the non-evolu-
tionary beliefs in terms of which they defined “species.” Regardless of
their definitions, he argued, what they called “species” evolved. His species
concept was therefore interestingly minimal: species were, for Darwin,
just what expert naturalists called “species.” By trying to talk about the
same things that his contemporaries were talking about, he hoped that his
language would conform satisfactorily enough for him to communicate
his position to them. [266]

Again and again, we are told by Beatty that Darwin “tried to get beyond de-
finitions to referents” (269), that “The evolution issue was accordingly, for
Darwin, an issue concerning the species so designated by naturalists” (274),

81
82 DARWIN AND THE NATURE OF SPECIES

that “he used the term [‘species’] in accordance with examples of its refer-
ential use by members of his naturalist community” (277), and moreover
that this “also allowed Darwin to communicate the position that the term
‘species’ was undefinable” (274).
I shall look more closely at the various aspects of Beatty’s strategy the-
ory in chapters 8 and 9. For the present, I want to focus on his claim that
Darwin simply followed the species designations of his fellow naturalists.
The other side of the coin to this claim is that Darwin did not try to change
any of the species designations of his fellow naturalists. Both claims, of
course, are factual claims, and accordingly they are to be decided by noth-
ing else but an appeal to the extant evidence. When we look at that evi-
dence, however, we cannot help but be struck by the fact that Darwin often
went against what his fellow naturalists called “species,” and often decided
one way or another in cases where they did not agree. This is a striking fact,
which creates a serious problem for anyone who claims that Darwin was a
species nominalist, taxa or category. But even more important is what is to
be learned when we look closely at those cases where Darwin went against
his fellow naturalists.
A good place to begin is with a close look at what Darwin wrote in the
Origin about primroses and cowslips. But first, it is important to notice a
remark made by Malcolm Kottler (1978), that “Both Darwin and Wallace
were fully aware of well-defined species. But they were more interested in
borderline cases . . . not for the purpose of proving that species did not re-
ally exist, but rather for the purpose of disproving special creation. If species
had been independently created, borderline cases should not exist. Yet such
doubtful forms were abundant” (297). This claim about the importance
of borderline cases, of course, is surely true. It was necessary for Darwin
that the existence of borderline cases was proved to exist so that varieties
could be claimed to be incipient species in accordance with his thesis of
gradualism. As Darwin put it in the Origin, “wherever many closely-allied
species occur, there will be found many forms which some naturalists rank
as distinct species, and some as varieties; these doubtful forms showing us
the steps in the process of modification” (404).
But even more important for Darwin’s argument against special cre-
ation, more important than borderline cases between varieties and species,
were cases that according to the general species concept of creationists
would have to be classified as both varieties and species, in other words,
cases that proved that the traditional criteria were incoherent and that there-
fore served as a falsification of the creationist species concept.
 Natural Selection and the Unity of Science 83

Such was the case of primroses and cowslips, and it explains why Dar-
win was so fond of referring to them. The problem is this: As Darwin tells
us in the Origin, primroses and cowslips are “united by many intermediate
links” (50) and so are “generally acknowledged to be merely varieties”
(485). In other words, according to the traditional, creationist view, “they
grant some little variability to each species, but when they meet with a
somewhat greater amount of difference between any two forms, they rank
both as species, unless they are enabled to connect them together by close
intermediate gradations” (297). Thus, part of the creationist species con-
cept involved the criterion that if any two forms are connected by interme-
diate gradations, no matter how different they may be, they must be
classified as conspecific varieties rather than as separate species. But another
important part of the creationist species concept was the sterility criterion.
As Darwin put it, “The view generally entertained by naturalists is that
species, when intercrossed, have been specially endowed with the quality of
sterility, in order to prevent the confusion of all organic forms” (245). And
again, as regards varieties, “a supposed variety if infertile [intersterile] in
any degree would generally be ranked as species” (271).
This is what makes primroses and cowslips so interesting. United by
many intermediate links and therefore ranked as varieties, Darwin on the
other hand also informs us that “according to very numerous experiments
made during several years by that most careful observer Gärtner, they can
be crossed only with much difficulty” (49–50), so that Gärtner “ranks them
as undoubted species” (268). Thus, according to creationist criteria, prim-
roses and cowslips must be ranked as both varieties and species.
Of even more interest and importance than the negative consequences
of primroses and cowslips for the creationist species concept, however, is
what Darwin’s classification of them reveals to us about his own species
concept. For, unlike most other naturalists, and therefore contrary to
Beatty’s strategy theory, Darwin classified primroses and cowslips as sepa-
rate species. As he put it in the Origin, he thought them “worthy of specific
names” and that “in this case scientific and common language will come
into accordance” (485).
At first sight, this is extremely odd. As Kottler (1978) points out, “In
the Origin . . . the only explicit distinction which he [Darwin] drew be-
tween variety and species was the presence or absence of intermediate gra-
dations, not the presence or absence of interbreeding” (297). As Darwin in
the Origin put it, “Hereafter we shall be compelled to acknowledge that
the only distinction between species and well-marked varieties is, that the
84 DARWIN AND THE NATURE OF SPECIES

latter are known, or believed, to be connected at the present day by inter-

mediate gradations, whereas species were formerly thus connected” (485).
This is, of course, and significantly for my analysis in chapter 3, a purely
horizontal criterion. And notably it is followed by Darwin in Descent (1871
I), where he takes sides on the then prevailing controversy over whether
the different human races are conspecific varieties (monogenism) or con-
generic species (polygenism), a debate that had an interesting relation to the
debate over human slavery (cf. Gould 1981, 69–72). Therein Darwin reaf-
firms that “Independently of blending from intercrossing, the complete ab-
sence, in a well-investigated region, of varieties linking together any two
closely-allied forms, is probably the most important of all the criterions of
their specific distinctness” (215). Accordingly “the most weighty of all the
arguments against treating the races of man as distinct species, is that they
graduate into each other, independently in many cases, as far as we can
judge, of their having intercrossed.” To this Darwin adds that “Every nat-
uralist who has had the misfortune to undertake the description of a group
of highly varying organisms, has encountered cases (I speak after experi-
ence) precisely like that of man; and if of a cautious disposition, he will
end up by uniting all the forms which graduate into each other as a single
species; for he will say to himself that he has no right to give names to ob-
jects which he cannot define” (226–227). Darwin’s own experience, of
course, relates mostly back to his work on barnacles. And indeed in his In-
troduction to his work on the Balanidae (Darwin 1854b), he says “In de-
termining what forms to call varieties, I have followed one common rule:
namely, the discovery of such closely allied, intermediate forms, that the ap-
plication of a specific name to any one step in the series, was obviously im-
possible; or, when such intermediate forms have not actually been found,
the knowledge that the differences of structure in question were such as, in
several allied forms, certainly arose from variation” (156).
What is even more problematic is that in both cases, namely, that of
primroses and cowslips and the races of man, Darwin thought the evidence
preponderant that they each originated from a single stock. In the Origin
Darwin tells us that primroses and cowslips “are united by many intermedi-
ate links, and it is very doubtful whether these links are hybrids; and there is,
it seems to me, an overwhelming amount of experimental evidence, showing
that they descend from common parents” (50). Similarly in Descent (1871 I)
Darwin tells us that “Those naturalists . . . who admit the principle of evo-
lution . . . will feel no doubt that all the races of man are descended from a
single primitive stock; whether or not they think fit to designate them as dis-
tinct species, for the sake of expressing their amount of difference” (229).1
 Natural Selection and the Unity of Science 85

To anyone who has read Darwin’s chapter on classification in the Ori-

gin, Chapter XIII, the phrase “amount of difference” should ring a bell. Al-
though Darwin tells us therein that “all true classification is genealogical”
and that classification “must be strictly genealogical in order to be natural,”
he immediately adds that “the amount of difference in the several branches
or groups . . . is expressed by the forms being ranked under different gen-
era, families, sections, or orders (420, second italics added).
Kevin Padian (1999) argues that “Darwin chose his verbs carefully”
(357), that the is refers not to Darwin’s own views on the matter but only
to the practices of his fellow naturalists, and that Darwin himself did not
champion dual criteria for classification (genealogy and similarity) but only
one criterion (genealogy), so that for Darwin “similarity is not in itself a cri-
terion for classification, but [merely] a means to understand genealogy”
(356). Padian also focuses on Darwin’s letter to Hooker (December 23,
1859) that I quoted in the previous chapter, in which Darwin wrote “ge-
nealogy by itself does not give classification” (Burkhardt and Smith 1991,
444), and argues that the context of the letter, which is Darwin’s response
to the evolutionary views of the French botanist C.V. Naudin, indicate that
Darwin had misunderstood Naudin, such that “Darwin did not mean
should when he said ‘genealogy alone does not give classification.’ In other
words, he meant that in practice it does not, and he could not see how
Naudin (as he thought) could say that it did” (360).
Padian’s claims, however, do not stand up to close scrutiny, and all the
evidence does indeed strongly point to the conclusion that Darwin thought
that similarity should be taken to be constitutive of taxa (species in particular),
though not exclusively similarity. To return to the passage in the Origin that
Padian reinterprets, the is does indeed refer to Darwin’s own views, and Dar-
win makes this clear only a couple of pages further, where he states “Thus,
on the view which I hold, the natural system is genealogical in its arrange-
ment, like a pedigree; but the degrees of modification which the different
groups have undergone, have to be expressed by ranking them under differ-
ent so-called genera, sub-families, families, sections, orders, and classes” (422,
italics added). Moreover, to Padian’s highly questionable reinterpretation of
Darwin’s letter to Hooker above, we need only add Darwin’s letter to Andrew
Murray (April 28, 1860), in which he wrote “In case of classification, de-
scent alone, as I believe I have shown, will not do; you must combine prin-
ciple of divergence of character & descent” (Burkhardt et al. 1993, 179).
Finally, what clinches the matter, especially with regard to species taxa, is
Darwin’s view on extinction, which as we have seen in the previous chapter
did not preclude the possibility that an extinct species could reappear.
86 DARWIN AND THE NATURE OF SPECIES

Returning to the issue of “amount of difference” in the Origin, we can

see the role it plays for Darwin in determining whether two forms connected
by intermediate gradations ought to be classified as varieties or species, when
he says “Hence, without quite rejecting the consideration of the present ex-
istence of intermediate gradations between any two forms, we shall be led to
weigh more carefully and to value higher the actual amount of difference be-
tween them” (485). The example he then gives is that of primroses and
cowslips! Similarly in Descent (1871 I) Darwin tells us that “Nevertheless it
must be confessed that there are forms, at least in the vegetable kingdom,
which we cannot avoid naming as species, but which are connected together,
independently of intercrossing, by numberless gradations” (227). In the
footnote to this passage (n. 18), he refers to examples given by Nägeli, also
Gray, but without referring to any species names. Interestingly, in a letter to
Lyell (May 8, 1860) Darwin wrote “With respect to Aster, I remember long
ago reading curious paper by Asa Gray & another on Aster, in which they
give cases of two forms so very distinct that they must consider them specif-
ically distinct, & yet perfectly united by intermediate varieties or links &
they admit that in almost every other case that this suffices to upset two
species as distinct” (Burkhardt et al. 1993, 196).2
What “amount of difference” in Darwin’s theory of classification
largely signifies, then, especially for his species concept, is that similarity
relations must be viewed as constitutive of taxa. Again, this conclusion is
entailed alone by Darwin’s view that extinction is not necessarily (logically)
forever. Nevertheless this is not to say that Darwin thought of species in
terms of overall similarity. Scholars such as Sulloway (1979, 43), Mayr
(1982, 210, 1991, 30), and Ruse (1987, 344) have interpreted Darwin this
way, but such a view cannot possibly hold up. Granted, in the Origin Dar-
win says “I look at the term species, as one arbitrarily given for the sake of
convenience to a set of individuals closely resembling each other” (52). But
this is Darwin speaking at his loosest, and possibly also strategically (cf.
chapter 8). When we look more closely, at his chapter on classification, for
example, we have seen in the previous chapter that he disassociates himself
from the view that classification is “placing together the forms most like
each other” (414), “the mere putting together and separating objects more
or less alike” (420). At the species level this was especially evident in his
work on barnacles. When Darwin discovered what he came to call “com-
plemental males,” males so small in relation to the hermaphrodites to which
they were attached (such that naturalists had thought they were parasites of
the latter), he nevertheless thought of them, in spite of their being “utterly
different in appearance and structure” from the hermaphrodites, as
 Natural Selection and the Unity of Science 87

“belonging to the same species!” (Darwin 1851b, 293). In the Origin the
same conclusion is maintained, and as we have seen in the previous chap-
ter other kinds of examples are added to illustrate the dominance of com-
mon descent over similarity, such as larval and adult stages of the same
individual (424). In addition, we have seen in the previous chapter what
Darwin thought of convergence, even at the species level. So although Dar-
win included similarity as constitutive of species, his concept was far from
that of overall similarity. Common descent was one of the relations that
put bounds on the similarity complexes. Another was the primacy of the
horizontal dimension, which, along with common descent, intergrading,
and amount of difference, helped to distinguish species from varieties.
Nevertheless, if the only difference between primroses and cowslips
being two species and the races of man being one species is amount of dif-
ference, then amount of difference sounds pretty arbitrary. And if that’s
what Darwin’s species concept comes down to, then it’s not much of an
objective species concept. This is a valid criticism. But it turns out that
there is much more involved in Darwin’s species concept than mere
amount of difference.
Part of what is involved is whether the amount of difference is relatively
“constant and distinct.” Early in the Origin Darwin wrote that “Those forms
which possess in some considerable degree the character of species, but which
are so closely similar to some other forms, or are so closely linked to them
by intermediate gradations, that naturalists do not like to rank them as dis-
tinct species, are in several respects the most important for us. We have every
reason to believe that many of these doubtful and closely-allied forms have
permanently retained their characters in their own country for a long time;
for as long, as far as we know, as have good and true species” (47). Similarly
in his concluding chapter he tells us that as a consequence of his view “Sys-
tematists will only have to decide (not that this will be easy) whether any
form be sufficiently constant and distinct from other forms, to be capable
of definition; and if definable, whether the differences be sufficiently
important to deserve a specific name” (484).
What makes the difference, then, what overrides both intermediate
gradations and common descent in Darwin’s species concept, is the fur-
ther criterion of “constant and distinct” characters, characters as “perma-
nent” (though not in the creationist sense) as the characters that demarcate
“good and true” species. This is largely why primroses and cowslips, in Dar-
win’s view, are two species instead of one. Indeed in Darwin’s view in the
Origin species of all kinds are only “well-marked and permanent varieties”
(133), or to vary the expression “only strongly marked and fixed varieties”
88 DARWIN AND THE NATURE OF SPECIES

(155). This, of course, given Darwin’s evolutionism, in particular his grad-

ualism, only makes sense in the horizontal dimension. From the vertical
perspective, in Darwin’s view, with the exception of living fossils there are
no permanent or fixed characters.
Although an improvement over mere “amount of difference,” charac-
ters “constant and distinct” might nevertheless also seem largely arbitrary,
given Darwin’s gradualism. In reply, however, it might be pointed out that
in the passages from the Origin quoted in the second paragraph above Dar-
win is dealing with borderline cases in the horizontal dimension, what are
often today called “messy situations.” Although such borderline cases are
entailed by Darwin’s view of evolution by natural selection, we saw in
c hapter 3 that Darwin believed that in most situations in the horizontal
dimension species are “tolerably well-defined objects,” that they “do not
at any one period present an inextricable chaos of varying and intermedi-
ate links” (177). Thus the existence of “good and true” species (horizontal)
in Darwin’s view is no more arbitrary, unreal, or invalidated than dusk and
dawn, for example, invalidate the reality of night and day.
The problem with this reply, however, is that in his correspondence
Darwin cited the “constant and distinct” criterion as his main criterion for
species delimitation. For example, in a letter to Hooker (July 30, 1856), on
the topic of barnacles, Darwin wrote “I differ from him [Lyell] in thinking
that those who believe that species are not fixed will multiply specific names:
I know in my own case my most frequent source of doubt was whether
others would not think this or that was a God-created Barnacle & surely de-
served a name. Otherwise I shd. only have thought whether the amount of
difference & permanence was sufficient to justify a name” (Burkhardt and
Smith 1990, 194). Moreover, years later in a letter to Hooker (October
22, 1864) Darwin wrote “The power of remaining for a good long period
constant, I look at as the essence of a species, combined with an apprecia-
ble amount of difference” (Burkhardt et al. 2001, 376). Citing this last pas-
sage, Poulton (1903) claimed that “Darwin regarded persistence of form as
an important criterion of a species” (91).
And yet in spite of this passage in Darwin’s letter, as well as what we
have seen in the Origin, there is still something more to Darwin’s “amount
of difference” than just relatively constant and distinct characters. Indeed
to think otherwise is to miss the key that unlocks a full understanding of
Darwin’s species concept. In short, it turns out that the crucial difference,
for Darwin, is whether those characters have been established by natural se-
lection; in other words, whether they are adaptations. In the case of prim-
roses and cowslips, Darwin reveals early in the Origin his reasons for going
 Natural Selection and the Unity of Science 89

against his fellow naturalists and ranking them as two distinct species
instead of one: “These plants differ considerably in appearance; they have
a different flavour and emit a different odour; they flower at slightly differ-
ent periods; they grow in somewhat different stations; they ascend moun-
tains to different heights; they have different geographical ranges; and lastly,
according to very numerous experiments made during several years by that
most careful observer Gärtner, they can be crossed only with much diffi-
culty. We could hardly wish for better evidence of the two forms being
specifically distinct” (49–50). The criterion that Darwin lists last here, as
we shall see in the next chapter, is a red herring, and can only have been
thrown in by Darwin for persuasive effect. Darwin’s real reasons are the
ones he lists before it. Although Darwin does not come out and say it, his
conclusion can only be, given that his book is about the Origin of Species
by Means of Natural Selection, that the characters which distinguish prim-
roses and cowslips as two species in his view are characters that have been
established by means of natural selection. That he does not come out and
say this is because in the Origin at that point he has not yet focused on nat-
ural selection; that focus comes two chapters later.
In the case of the races of man, on the other hand, Darwin in Descent
(1871 I) is more explicit. In the penultimate passage he tell us that

as far as we are enabled to judge (although always liable to error on this

head) not one of the external differences between the races of man are of
any direct or special service to him. . . . The variability of all the charac-
teristic differences between the races, before referred to, likewise indi-
cates that these differences cannot be of much importance; for, had they
been important, they would long ago have been either fixed and pre-
served, or eliminated. In this respect man resembles those forms, called
by naturalists protean or polymorphic, which have remained extremely
variable, owing, as it seems, to their variations being of an indifferent
nature, and consequently to their having escaped the action of natural
selection. [248–249]

This is the key. We can see it in the title of Darwin’s most famous
book, On the Origin of Species by Means of Natural Selection. Moreover in
the Origin Darwin tells us what the connection is between “constant and
distinct characters” and natural selection, the connection being that natural
selection is primarily responsible for the fixed state. Beginning (as Darwin
typically does) with the evidence from domesticated animals, he says “if
any part, or the whole animal, be neglected and no selection be applied, that
part (for instance, the comb in the Dorking fowl) or the whole breed will
90 DARWIN AND THE NATURE OF SPECIES

cease to have a nearly uniform character. The breed will then be said to
have degenerated” (152). Looking to nature, he then goes on to say “In
rudimentary organs, and in those which have been but little specialized for
any particular purpose, and perhaps in polymorphic groups, we see a nearly
parallel natural case; for in such cases Natural Selection either has not or
cannot have come into full play, and thus the organisation is left in a fluc-
tuating condition” (152). Again he says “That the struggle between Natural
Selection on the one hand, and the tendency to reversion and variability on
the other hand, will in the course of time cease; and that the most abnor-
mally developed organs may be made constant, I can see no reason to
doubt” (153–154). Granted, though Darwin then goes on in the next few
pages to argue that “specific characters are more variable than generic”
(154), the point remains for him that species are “only strongly marked
and fixed varieties” (155). This, moreover, will prove significant when, in
chapter 7, we examine Darwin’s view on the nature of varieties and how
they differ from species.
As the key to Darwin’s species concept, we can now also see why he
would give such high though not exclusive importance to common descent
in his species concept, as we have seen in c hapter 4. Common descent
means common inheritance, and it is by common inheritance that adapta-
tions get shared. Conversely, it was for Darwin highly unlikely that the
same adaptation could arise separately by saltation, as we shall see a little
later in this chapter, or that adaptations could evolve separately in differ-
ent lines of descent, as we have seen in the previous chapter on the topic of
convergence at the species level and as we shall see again in the next chap-
ter when we look at Darwin on niches.3
As the key we can also see why Darwin would think of species primar-
ily though not exclusively as horizontal rather than vertical entities, as we
have seen in chapter 3. Species ontologically for Darwin could not possi-
bly be lineages or clades because adaptations come and go over time in a sin-
gle lineage or clade. When remaining relatively unchanged, however, their
reality could be extended over long periods of time, as with living fossils or
species F in Darwin’s diagram in the Origin.
Returning to the matter of examples, so far we have looked at the cases
of primroses and cowslips and the races of man. There are further cases that
not only go against Beatty’s claims but further support the interpretation
developed above. A prime example is the case of domestic pigeons. In a let-
ter to Huxley (December 13, 1859) Darwin supplied an enclosure he had
written for Huxley on pigeon breeding. In that enclosure he included some
quotations from the writings of John Matthews Eaton. Although Eaton
 Natural Selection and the Unity of Science 91

preferred to refer to domestic pigeons as “species,” Darwin in his enclosure

referred to the “Breeds” of pigeons, remarking that if Huxley could see the
drawings which Darwin has he “would have grand display of extremes of di-
versity” (Burkhardt and Smith 1991, 428). Why would Darwin group all
the breeds of pigeons together as one species (as he did in the Origin, 20–28),
contrary to the breeder Eaton, if he thought that the designation of species
and varieties was arbitrary? This raises an interesting question. As we have
seen earlier in this chapter, Darwin ranked primroses and cowslips as two
species instead of one, even though they were derived from a common stock.
In the case of domestic pigeons, with their great diversity, Darwin tells us in
the Origin “Altogether at least a score of pigeons might be chosen, which if
shown to an ornithologist, and he were told that they were wild birds, would
certainly, I think, be ranked by him as well-defined species. Moreover, I do
not believe that any ornithologist would place the English carrier, the short-
faced tumbler, the runt, the barb, pouter, and fantail in the same genus”
(22–23). Darwin also believed that all the pigeon breeds derived from a
common stock, the rock pigeon, Columba livia (23), and yet, unlike prim-
roses and cowslips, and in spite of their great diversity, he did not rank any
of the breeds of domestic pigeons as specifically distinct. So if common de-
scent and being well-defined are not sufficient criteria for a species distinc-
tion, and they clearly are not in this case, then what for Darwin is? Again the
answer appears to fit quite perfectly with my theory above. Contrary to the
characters that functionally distinguish primroses and cowslips, which as we
saw Darwin thought were adaptations produced by natural selection, the
characters that distinguish pigeon breeds were produced by artificial selec-
tion, the product of mere human fancy, and thus were functionally super-
ficial. As Darwin put it in Variation (1868 I) at the end of his two chapters
on pigeons, “It is not likely that characters selected by the caprice of man
should resemble differences preserved under natural conditions either from
being of direct service to each species, or from standing in correlation with
other modified and serviceable structures” (233).
This connects with what Darwin wrote in the Origin. Although there
he does indeed state that varieties or races produced by artificial selection
“show adaptation in their structure or in their habits to man’s wants or fan-
cies” (38, italics mine), he nevertheless adds that artificial selection focuses
only on “external and visible characters,” whereas natural selection acts “on
every shade of constitutional difference” (83). Furthermore, “Man selects
only for his own good; Nature only for that of the being which she tends.”
Finally, the wishes and efforts of man are “fleeting,” his time is “short,” so
that “how poor will his products be, compared with those accumulated by
92 DARWIN AND THE NATURE OF SPECIES

nature during whole geological periods” (84). Thus, for Darwin, “nature’s
productions should be far ‘truer’ in character than man’s productions” and
“should be infinitely better adapted to the most complex conditions of life”
(84). Unlike domestic productions, then, which are produced by artificial
selection and accordingly are superficial, and furthermore are produced not
for the good of the organism but only for mere human fancy, natural
selection produces genuine adaptations and consequently real species.
Returning to organisms in a state of nature, there are yet further exam-
ples that go against Beatty but that support the above view. For example,
in the case of primroses, George Bentham, in his Handbook of British Flora,
published in 1858, made Primula scotica a variety of Primula farinosa, but
Darwin seemed to disagree. As he wrote in a letter to Daniel Oliver (Feb-
ruary 20, 1863), “would it not be worth while to tell him of Mr Scotts ob-
servation; for there can be no doubt that this difference indicates an
important functional difference” (Burkhardt et al. 1999, 159). Indeed in his
book on cross-fertilization Darwin (1876b) followed Lindley and treated P.
scotica as a species (362).
Further examples come from Darwin’s book on orchids (1862a). Right
from the start, Darwin states “The object of the following work is to show
that the contrivances by which orchids are fertilized, are as varied and al-
most as perfect as any of the most beautiful adaptations in the animal king-
dom” (1). Moreover in the above letter to Oliver, Darwin wrote
“Treviranus in his Review of the Orchids does not seem to appreciate at all
the prettiness of the adaptations, which seems to me the cream of the case”
(159). Adaptations also appear in that work as the key for specific identity
in orchids, which sometimes set Darwin against the species designations of
his fellow naturalists. A good example is the case of Habenaria chlorantha,
which Bentham ranked as a variety of H. bifolia. In a letter to Bentham
(June 22, 1861), Darwin wrote “I must think that you are mistaken in
ranking Hab. chlorantha as a var. of H. bifolia: the pollen-masses & stigma
differ more than in most of the best species of Orchis” (Burkhardt et al.
1994, 185). And indeed in his orchid book (1862a) Darwin explicitly calls
these characters, which he used to treat these forms as distinct species, “cases
of adaptation” (44, cf. 69–73).
Another orchid example is Ophrys arachnites, which Darwin tells us “is
considered by some botanists as only a variety of the Bee Ophrys [O. apifera],
by others as a distinct species” (51). He then goes on to point out the struc-
tural differences, stating in conclusion that “This plant, therefore, differs in
every important respect from O. apifera, and seems to be much more closely
 Natural Selection and the Unity of Science 93

allied to O. aranifera” (52), the implication being that Darwin himself

considers O. arachnites a species because of its unique adaptations.
A related example is given in Darwin’s description of Ophrys apifera,
namely, that in “North Italy Ophrys apifera, aranifera, arachnites, and
scolopax are connected by so many and such close intermediate links, that
all seem to form a single species in accordance with the belief of Linnaeus,
who grouped them all together under the name of Ophrys insectifera”
(58–59). Darwin then goes on to implicitly split them. First, he says “The
case therefore is an interesting one, as here we have forms which may be and
generally have been ranked as true species, but which in North Italy have
not as yet been fully differentiated.” He then adds “Whether we rank the
several forms of Ophrys as closely allied species or as mere varieties of the
same species, it is remarkable that they should differ in a character of such
physiological importance as the flowers of some being plainly adapted for
self-fertilization, whilst the flowers of others are strictly adapted for cross-
fertilization, being utterly sterile if not visited by insects” (59).
Further examples come from cases of sudden origin (saltation), which
some naturalists took to be true species but which Darwin in every case re-
fused to accept. In chapter 3 (note 2) we have seen Darwin’s response to
William Harvey’s case of Begonia frigida, which Darwin recognized as a
distinct form but never once called a “species.”
Of equal if not greater relevance are two cases discussed in Variation
(1868 I). The first is that of the Himalayan breed of rabbit, a distinct white
breed with red eyes and brownish-black ears, nose, feet, and upper side of
the tail, which, it was found, “notwithstanding their sudden origin, if kept
separate, bred perfectly true,” so that “Some good observers . . . stoutly
maintained that they formed a new species” (113), giving them the name
Lepus nigripes. Darwin, however, rejected their specific status, dismissing
them as a case of “reversion” to the “not improbable” originators of their
silver-grey parental species, namely, a supposed cross between “a black and
albino variety” (115).
The second case from Variation is that of the “japanned” or “black-
shouldered” form of peacock (peafowl), which appeared independently in
Great Britain from common peafowl in at least seven well-authenticated
cases (306–307), which “propagate their kind quite truly” (305), and which
one “high authority,” Philip Sclater, named a “distinct species, viz. Pavo
nigripennis” (305). Darwin, however, granting that he has “heard of no
other such case in the animal or vegetable kingdom,” argues that the form
is a “strongly marked variety or ‘sport,’ which tends at all times and in many
94 DARWIN AND THE NATURE OF SPECIES

places to reappear” (307). Interestingly, a number of years earlier, in a let-

ter to Sclater (May 14, 1862), Darwin wrote “With four cases now recorded
I would wager the P. nigripennis will prove a variety,—hardly more surpris-
ing in its origin, than the so-called Himalayan rabbit” (Burkhardt et al.
1997, 193).
The above three cases provide an interesting set of examples since, in
arguing against the species ascriptions of his fellow naturalists, Darwin is
not appealing to adaptations, as in the cases of primroses, cowslips, and or-
chids, but to the lack of new adaptations, as in the cases of humans and pi-
geons. But unlike the latter cases, the origins of the forms in the previous
three paragraphs were by saltation. And we know from the Origin that Dar-
win thought it highly unlikely that an adaptation would ever result by salta-
tion. Monstrosities, he said, which he took to mean “some considerable
deviation of structure in one part,” are either, he continued, “injurious to
or not useful to the species” and are “not generally propagated” (44). In-
stead for Darwin it was slow, gradual, cumulative natural selection that
produced adaptations, not saltations. Again as he put it in the Origin, this
time appealing to the gradualism and uniformitarianism of the “modern ge-
ology” of Lyell, “so will natural selection, if it be a true principle, banish the
belief of the continued creation of new organic beings, or of any great and
sudden modification of their structure” (95–96; cf. Stauffer 1975, 223).
Indeed Darwin just could not see adaptations arising by saltation. As he
put it in a letter to Gray (August 11, 1860), “Please tell him [Prof. Par-
sons] that I reflected much on chance of favourable monstrosities (i.e. great
& sudden variations) arising. I have, of course, no objection to them; in-
deed it wd. be great aid; but I did not allude to subject [in the Origin], for
after much labour I could find nothing which satisfied me of the probabil-
ity of such occurrences. There seems to me in almost every case too much,
too complex, & too beautiful adaptation in every structure to believe in its
sudden production” (Burkhardt et al. 1993, 317). Indeed to think other-
wise, for Darwin, smacked of miracles and creationism, not formation by
natural law.
In sum, all of the above examples should be sufficient to put to rest
the view that Darwin simply followed what his fellow naturalists called
“species.” In the next chapter I shall give an explicit reconstruction of Dar-
win’s species concept, in the form of a definition, after examining what he
did not include in it. For the remainder of this chapter I want to focus on
Darwin’s species concept, characterized as we have seen mainly by natural
selection and adaptation, as fulfilling the vera causa (true cause) ideal of the
philosophy of science of his day.
 Natural Selection and the Unity of Science 95

Darwin’s world, as is well known, the world of the science of his day,
was a world characterized by discovering laws of nature. Darwin did not re-
ject this world but fully embraced it. Indeed Darwin himself defined nature
in terms of laws. For example, in Natural Selection (Stauffer 1975) Darwin
wrote “By nature, I mean the laws ordained by God to govern the Uni-
verse” (224). In Variation (1868 I) Darwin wrote, with the above deism
now noticeably missing, “I mean by nature only the aggregate action and
product of many natural laws” (7). Accordingly Darwin also defined science
in terms of laws. In his autobiography (1876a) he wrote “science consists
in grouping facts so that general laws or conclusions may be drawn from
them” (70).
Although Darwin would have simply imbibed this emphasis on laws
and the discovery of them by science in his early education, there were two
books in particular, and to a lesser extent a third, which greatly influenced
him. The third would be Lyell’s Principles of Geology, in particular the first
and third volumes, both of which were read by Darwin during his Beagle
voyage. In those volumes Lyell repeatedly appealed to laws of geology in his
argument for the gradual transformation of geological landscapes and seas,
including fossils, all of which appealed to processes operating in the here
and now. In the first volume (1830), for example, Lyell states right at the
outset that “Geology . . . enquires into the causes of these changes,” the
changes being those that have taken place both in the organic and inor-
ganic worlds, and that it aims at discovering “the laws now governing its an-
imate and inanimate productions” (1). In the third volume (1833) Lyell
gives a specific example, that of “the laws of earthquakes” which throw light
on the “origin of mountains” (5). Interestingly Lyell claimed that geology
should be restricted “in the first instance, to known causes, and then spec-
ulating on those which may be in activity in regions inaccessible to us” (3).
But by far the greatest influence on Darwin for the topic of laws was
John Herschel’s A Preliminary Discourse on the Study of Natural History
(1830). Darwin began reading this book shortly after finishing his B.A. at
Cambridge in late January 1831, almost a year before beginning his Beagle
voyage (begun on December 31, 1831, and finished on October 2, 1836).
Darwin read the book again, according to his reading notebooks, in 1838
(Burkhardt and Smith 1988, 456), very shortly after he first read Malthus
(September 28 to October 3), which of course helped generate his earliest
formulation of natural selection (Barrett et al. 1987, 375–376). In his au-
tobiography (1876a) Darwin wrote that Herschel’s book, along with Hum-
boldt’s Personal Narrative of Travels, “stirred up in me a burning zeal to
add even the most humble contribution to the noble structure of natural
96 DARWIN AND THE NATURE OF SPECIES

science. No one or a dozen other books influenced me nearly so much as

these two” (68).4
Herschel’s book was quickly recognized by his contemporaries as the
authoritative treatise on scientific method. What is especially important for
our purposes is what Herschel wrote about causes and laws. He argued that
the aim of science is to find explanations in terms of “an immediate pro-
ducing cause,” and that “If that cannot be ascertained, the next is to gener-
alize the phenomenon, and include it, with others analogous to it, in the
expression of some law, in the hope that its consideration, in a more ad-
vanced state of knowledge, may lead to the discovery of an adequate prox-
imate cause.” Explicitly following Newton, Herschel applied the term “veræ
causæ,” by which he meant “causes recognized as having a real existence in
nature, and not being mere hypotheses or figments of the mind” (144).
A further important influence on Darwin’s concept of laws of nature
came from William Whewell’s History of the Inductive Sciences (1837),
which from Darwin’s reading notebooks we know he had read in 1838
(Burkhardt and Smith 1988, 456), having previously skimmed parts of it
(Burkhardt and Smith 1986, 186), and which again (his full reading) was
very shortly after he read Malthus. Darwin was possibly also influenced by
Whewell’s Philosophy of the Inductive Sciences (1840), which we know from
his reading notebooks he had planned to read, noting apparently in 1841,
after reading a review of it by Herschel, that “I see I must study” (Burkhardt
and Smith 1988, 446), but we don’t know if he even cracked the book
open at all.
Whewell differed from Herschel in not requiring direct or even analog-
ical evidence of verae causae, but instead emphasized a consilience of induc-
tions. By this term he referred to the proof of a vera causa being obtained
when it explains inductions (generalizations) from a wide variety of areas.
As Michael Ruse (1979, 176–180) points out, Darwin employed both
approaches to verae causae, the direct and analogical conception of Her-
schel and Lyell and the consilience conception of Whewell. In the Origin
the Herschelian approach is to be found in the first chapter, with Darwin’s
examination of variation under domestication and artificial breeding. This
Darwin called “the best and safest clue” (4) for understanding the origin of
species in nature. Darwin then goes on to make the case for natural selec-
tion in nature, and further for its production of varieties from chance vari-
ations in individuals and for species from varieties. In the remainder of the
Origin, Darwin employed the Whewellian approach to verae causae, amass-
ing evidence from a widely different group of facts (instincts, the fossil
record, biogeography, homology, embryology, rudimentary organs, and
 Natural Selection and the Unity of Science 97

non-evolutionary classifications), each of them an induction and collec-

tively a consilience of inductions (cf. Ruse 1975; Hodge 1977; and Ruse
2000 vs. Hodge 2000 on Darwin’s debt to Whewell). Darwin, however,
did not explicitly bring out this feature of his argument in the Origin, gloss-
ing it over instead by stating that “this volume is one long argument” (459;
cf. 5–6).
Ruse (1979, 235–236) points out that as time went on and Darwin’s
“best and safest clue” came more and more under attack, Darwin relied in-
creasingly on the Whewellian conception of verae causae. In fact, however,
Darwin began his repeated appeal to consilience slightly before the publi-
cation of the Origin, commencing with his letters to those who received
advance copies, and then continuing this appeal for quite some time (cf.
Burkhardt and Smith 1991, 369, 374, 403, 404, 422, 435, 449, 460,
Burkhardt et al. 1993, 76, 84, 91, 123, 129, 137, 195, 266, Burkhardt et
al. 1994, 52, 54, 96, 99, 108, 135, Burkhardt et al. 1997, 255, Burkhardt
et al. 1999, 155). Perhaps the best example of these passages is Darwin’s let-
ter to Samuel Pickworth Woodward (March 6, 1860), for the way he uses
both concepts of vera causa:

The fair way to view the argument of my book, I think, is to look at Nat-
ural Selection as a mere hypothesis (though rendered in some degree
probable by the analogy of method of production of domestic races; &
by what we know of the struggle for existence) & then to judge whether
the mere hypothesis explains a large body of facts in Geographical Dis-
tribution, Geological Succession, & more especially in Classification, Ho-
mology, Embryology, Rudimentary Organs The hypothesis to me does
seem to explain several independent large classes of facts; & this being so,
I view the hypothesis as a theory having a high degree of probability of
truth. [Burkhardt et al. 1993, 123]

Darwin would later write, in a letter to Alexander Goodman Moore (March

8, 1861), comparing the matter to the belief in ether and the undulatory
theory of light, “which is now universally admitted as a true theory,” that
similarly because evolution by natural selection explains many diverse
classes of facts “therefore I believe in its truth” (Burkhardt et al. 1994, 50).5
Returning to the influence of Herschel, Silvan Schweber (1985) has
suggested that there were many other aspects of Herschel’s influence on
Darwin besides the vera causa ideal, including that for Herschel good sci-
ence involves the decompounding and resolution of complex phenomena
into simpler ones, as in chemistry with elements, so that, as Schweber puts
it, “I believe Darwin was sensitive to these remarks of Herschel when he
98 DARWIN AND THE NATURE OF SPECIES

pondered what constituted an ‘element’ in his theory of natural selection.

Were individuals the ‘elements,’ or were varieties and species ‘elements’?”
(48). Beyond this tantalizing glimpse, however, apparently related to the
issue of the units of selection, Schweber ventures no further. What I would
like to add at this point is a further influence that Schweber might have
glimpsed but alas has missed, along with Ruse and Hodge and everyone
else, namely, the effect on Darwin’s mature species concept of what Her-
schel wrote on the relation between verae causae and natural classification.
It will no doubt be helpful to begin with a quick look at Herschel’s
(1830) concept of laws of nature, since as we have seen he intimately con-
nects it with the concept of vera causa. Herschel’s concept will prove to il-
luminate Darwin’s concept of laws as revealed in the examples examined in
chapter 2 but especially with the example examined so far in this chapter,
namely, natural selection.
Herschel had two great categories of laws that he subsumed under laws
of nature and that he stated as follows: “We may therefore regard a law of
nature either, 1st, as a general proposition, announcing, in abstract terms,
a whole group of particular facts relating to the behavior of natural agents
in proposed circumstances; or, 2dly, as a proposition announcing that a
whole class of individuals agreeing in one character agree also in another”
(100). The latter he called “the relation of constant association, inasmuch as
it asserts that in whatever individual the one character is found, the other
will invariably be found also” (101). The other category is much more im-
portant for our purpose (and also for Herschel), namely, “the consideration
of a proximate, if not an ultimate, cause” (101). The classic example, of
course, for Herschel as well as for others was Newton’s law of gravity, which
Herschel called both a “force” and a “real cause” (198). Causal laws for
Herschel are the explanatory laws (144), and he conceived of these as being
essentially conditional in nature, such that “if such a case arise, such a course
shall be followed” (36). More importantly he conceived of these as being
eternally fixed, applying to kinds that might not yet or that might not ever
have any members, as with chemical elements and compounds, such that
“no chemist can doubt that it is already fixed what they will do when the
case does occur. They will obey certain laws, of which we know nothing at
present, but which must be already fixed, or they could not be laws” (36).
Compare all of this now with Darwin’s concept of laws, including espe-
cially natural selection. In his abstract of Macculloch’s Proofs and Illustrations
of the Attributes of God (Barrett et al. 1987, 632–641), written in late 1838,
he complains about using the deity to explain structures in organisms: “The
explanation of types of structure in classes—as resulting from the will of the
 Natural Selection and the Unity of Science 99

deity, to create animals on certain plans.—is no explanation—it has not the

character of a physical law, & is therefore utterly useless—it foretells nothing
because we know nothing of the will of the Deity, how it acts & whether
constant or inconstant like that of Man.—the cause given we know not the
effect” (55). Here we see both the importance of explanation and prediction,
themes that would recur throughout Darwin’s writings where he mentions
laws. For example, in his autobiography (1876a), in criticism of Herbert
Spencer’s laws, Darwin wrote “they do not seem to me to be of any strictly
scientific use. They partake more of the nature of definitions than of laws of
nature. They do not aid one in predicting what will happen in any particu-
lar case” (109). Similarly Paley’s explanation of adaptations “fails, now that
the law of Natural Selection has been discovered.” Indeed Darwin immedi-
ately adds that “Everything in nature is the result of fixed laws” (87). We can
also see the conditional nature in Darwin’s concept of natural selection as
law, in for instance his letter to Charles Bunbury (February 9, 1860), in which
Darwin dealt explicitly with whether natural selection is a “vera causa,” and
in which he wrote “Natural selection seems to me in so far in itself not be
quite hypothetical, in as much if there be variability & a struggle for life, I
cannot see how it can fail to come into play to some extent” (Burkhardt et al.
1993, 76). He then goes on to emphasize the importance “in giving a ratio-
nal, instead of theological explanation of many known facts” (77).
Darwin also subscribed to laws of association, as with what came to be
known as the Darwin-Knight law, that “it is a general law of nature (utterly
ignorant though we may be of the meaning of the law) that no organic
being self-fertilises itself for an eternity of generations; but that a cross with
another individual is occasionally—perhaps at very long intervals—indis-
pensable” (Darwin 1859, 97; cf. Darwin 1858, 21). But what is most im-
portant for our purposes is whether he thought of natural selection as a
force, so that the law of natural selection is a force law akin to Herschel’s
acceptance of Newtonian gravity as a force law. As we have seen in chap-
ter 2, Darwin sometimes in the Origin calls natural selection a “power,”
while other times he calls it a “process.” Darwin’s repeated use of the word
“power” might indicate that he thought of natural selection as a force anal-
ogous to gravity or magnetism, but in a letter to Gray (November 29, 1857)
Darwin made it clear that he thought of natural selection, as it is often
thought of today (e.g., Endler 1986, 29–33; Haynes 1987, 5), not as a force
but as a process, in fact as the result of a number of processes: “I had not
thought of your objection of my using the term ‘natural Selection’ as an
agent; I use it much as a geologist does the word Denudation, for an agent,
expressing the result of several combined actions” (Burkhardt and Smith
100 DARWIN AND THE NATURE OF SPECIES

1990, 492). Darwin’s use of the word “agent” is interesting in relation to

Herschel’s use, for although Herschel seemed to restrict the word “agent”
to force laws, such as Newton’s law of gravity, Darwin expanded the con-
cept to apply to process laws as well, such as natural selection.6
Darwin, of course, argued that species are produced by laws, begin-
ning early in his career as an evolutionist. In, for example, Notebook E
(Barrett et al. 1987), he says “the laws of variation of races, may be impor-
tant in understanding laws of specific change.—When the laws of change
are known.——then primary forms may be speculated on, & laws of life,—
the end of Natural History, will be approximated to” (54). Similarly in a
letter to Hooker (July 30, 1856) he wrote “I am, also, surprised at his
[Lyell’s] thinking it immaterial whether species are absolute or not: when-
ever it is proved that they all species are produced by generation, by laws
of change what good evidence we shall have of the gaps in formations. And
what a science Natural History will be, when we are in our graves, when all
the laws of change are thought one of the most important parts of Natural
History” (Burkhardt and Smith 1990, 194).
Since species for Darwin are produced by laws, in particular natural
selection (recall the title of the Origin), this connects to the issue of laws and
classification, and again to Herschel, for in Herschel (1830) we find a very
clear connection between verae causae and classification. In his fifth chap-
ter devoted to the topic, titled “Of the Classification of Natural Objects
and Phenomena, and of Nomenclature,” Herschel tells us not only, as one
would expect, that classification is an important part of science, but much
more importantly he distinguishes between natural and artificial systems of
classification. Although both classes are groups “bound together by gen-
eral resemblances” (135), the most basic of which are species (chemical and
biological), it is evident from Herschel’s chapter that the main difference
between a natural and an artificial system is that the former is based on
verae causae whereas the latter is not. Although Herschel has very little to
say about biological species—he clearly thinks of them in the same way as
chemical species, as “distinguished from each other by essential differences”
(136)—his concept of natural classification comes out quite clearly in the
case of mineral species. Lamenting that “There is no science in which the
evils resulting from a rage for nomenclature have been felt to such an ex-
tent as in mineralogy” (139), his complaint nevertheless is not so much
that the names for minerals have not been fixed, that “there is scarcely one
[mineral species] which has not four or five names in different books”
(139). Instead his main complaint is that many mineralogists raise mineral
species into genera so that their resulting mineral species are based on
 Natural Selection and the Unity of Science 101

arbitrary rather than on nomic differences, in his words that “species too
definite to admit of mistake, are actually rendered generic, and extended to
whole groups, comprising objects agreeing in nothing but the arbitrary
heads of classifications from which the most important natural relations
are professedly and purposely rejected” (140).
This is the equivalent of the complaint against splitters in biology, as
opposed to lumpers, a debate that raged in Darwin’s time (and continues
in our own).7 In the case of mineral species, it is evident that the “most im-
portant relations” that Herschel thought the splitters overlook are the laws
of crystallography. These laws play a repeated and prominent role through-
out Herschel’s Discourse. In his section on mineralogy he tells us that crys-
tallography, with the aid of inventions such as the reflecting goniometer,
“furnished the mineralogist with the means of reducing all the forms of
which a mineral is susceptible under one general type, or primitive form”
(292). Accordingly, each mineral species has a peculiar “cleavage,” a char-
acter, he tells us, “in the highest degree geometrical, and affording, as might
naturally be supposed, the strongest evidence of its necessary connection
with the intimate constitution of the substance” (291). He further tells us
that until such cleavages were discovered by crystallography, the main name
being Abbé Haüy (240), “no mineralogist could give any correct account
of the real distinction between one mineral and another” (291). In his sec-
tion on crystallography (239–245) he provides a short history, leading up
to the present state of the science, repeatedly referring to the “laws of crys-
tallography,” which he earlier referred to as “laws . . . which limit the forms
assumed by natural substances . . . to precise geometrical figures, with fixed
angles and proportions” (123). Nevertheless these laws are only laws of
“constant association,” to use his term examined above, so that, even
though they allow a certain amount of novel prediction (240), they can
only point to causal/mechanical laws that are not yet known (245). Even
so, crystallography in Herschel’s view developed to the point where it could
legitimately be called a “separate and independent branch of science,” since
it applies not only to mineralogy but to other areas, notably chemistry and
optics (293).
This lesson could hardly have been lost on Darwin. Not only had Dar-
win read Herschel’s Discourse twice, and not only had Lyell emphasized the
importance of crystallography for geology (Lyell 1830, 2, 1833, 66), but the
history of mineralogy (including the importance of Haüy, the nature of
cleavages, and the laws of crystallography) is repeated in even greater detail
in Whewell’s History (1837). Even during his student days at Cambridge,
Darwin must have known about the laws of crystallography, either from his
102 DARWIN AND THE NATURE OF SPECIES

main mentor John Stevens Henslow, professor of botany but previously

professor of mineralogy (indeed Henslow had taught Darwin’s brother
Erasmus mineralogy), or from Adam Sedgwick, professor of geology, from
whom Darwin received a crash course in geology and with whom he went
on a field trip just before his Beagle voyage. Indeed, Darwin had even
brought with him on the Beagle a reflecting goniometer, though he was not
expert enough at using it and wrote to Henslow for a book to help
(Burkhardt and Smith 1985, 251, 252 n. 3, 352, 395; cf. 3 for an early
reference to a “small ganiometer” by Erasmus so that “we shall be able to
separate the different crystalls in your cab”).
Darwin, of course, went on to become one of the greatest geologists of
his time, although the public normally thinks of him only as a theoretical
biologist. At any rate, the point that needs to be made is that Darwin, by
appealing to the vera causa of natural selection in the origin of species, as
well, as we have seen in the previous chapter, to other verae causae such as
“the vera causa of community of descent,” was attempting to do for biolog-
ical classification what others had done for other sciences, namely, he was
applying laws to distinguish real from unreal species.8 Or more generally,
by employing the vera causa ideal Darwin was attempting to bring biol-
ogy, both classification and explanation of the origin of species, into the
unity of science.
Indeed comparisons between Herschel on crystallography and Darwin
on natural selection and classification abound. One further comparison is
that Herschel, as we have seen above, argued that the laws of crystallogra-
phy apply to other sciences such as chemistry and optics, not only to crys-
tallography, while in chapter 3 we saw that Darwin applied the vera causa
of natural selection to linguistics, not just to biology. Of course there are
also profound differences. For one, Herschel was an essentialist with re-
gard to species, whether chemical, mineral, or biological, whereas Darwin
was not with regard to biological species. Moreover, genealogy means noth-
ing for the delimitation of species in chemistry or mineralogy, whereas for
Darwin, as we have seen in the previous chapter, it meant a lot indeed.9 But
to focus on such differences is to miss what is really important, to miss what
Darwin was really trying to do. In this vein Darwin wrote in a letter to
Lyell (August 21, 1861) that “in the case of species . . . their formation has
hitherto been viewed as beyond law,—in fact this branch of science is still
with most people under its theological phase of development” (Burkhardt
et al. 1994, 238), which compares with what he wrote to Lyell a few weeks
earlier (August 1), that “I must think that such views of Asa Gray & Her-
 Natural Selection and the Unity of Science 103

schel merely show that the subject in their minds is in Comte’s theological
stage of science” (226–227).
We can see, then, that Darwin connected one of his fundamental in-
sights, that adaptations are produced by natural selection, with the reality
of species (and also varieties, as we shall see in chapter 7). He could have
easily titled his book On the Origin of Adaptations by Means of Natural Se-
lection. But he didn’t. He titled it On the Origin of Species because he really
did believe in the objective existence of species. Darwin’s focus, then, was
not just on adaptations, but on the broader framework of evolution, and for
that he needed not only adaptations but varieties and species. Character
evolution was not simply his focus, and it could not be, because he also
wanted to argue for and explain the branching evolution of species.
So Darwin connected the production of adaptations by natural selec-
tion with the reality of species. But he did more. By employing the vera
causa law of natural selection to explain both the existence of adaptations
and the reality of species, Darwin brought evolutionary biology (and ulti-
mately the rest of biology) into the framework of the natural sciences, which
were based on laws.
An interesting question remains, however, whether Darwin thought
that natural selection, and only natural selection, produces adaptations.
There is, of course, the passage at the very end of the Introduction to the
Origin in which Darwin says “I am convinced that Natural Selection has
been the main but not exclusive means of modification” (6). This was a
claim that Darwin repeated throughout his correspondence. One might
naturally think, then, that Darwin was not a real Darwinian, and one some-
times hears this, because he allowed for the inheritance of acquired charac-
teristics, not in the case of mutilations but in the cases of habit and use and
disuse. But one has to keep in mind that modification is a much wider con-
cept than adaptation, so that Darwin’s claim above does not necessarily
commit him to allowing processes other than natural selection a causal role
in the production of adaptations.
This is indeed a contentious issue. Robert Richards (1992), for exam-
ple, contends that right from the beginning Darwin allowed for a number
of mechanisms to explain adaptations, but that after he hit on natural se-
lection these mechanisms, as in the Origin, could no longer be central, and
were retained only as “auxiliary mechanisms” (84). Richards goes on to say,
however, that Darwin “asserted that acquired habit, like selection, could
act more immediately to introduce adaptations; but he thought his primary
device would always retain the upper hand” (n. 43). Richards goes too far,
104 DARWIN AND THE NATURE OF SPECIES

however. The passage from the Origin that he cites in support of his view
is the following: “On the whole, I think we may conclude that habit, use,
and disuse, have, in some cases, played a considerable part in the modifi-
cation of the constitution, and of the structure of various organs; but that
the effects of use and disuse have often been largely combined with, and
sometimes overmastered by, the natural selection of innate difference”
(142–143).
This passage is hardly a conclusive statement about the ability of
Lamarckian mechanisms alone for “introducing” adaptations. In fact, Dar-
win in the above passage does not even mention adaptations, but only mod-
ification, which is not the same thing. In fact there is plenty of evidence that
goes against Richards’ interpretation. Darwin always seems to emphasize
natural selection as being responsible for adaptations, possibly mixed in
some instances with Lamarckian mechanisms.
For a start, there is chapter V of the Origin, titled “Laws of Variation.”
In that chapter Darwin reasserts right from the start his belief that variations
are not produced by “chance” but have causes (laws), and he attempts to in-
vestigate the various causes, as well as the various relations between varia-
tions, such as what he called in c hapter I “the mysterious laws of the
correlation of growth” (12). It is in this chapter, on the causes of variations
(which is a preliminary matter, not to be confused with adaptations per se),
that the passage made much of by Richards is to be found. We find, for a
start, that Darwin examines the effect of climate and food on the produc-
tion of variations, and concludes that they probably have some effect (more
in plants than in animals), but also that they “cannot have produced the
many striking and complex co-adaptations of structure between one or-
ganic being and another, which we see everywhere throughout nature”
(132). He then gets into the effects of use and disuse, rejecting first “that
mutilations are ever inherited” (135). To use and disuse proper he admits
heritable variation (134), but in each and every case of adaptation he never
allows only use and disuse; instead he argues that natural selection had to
play a big part. Darwin then turns to acclimatization, to the observation
that “each species is adapted to the climate of its own home” (139). But
here Darwin says “the degree of adaptation . . . is often overrated” (139).
Moreover with regard to the ranges of species he takes “competition of
other organic beings” to be equally or even more important than climate
(140). He then questions whether habit alone could be responsible for ac-
climatization, calling it “mere habit,” and concludes with the passage we
have seen quoted by Richards. To this we should add what Darwin wrote
in the summary of chapter I, the chapter titled “Variation Under Domes-
 Natural Selection and the Unity of Science 105

tication,” that “Variability is governed by many unknown laws, more espe-

cially by that of correlation of growth. Something may be attributed to the
direct action of the conditions of life. Something must be attributed to use
and disuse. The final result is rendered infinitely complex” (43). This is all
about the causes of variation. But what has that to do with natural selection?
Everything! Since “unless profitable variations do occur, natural selection
can do nothing” (82).
There is further evidence that Darwin thought that natural selection
must be the main explanation for adaptations. There is his “Historical
Sketch” (Burkhardt et al. 1993, 572–576), added to the American edition
of the Origin and to all subsequent editions. Where Darwin discusses
Lamarck, he says “To this latter agency [‘use and disuse or the effects of
habit’] he seems to attribute all the beautiful adaptations in nature” (573).
He then adds in a footnote that his grandfather “anticipated these erro-
neous views in his Zoonomia.” Turning a little later to the author of Ves-
tiges, Darwin states that his appeal to “vital forces” and “sudden leaps”
“cannot . . . account in a scientific sense for the numerous and beautiful
co-adaptations, which we see throughout nature” (574; cf. Origin 3–4).
There is more. In a letter to Gray (September 5, 1857), for example,
Darwin wrote “The facts which kept me longest scientifically orthodox are
those of adaptation . . . . To talk of climate or Lamarckian habit produc-
ing such adaptations to other organic beings is futile. This difficulty, I be-
lieve I have surmounted. . . . I will enclose . . . the briefest abstract of my
notions on the means by which nature makes her species” (Burkhardt and
Smith 1990, 445–446). In a letter to Hooker (July 2, 1859) Darwin wrote
“to explain adaptations . . . this point has always seemed to me the turning
point of the theory of Natural Selection” (Burkhardt and Smith 1991, 316).
In a letter to William Harvey (September 20–24, 1860) he wrote “as if I
said [in the Origin] that Natural Selection was the sole agency of modifi-
cation; whereas I have over & over again, ad nauseam, directly said & by
order of precedence implied (what seems to me obvious) that selection can
do nothing without previous variability. . . . I consider Natural Selection
as of such high importance, because it accumulates successive variations in
any profitable direction; & thus adapts each new being to its complex con-
ditions of life” (Burkhardt et al. 1993, 371). Finally, in a letter to Hugh Fal-
coner (October 1, 1862) he wrote “Let me explain how it arose that I laid
so much stress on Natural Selection, and I still think justly: I came to think
from Geographical Distribution &c. &c. that species probably change; but
for years I was stopped dead by my utter incapability of seeing how every
part of each creature (a wood-pecker or swallow for instance) had become
106 DARWIN AND THE NATURE OF SPECIES

adapted to its conditions of life. This seemed to me and does still seem the
problem to solve, and I think natural selection solves it, as artificial selec-
tion solves the adaptation of domestic races for man’s use” (Burkhardt et al.
1997, 440–441).
On the other hand, interestingly, Darwin wrote in a letter to Hooker
(May 29, 1860) “Why I do not believe so much as you do in Physical agen-
cies, is that I see in almost every organism (though far more clearly in ani-
mals than in plants) adaptation, & this except in rare instance must, I shd.
think, be due to selection” (Burkhardt et al. 1993, 230).
In sum, then, on the topic of adaptations it would seem that Darwin
should be placed close to but not quite beside the modern arch-selection-
ist Richard Dawkins (1986, 288–318) in his argument that adaptive com-
plexity is explicable not by habit or use and disuse or mutation or theistic
design but “only by Darwinian selection.”
At any rate, in solving the problem of adaptation, Darwin also, in his
mind, solved the problem of both the origin and nature of species, and also
the problem of uniting biology with the rest of natural science. Adaptation
was the key in each and every case. Whether species should be so conceived,
of course, is another matter. But that is a topic to be discussed in the next
chapter, after we examine what Darwin did not include in his species concept.
 Chapter 6

Not Sterility, Fertility, or Niches

We have seen in the previous chapter, specifically in the case of primroses

and cowslips, that Darwin gave sterility between the two forms as one of his
reasons (the last) for why they should be considered specifically distinct.
But how seriously should we take this? We shall see in this chapter that we
should not take it seriously at all. The other side of the coin, fertility, which
I shall take to be shorthand for intrafertility, is we shall see no more a part
of Darwin’s species concept than sterility. In the case of primroses and
cowslips, we have also seen Darwin refer to the fact that they inhabit dif-
ferent “stations.” We shall look at what Darwin meant by this term, and we
shall see that Darwin no more thought of species as inhabitants of ecolog-
ical niches than he thought that they are intersterile or intrafertile. All of this
is interesting for a number of reasons, one of which has to do with recon-
structing Darwin’s species concept, which I shall do at the end of this chap-
ter, followed by a discussion of its strengths and weaknesses. A further
reason for interest in what Darwin has to say about the above criteria has
to do with modern species concepts, in other words, why Darwin would re-
ject modern species concepts based on reproductive isolation, or fertility,
or niches. Modern debaters on these topics make some interesting argu-
ments, but Darwin has some interesting perspectives of his own which, sur-
prisingly, seem universally to have been overlooked, perspectives that in
some cases clinch the modern critiques. This chapter, therefore, will func-
tion at a number of levels.

107
108 DARWIN AND THE NATURE OF SPECIES

Let us begin with sterility. We have seen in previous chapters, and we

shall see again, that some modern authors believe that Darwin, in his ma-
ture writings, came close to something like the modern biological species
concept. In Darwin’s day, of course, the phrase “reproductive isolation
mechanisms” was never used. Instead, sterility was the concept and it was
a common idea. Lyell (1832) put it clear enough as “nature has forbidden
the intermixture of the descendants of distinct original stocks, or has, at
least, entailed sterility on their offspring, thereby preventing their being
confounded together” (19). A little earlier Prichard (1813) raised the ques-
tion whether nature has provided for the instinctual repugnance between
species, “rather than by any more absolute decree” (10), but concluded
“by the result of numerous experiments” that “the absolute sterility of such
mixed offspring, must be held to be a law established in nature, and to it,
rather than to any supposed agency of instinct, must be attributed the uni-
versal preservation of distinct species” (12–13). All of this finds a clear
echo in Darwin’s Origin, where he writes “The view generally entertained
by naturalists is that species, when intercrossed, have been specially en-
dowed with the quality of sterility, in order to prevent the confusion of all
organic forms” (245).
As Mayr (1970) points out, sterility is only one of the reproductive iso-
lation mechanisms in the biological species concept. According to this con-
cept, a species is one or more actually or potentially interbreeding populations
separated from other species by reproductive isolation mechanisms (12).
These mechanisms, all of which “have a partially genetic base” (56), divide
into two classes (57), namely, premating (seasonal and habitat isolation, etho-
logical isolation, and mechanical isolation, an example of the latter being no
sperm transfer), and postmating (gametic mortality, zygotic mortality, hy-
brid inviability, and hybrid sterility). The biological species concept, as Mayr
further points out, is a “relational” concept, like “brother,” such that it is log-
ically impossible for the world to have only one species (13–14).
Mayr’s biological species concept emphasizes what keeps species apart.
There are a number of other species concepts, however, which emphasize
what keeps a species together. Perhaps the most important of these is Hugh
Paterson’s (1985) recognition species concept. Originally Paterson charac-
terized a species in terms of what he called a “specific-mate recognition sys-
tem,” defined as a signaling system between appropriate mating partners or
their cells. Each species for Paterson had its own distinct specific-mate
recognition system. However, due to the fact that many good species share
a common specific-mate recognition system (as with several species of
orchids from different genera), Paterson in his 1985 paper (which still
 Not Sterility, Fertility, or Niches 109

retained the name of his older species concept) expanded the original def-
inition to something more inclusive, namely, a distinct fertilization system
(which includes the specific-mate recognition system as a subclass). (For
further details on Mayr’s and Paterson’s species concepts, and for a detailed
critique, cf. Stamos 2003, 192–205.)
A number of other biologists have defined species in terms of ecologi-
cal niches, each species being kept together by a niche. I will mention a few
of these when we come to look at what Darwin had to say that is relevant
to the matter. What we shall see in this chapter is that not only did Dar-
win in his mature writings not come close to defining species in terms of
sterility, or fertility, or niches, but perhaps even more importantly he raised
serious difficulties for each of these approaches, difficulties that might be
judged to completely undermine them, even though his concept of repro-
ductive isolation mechanisms, fertilization systems, and ecological niches
was more limited than those of the relevant modern thinkers.
A good place to begin is with what Darwin thought about dogs. It is
clear that Darwin thought that the various wild species from which domes-
tic dogs were derived were intersterile to some degree. This comes out per-
fectly clear in two of his letters to Lyell written in 1859 (Burkhardt and
Smith 1991). In the first letter (October 25), Darwin summarizes the the-
ory of Pyotr Pallas, which includes the view that the wild species (which
Darwin expands in number) have a “tendency to sterility” that “when long
domesticated they lose” (357). In the second letter (October 31), Darwin
distinguishes between which parts of Pallas’ theory he accepts and which
parts he does not, and it is sufficient for our purposes that he writes “I must
think that the sterility which they would probably have evinced if crossed
before being domesticated, has been eliminated” (363).1 In his correspon-
dence, moreover, Darwin makes it clear that he thinks the wild parental
species (plural) of domestic dogs were themselves descended from a single
primitive species, in other words, that though domestic dogs as a group are
polyphyletic, their parental species as a group are monophyletic. This is
clear from his letter to his sister Caroline Wedgwood (November 21, 1859),
in which in reply to her query on the domestic dogs issue he wrote “It is a
distinct question whether these wild species have descended from one abo-
riginal stock as I believe has been the case” (386). Thus it would seem to
be Darwin’s view that intersterility was the constitutive factor that divided
the aboriginal stock into a number of wild species. In other words, from the
case of Darwin on dogs it certainly looks as if Darwin took reproductive iso-
lation (in the strict sense of sterility) to be a constitutive factor in the on-
tology of species (combined of course with other factors such as common
110 DARWIN AND THE NATURE OF SPECIES

descent). This conclusion appears only stronger if we take what Darwin

says about primroses and cowslips examined at the beginning of the previ-
ous chapter.
And indeed those few authors who have interpreted Darwin in his ma-
ture period as a species realist of some sort have advocated this conclusion
(regarding sexual taxa) in one way or another. Among the strategy theorists,
it is implied by Beatty (1985) in his interpretation of Darwin as believing
that what his fellow naturalists called “species” were really “chunks with[in]
the genealogical nexus of life” (278). In my first paper on Darwin (Stamos
1996), I explicitly held that Darwin’s mature species concept involved
sterility/fertility to a high degree, such that “It is the evidence that these
two criteria [‘interspecific sterility and intraspecific fertility’] are almost uni-
versal that allows Darwin to steer a course right down the middle between
species nominalism and species essentialism; in other words, on the one
hand it allows for species (for the most part) to be objectively real, while on
the other hand it allows for the indefinite mutability of species—both of
which are required for his theory” (140).
But it is the earlier theorists, namely, Ghiselin and Kottler, who pro-
vide the most interesting arguments and examples in this vein. What we
shall see in this chapter, however, is that if we take a closer and more crit-
ical look at their examples than they did, then we shall gain insights that in
turn will require us to reinterpret what Darwin says about dogs above and
primroses and cowslips in the previous chapter.
Beginning with Ghiselin (1969), he focuses on a paper of Darwin’s on
primroses and cowslips (Darwin 1869) surprisingly not included in Bar-
rett’s (1977) anthology. According to Ghiselin, in this paper Darwin

demonstrates that the intermediate form, or oxlip, is a sterile hybrid, and

supports this inference by showing that the oxlip occurs where the parent
species are present, but not otherwise. The third species is shown to be
sterile when crossed with the others, and to be distinct in morphology and
in geographical range. In summing up his observations, Darwin says that
the various forms “are all descended, from the same primordial form, yet,
from the facts which have been given, we may conclude that they are as
fixed in character as the very many other forms which are universally
ranked as species. Consequently they have as good a right to receive dis-
tinct specific names as have, for instance, the ass, quagga, and zebra. [100]

“When we realize that the analogy with horses,” continues Ghiselin, “is an
oblique reference to mules, we see that Darwin was stressing reproductive
distinctness” (100). Ghiselin nevertheless qualifies this interpretation on
 Not Sterility, Fertility, or Niches 111

the very next page: “Let it not be thought, however, that Darwin supported
the biological species definition in its strictly modern sense. There is no
solid evidence that he conceived of species as reproductively isolated pop-
ulations. His emphasis lay more with the distinctness of the individuals in
different species in terms of their biologically important characteristics, and
also with the genealogical interrelationships of the individuals within each
species” (101).
It must be somewhat odd, however, if Ghiselin’s interpretation is cor-
rect, that Darwin (as we shall see) would sometimes stress reproductive
distinctness and sometimes not, and with no apparent rhyme or reason.
Certainly a more charitable interpretation is desirable. And in fact it is
possible to reinterpret Ghiselin’s evidence as not stressing reproductive
distinctness at all, which would make Darwin more internally consistent.
This is the interpretation of Darwin’s mature species concept that I wish
to develop and defend here, namely, that for Darwin sterility, and its cor-
relative fertility, are not constitutive of species distinctness and species
unity respectively, but instead merely serve as probable evidence of such.
(I shall return later to what Ghiselin above calls Darwin’s “oblique refer-
ence to mules.”)
The evidence supporting this radical reinterpretation is pervasive.
There is, first of all, Darwin’s repeated efforts in the Origin toward the con-
clusion that sterility between species “is not a special endowment, but is
incidental on slowly acquired modifications, more especially in the repro-
ductive systems of the forms which are crossed” (272). As Darwin more
fully put it,

The importance of the fact that hybrids are very generally sterile, has, I
think, been much underrated by some late writers. On the theory of nat-
ural selection the case is especially important, inasmuch as the sterility of
hybrids could not possibly be of any advantage to them, and therefore
could not have been acquired by the continued preservation of successive
profitable degrees of sterility. I hope, however, to be able to show that
sterility is not a specially acquired or endowed quality, but is incidental
on other acquired differences. [245]

Given, then, that in Darwin’s view sterility between species cannot be

a direct consequence of natural selection, unlike other characters such as
beaks and eyes, so that it is therefore not an adaptation, and given what we
have seen in the previous chapter, it is legitimate to hypothesize that in
Darwin’s view any property that is not an adaptive consequence of natural
selection is not a species-specific property, it is not part of what a species is.2
112 DARWIN AND THE NATURE OF SPECIES

This conclusion seems to further recommend itself by the fact that

Darwin allows for intraspecific sterility, not just in the sense of sterile castes
but more importantly between fertile individuals of the same species. In
the Origin Darwin gives a number of examples of conspecific varieties with
“a certain amount of sterility” (269) between them. Perhaps the most in-
teresting case is the white and yellow varieties of nine species of the genus
Verbascum as studied “by so good an observer and so hostile a witness, as
Gärtner,” hostile because like so many others he considered “fertility and
sterility as safe criterions of specific distinction” (270). With these varieties
no one (including Darwin) doubted that they are varieties, since the yellow
and white varieties of each species “present no other difference besides the
mere colour of the flower; and one variety can sometimes be raised from the
seed of the other” (271). What is so remarkable about them is that within
each species there was found to be much less fertility when the white and
yellow varieties were crossed compared to when the crosses were confined
to within each of the two varieties (cf. Stauffer 1975, 406–407; Darwin
1876c, 258–259). Similar results are reported by Darwin from the experi-
ments by Gärtner on maize, Girou de Buzareingues on gourds, Kölreuter
on tobacco, and Darwin’s own observations on hollyhocks, which he sus-
pects “present analogous facts” (271).3
Of further interest are two papers of Darwin’s, one on the sexual rela-
tions within and between the dimorphic forms (short-styled and long-
styled) of primroses and cowslips (Darwin 1862b), the other on the sexual
relations within and between the trimorphic forms (short-styled, mid-
styled, and long-styled) of the purple loose-strife, Lythrum salicaria (Dar-
win 1865). In the former case, Darwin wrote in conclusion that “The
simple fact of two individuals of the same undoubted species, when homo-
morphically united [long-styled with long-styled, short-styled with short-
styled], being as sterile as are many distinct species when crossed, will
surprise those who look at sterility as a special endowment to keep created
species distinct” (Darwin 1962b, 61). In the latter case, the case of Lythrum
salicaria, “in each of the three forms the fertile unions are all heteromorphic,
the appropriate pollen coming from the stamens of corresponding length
borne by the other two forms, and . . . the homomorphic unions of the fe-
males with their own two sets of males are always more or less sterile” (Dar-
win 1865, 120). In the concluding paragraph of the paper he wrote that
“differences in sexual nature have been thought to be the very touchstone
of specific distinction. We now see that such sexual differences—the greater
or less power of fertilizing and being fertilized—may characterize and keep
separate the coexisting individuals of the same species, in the same manner
 Not Sterility, Fertility, or Niches 113

as they characterize and have kept separate those groups of individuals, pro-
duced from common parents during the lapse of ages or in different re-
gions, which we rank and denominate as distinct species” (128). And in
the concluding paragraph of a much later paper (Darwin 1880), he wrote
that “mutual sterility is no safe and immutable criterion of specific differ-
ence. We have, however, much better evidence on this head, in the fact of
two individuals of the same form of heterostyled plants, which belong to
the same species as certainly as do two individuals of any species, yielding
when crossed fewer seeds than the normal number, and the plants raised
from such seeds being, in the case of Lythrum salicaria, as sterile as are the
most sterile hybrids” (220).4
Moreover we have already seen in the case of domestic dogs and cat-
tle, and also in the theoretical case of convergence at the species level, that
Darwin did not take interbreeding or interfertility as a criterion of species
unity. And given this fact and the previous, it becomes difficult to main-
tain the view that Darwin’s mature species concept included either repro-
ductive isolation or interbreeding as constitutive of species.
If not constitutive, Darwin nevertheless held, as he put it in the Ori-
gin, both that the fertility between conspecific varieties is “almost invariably
the case” (268) and that sterility barriers between different species “can-
not, under our present state of knowledge, be considered as absolutely uni-
versal” (254). That neither are absolutely universal but almost universal is
quite sufficient for them to serve as good preliminary evidence of species
distinctness, even though, as we have seen, they are not constitutive prop-
erties of species.
The same conclusion may be inferred from Darwin’s discussion in the
Origin on “systematic affinity,” which he defines as “the resemblance be-
tween species in structure and in constitution, more especially in the struc-
ture of parts which are of high physiological importance and which differ
little in the allied species” (256). Of the covariance between the degree of
systematic affinity and the degree of hybridity, Darwin affirms that they do
largely covary, but he then immediately adds that “the correspondence be-
tween systematic affinity and the facility of crossing is by no means strict”
(257), indeed that “the capacity in any two species to cross is often com-
pletely independent of their systematic affinity” (258). Sometimes, as Dar-
win points out, there is a very high degree of systematic affinity between
two species but very little if any interfertility, while sometimes there is a
very high degree of interfertility but low systematic affinity. As such, then,
systematic affinity can do no more than merely serve as preliminary evi-
dence of interfertility. Thus, to apply once again the modern distinction
114 DARWIN AND THE NATURE OF SPECIES

employed in Chapter 4, intraspecific fertility and interspecific sterility are

for Darwin fallible diagnostic criteria of species distinctness, and they are
fallible because although usually found in and between distinct species they
are not constitutive characters of species.
Instead of fertility and sterility, then, the characters that do count for
Darwin as being constitutive of species are relatively constant and distinct
characters (morphological, ecological, and behavioral), characters moreover
of high physiological importance fixed as such by natural selection, all of
which we have seen in chapter 5. Sterility between species, on the other
hand, is not a character produced by natural selection, it is not adaptive,
and therefore should not count in a vera causa classification. Instead for
Darwin sterility should only serve as preliminary evidence.
Much the same goes for fertility, although this is far more problematic
and we have to be very careful here. In the Origin Darwin refers to hybrid
vigor, inbreeding depression, and the necessity of a certain amount of cross-
ing for hermaphrodites (267), and it might naturally be supposed that he
thought of this as adaptive, as something selected for. However, only a few
pages later, Darwin seems to assert the opposite and to view fertility in the
same category as sterility, not as something selected for but as a consequence
of selection:

I do not think that the very general fertility of varieties can be proved to
be of universal occurrence, or to form a fundamental distinction between
varieties and species. The general fertility of varieties does not seem to
me sufficient to overthrow the view which I have taken with respect to
the very general, but not invariable, sterility of first crosses and of hy-
brids, namely, that it is not a special endowment, but is incidental on
slowly acquired modifications, more especially in the reproductive sys-
tems of the forms which are crossed. [272]

At any rate, to understand Darwin on this matter we have to make a dis-

tinction between fertilization mechanisms, which as we have seen in the
previous chapter on the topic of orchids Darwin clearly thought were adap-
tations, and fertility per se, whether self-fertility in hermaphrodites or in-
traspecific fertility. Just because Darwin thought fertilization mechanisms
were adaptations, we cannot therefore assume that he also thought that fer-
tility per se was adaptive. On the contrary, there is plenty of evidence that
should direct us toward the opposite conclusion, that fertility per se was not
for Darwin adaptive, or often not adaptive. Some of that evidence comes
from what Darwin would later claim about self-sterility in plants. Darwin
was well aware that this was a matter of degree throughout the plant king-
 Not Sterility, Fertility, or Niches 115

dom, the opposite end being the so-called Darwin-Knight law (that no
plant self-fertilizes perpetually). In an interesting discussion, Darwin
(1876b) points out that it would be natural to suppose that self-sterility
had been gradually evolved by natural selection. However, he then goes on
to reject this view, his main reason being “the immediate and powerful ef-
fect of changed conditions in either causing or in removing self-sterility.
We are not therefore justified in admitting that the peculiar state of the re-
productive system has been gradually acquired through Natural Selection;
but we must look at it as an incidental result dependent on the conditions
to which the plants have been subjected, like the ordinary sterility caused
in the case of animals by confinement, and in the case of plants by too
much manure, heat, etc.” (346). By parity of reasoning, Darwin may well
have held the same view about much of intraspecific fertility, that it is an
incidental result dependent on the conditions of life. We have to recall that
in Darwin’s view natural selection produces adaptations only for the good
of the individual, a claim repeated throughout the Origin (e.g., 83–86), or
of the community only when that contributes to the good of each individ-
ual in the community (87), where by “good” Darwin clearly meant “wel-
fare” (127). Darwin, therefore, might well have held the view that, just as
interspecific sterility does the individual no good, so likewise intraspecific
fertility does or often does the individual no good, so that combined with
the evidence from the effect on reproductive systems by the conditions of
life, there is no good reason to suppose that fertility is always selected for.
What further supports this interpretation is Darwin’s claim that fertil-
ity can actually oppose natural selection. In the Origin he says “Intercross-
ing plays a very important part in nature in keeping the individuals of the
same species, or of the same variety, true and uniform in character” (103),
to which he adds “The process [of natural selection] will often be greatly
retarded by free intercrossing. Many will exclaim that these several causes
are amply sufficient wholly to stop the action of natural selection. I do not
believe so” (108).
We should also return to Darwin’s (1859) prime example of the white
and yellow varieties of the nine species of the genus Verbascum. It was fur-
ther found by Gärtner that “when yellow and white varieties of one species
are crossed with yellow and white varieties of a distinct species [of Verbas-
cum], more seed is produced by the crosses between the same coloured
flowers, than between those which are differently coloured” (270–271; cf.
Stauffer 1975, 406–407; Darwin 1876c, 258–259).
With this new interpretation, that for Darwin sterility and fertility are
generally good evidence of species distinctness but are not constitutive of
116 DARWIN AND THE NATURE OF SPECIES

species distinctness, we may look with new eyes at the evidence adduced in
favor of the interpretation that Darwin’s mature species concept involved
to some extent both “reproductive distinctness,” as Ghiselin put it above,
and “the genealogical interrelationships of the individuals within each
species.” With Ghiselin’s example of Darwin’s (1869) reference to the ass,
quagga, and zebra, it is difficult, especially following the analysis above, to
see how it is an oblique reference to mules, and therefore as stressing repro-
ductive distinctness. Whatever the degree of sterility or fertility between
these three species, Darwin need only have been referring to their relatively
constant and distinct characters (as indeed he does in Ghiselin’s quotation).
Interestingly, in one of his later papers Darwin (1880) remarked on the
perfect fertility between the common and Chinese goose, which, because
of their distinctness in many characters, elucidated by Darwin in his paper,
some authors had placed in different genera. But because of the discovered
perfect fertility between them, one author in particular had suggested that
the Chinese goose is only a domestic variety of the common goose. “But it
would, I believe,” replied Darwin, “be quite impossible to find so many
concurrent and constant points of difference as the above, between any two
domesticated varieties of the same species. If these two species are classed
as varieties, so might the horse and ass, or the hare and rabbit” (219–220,
italics in original). Far from being an oblique reference to reproductive dis-
tinctness (especially since in the goose example there is nothing compara-
ble to mules), the reference here is to constant character differences,
precisely in accordance with the analysis above.5
Perhaps more challenging than the evidence brought forward by Ghis-
elin is the evidence brought forward by Kottler (1978), according to whom
“passages from the Origin and later works, as well as Darwin’s taxonomic
practice, suggest that he did appreciate the importance of biological crite-
ria, including reproductive isolation” (292). Kottler’s main piece of evi-
dence is Darwin’s reference to sibling (cryptic) species (both terms are of
twentieth century vintage), species that are so morphologically similar that
their minor differences were either missed or ignored until it was discovered
that they were reproductively isolated from one another (hence they were
originally thought to be one species). In particular, Kottler focuses on Dar-
win’s repeated reference to willow wrens, referred to especially in his unfin-
ished Natural Selection and also in his review of H.W. Bates’ paper on
mimicry in butterflies.
My difficulty with Kottler’s use of these references, however, is that
there is no mention by Darwin of any kind of reproductive isolation, only
the maintenance of distinct characters. In the passage from Natural Selec-
 Not Sterility, Fertility, or Niches 117

tion that Kotter quotes (1978, 280 n. 9), Darwin wrote “In species we
should remember how extremely close some undoubtedly distinct forms
are, as many plants, & as in some of the willow wrens, which are so close
that the most experienced ornithologists can hardly distinguish them except
by their voice, & the materials with which they line their nests; yet as these
wrens inhabit the same country & always exhibit the same difference, no
one can doubt that they are good species” (Stauffer 1975, 99).
Of course one might infer that Darwin is implicitly referring to repro-
ductive isolation here. One area of support comes from his transmutation
notebooks, specifically Notebook B (Barrett et al. 1987, 224), wherein Dar-
win gave one of his definitions of species, which implicitly includes willow
wrens: “Definition of Species: one that remains at large with constant char-
acters, together with beings of very near structure.—Hence species may be
good ones & differ scarcely in any external character:—For instance two
wrens forced to haunt two islands one with one kind of herbage & one with
other, might change organization of stomach & hence remain distinct”
(213). Given that on the previous page Darwin wrote that “between Species
from moderately distant countries, there is no test but generation . . .
whether good species” (212), it is with good reason that Darwin scholars
such as Kottler (1978, 279–280), Sulloway (1979, 29), and Mayr (1982,
266) use this passage to interpret Darwin’s early species concept as includ-
ing sibling species.
In further support of Kottler’s interpretation of the willow wren refer-
ence in Natural Selection, it should be pointed out that the concept of sib-
ling species is also to be found in the Origin, where Darwin affirms that
“many species there are, which, though resembling each other most closely,
are utterly sterile when intercrossed” (268). Moreover Kottler’s interpreta-
tion may seem natural given the modern prevalence of the biological species
concept, with its inclusion of sibling species. But of course one must
always be careful not to allow the classic mistake of letting a highly influ-
ential modern scientific theory or definition influence one’s historical in-
terpretation of an earlier scientist. And this is possibly what we have here.6
Although to modern eyes it might look like Darwin, in his reference to
willow wrens in Natural Selection, is referring to sibling species and thereby
implicitly employing them in his mature species concept, there is, again, no
actual reference to reproductive isolation. And given the extended analysis
in the present chapter on Darwin’s rejection of sterility as constitutive of
specific difference, the only warranted conclusion is that the implication of
reproductive isolation is just not there. All that mattered to Darwin in terms
of his mature species concept, as follows from the present chapter and
118 DARWIN AND THE NATURE OF SPECIES

my analysis in the previous chapter, was that the two species have at least
one character difference, in particular, a different adaptation, and that the
difference has persisted in spite of the two species overlapping in range. In-
deed, in Natural Selection this comes out at the end of the chapter quoted
from above. Therein Darwin distinguishes between species and varieties,
the former in his view “being much more constant in all their characters,”
this constancy being due mainly to “the several causes of variability having
acted less energetically on the two species under comparison than on the
one species yielding the two or more varieties; and partly to the characters
of the two species having been long inherited, & by this very cause having
become more fixed” (Stauffer 1975, 165). Later in the same work, in his
chapter on natural selection, Darwin connects this constancy of characters
with natural selection:

In regard to the difference between varieties and species, I may add that
varieties differ from each other & their parents, chiefly in what natural-
ists call unimportant respects, as size, colour, proportions &c; but species
differ from each other in these same respects, only generally in a greater
degree, & in addition in what naturalists consider more important re-
spects. But we have seen in Ch. IV, that varieties do occasionally, though
rarely, very [sic, vary] slightly in such important respects; and in so far as
differences in important physiological characters generally stand in di-
rect relation to different habits of life, modifications however slight in
such characters would be very apt to be picked out by natural selection
& so augmented, thus to fit the modified descendants from the same par-
ent to fill as many & as widely different places in nature as possible.
[Stauffer 1975, 244]7

Indeed Darwin is not even committed to reproductive isolation keep-

ing two closely related, overlapping forms distinct. Although in Natural
Selection he admits that intercrossing “will obviously retard, perhaps oblit-
erate, the process of selection by dragging back the offspring of a selected
variety towards its parental type” (Stauffer 1975, 255), he nevertheless im-
mediately adds that this is a topic filled with “doubt.” As he put it in the
concluding section of his chapter on natural selection, “Intercrossing will
prevent or retard the process of natural selection; but here we are involved
in much doubt.” The doubt surrounds the weighing of several contingen-
cies responsible for “structural change” (271), or, as he a little later put it,
“the several contingencies favorable to natural selection.” “I am inclined,”
wrote Darwin, “to rank changed relations or associations between the in-
habitants of a country from opening up new places in its polity, as the most
 Not Sterility, Fertility, or Niches 119

important element of success.” Equally significant is that a few lines later

he wrote that “A diminished amount of intercrossing is probably the least
important element” (273).8
The second piece of evidence adduced by Kottler (1978) is from Dar-
win’s review of Henry Walter Bates’ (1862) paper on mimicry in butterflies.
In the second last paragraph of his otherwise favorable review, Darwin
(1863c) remarks that Bates “ought . . . to have given in every case his rea-
sons in full for believing that the closely allied and co-existing forms, with
which his varieties do not pair, are not distinct species. Naturalists should
always bear in mind such cases as those of our own willow wrens, two of
which are so closely alike that experienced ornithologists can with diffi-
culty distinguish them, excepting by the materials of which their nests are
built; yet these are certainly as distinct species as any in the world” (92).
Kottler’s footnote in which this passage is quoted and discussed (Kottler
1978, 280 n. 9) is a continuation of his discussion on Darwin’s “biologi-
cal” species concept in the transmutation notebooks and Darwin’s exam-
ple of willow wrens, which according to Kottler illustrates Darwin’s
inclusion of sibling species in that concept. Although Kottler in his foot-
note seems more concerned to point out that Darwin in the final page of
his review was prodding Bates to provide more evidence of “preferential
mating,” mating within though not between overlapping varieties of the
same species, it is obvious that Kottler’s use of the willow wrens reference
is meant to support his claim that even in Darwin’s mature period his
realist species concept included reproductive isolation to some degree.
When I read the above passage from Darwin’s review, however, I am
instead reminded of a particular passage from the Origin that we have seen
before, viz., “Those forms which possess in some considerable degree the
character of species, but which are so closely similar to some other forms . . .
that naturalists do not like to rank them as distinct species, are in several
respects the most important for us. We have every reason to believe that
many of these . . . closely-allied forms have permanently retained their char-
acters in their own country for a long time; for as long, as far as we know,
as have good and true species” (47). This passage, it seems to me, has those
like Bates written all over it. Although Bates adduced evidence of preferen-
tial mating in overlapping conspecific varieties, he nevertheless categorized
those varieties as varieties, not as species. This is especially clear in the case
of Mechanitis polymnia, which Bates says helped destroy his belief in the
constancy of species (530), and which Darwin accordingly refers to in the
third last paragraph in his review as a case that “well deserves study: after
describing its several varieties, Mr. Bates adds, ‘these facts seem to teach us
120 DARWIN AND THE NATURE OF SPECIES

that, in this and similar cases, a new species originates in a local variety . . .
from one variable and widely distributed species’” (92). Bates tells us that
M. polymnia is highly polymorphic (500 n*) and generally presents only
one local form in a district (531 n*). What is especially interesting about
M. polymnia, however, is not that some of its varieties, for example, the
predominant M. Egaënsis (Bates normally used binomials for varieties), ex-
hibit preferential mating (529), but rather its varieties (or rather the mem-
bers of those varieties) are uniquely adaptive. The unique adaptation(s) of
each variety, however, is not that of mimicry, which Bates, of course,
thought of as an adaptive product of natural selection (508). M. polymnia
is one of those species that is mimicked (502, 503, 564, 566). Nevertheless
Bates thinks of its varieties as differently adapted, by selection, “the selected
ones being different in different districts,” though “What these [the adap-
tive] conditions were, or have been, was not revealed by the facts” (511).
What Darwin is doing at the end of his review, then, it seems to me,
is challenging Bates’ species concept on a matter of theoretical consistency.
At one point Bates himself seems to equate a new species with a new adap-
tation, when he says the case of mimetic species “offers a most beautiful
proof of the truth of the theory of natural selection. It also shows that a
new adaptation, or the formation of a new species, is not effected by great
and sudden change, but by numerous small steps of natural variation and
selection” (513). But in practice, as in the case of M. Egaënsis, Bates still
considered it a variety of M. polymnia, rather than a species in its own
right, in spite of its having a unique adaptation. Part of his reason was the
existence of connecting forms (530), which unlike Darwin in cases such
as primroses and cowslips Bates repeatedly took as a criterion of conspe-
cific inclusion (501, 532, 546, 555). The other reason was that it did not
meet what Bates considered to be the test of species attainment: “The new
species cannot be proved to be established as such, until it be found in
company with a sister form which has had a similar origin, and maintain-
ing itself perfectly distinct from it” (530). What matters to Darwin, on
the other hand, is the permanence-of-characters criterion, characters more-
over selected and made relatively constant by natural selection, and this is
evident in the passage quoted above from Darwin’s review. It is evident in
his reference to willow wrens, and also in his own examples of preferen-
tial mating, such as “that two herds of differently coloured deer long pre-
served themselves distinct in the New Forest” (92).9 The preferential
mating, however, is not what Darwin’s species concept is about, nor his
challenge to Bates, since preferential mating would for Darwin be no more
 Not Sterility, Fertility, or Niches 121

adaptive by selection than would sterility. Instead his challenge to Bates is

to explain why his varieties, each with a unique adaptation, are not classi-
fied by Bates as distinct species. Of course other interpretations of Dar-
win’s challenge to Bates are possible, but the above interpretation is the
most consistent with everything else we have seen and is therefore the one
to be preferred.
Having rejected sterility and fertility as parts of Darwin’s species con-
cept, it remains to consider whether he would have accepted a niche com-
ponent. A number of modern biologists have defined species either in terms
of the occupants of a niche or as having a niche component. Ehrlich and
Raven (1969), for example, impressed by the weakness of gene flow within
many good species, by the amount of gene flow between many good
species, and by the sheer lack of it in asexual species, defined species in
terms of “similar selective regimes” (61). Van Valen (1976) is another ex-
ample motivated by much the same reasons. For him a species is a lineage
or set of lineages that occupies an “adaptive zone” minimally different from
any other lineage in its range (70). Others still, such as Simpson (1961), de-
fined a species not as occupying a niche throughout its entire existence (re-
call that Simpson conceived of species as vertical entities), so that the
demarcation of a species vertically would not be by a niche, but rather only
as having at any one time a unique adaptive “role,” which he equated with
a “niche” (154–155). With each of these biologists the niche is conceived
to be a property of the environment. But in some species concepts the niche
concept is more modern, conceiving of a niche as the actual or potential re-
source utilization of a population or species, so that the niche is not a prop-
erty of the environment (hence no empty niches) but rather a property of
the population or species. Thus Templeton (1989), for example, defines a
species as an evolutionary lineage “having the potential for phenotypic co-
hesion through intrinsic cohesion mechanisms” (12), and he divides these
cohesion mechanisms into basically two categories, the one being the ex-
change of genes via sexual reproduction, the other being the fundamental
(potential resource utilization) niche of the species. (For further details on
ecological niche species concepts as well as on ecological niche concepts, cf.
Stamos 2003, 144–156.)
Did Darwin have any of this in his species concept, and would he have
endorsed any of this? Although the actual use of the word “niche” as a con-
cept in ecology did not begin until the early 1900s, the concept was clearly
in circulation in Darwin’s time and even a little before it. Lyell (1832), for
example, used the phrase “the economy of Nature” (42), and after quoting
122 DARWIN AND THE NATURE OF SPECIES

from an 1820 essay by Augustin Pyramus de Candolle, in which the latter

distinguished “stations” from “habitations,” Lyell wrote

we may remind the geological reader that station indicates the peculiar
nature of the locality where each species is accustomed to grow, and has
reference to climate, soil, humidity, light, elevation above the sea level,
and other analogous circumstances; whereas by habitation is meant a gen-
eral indication of the country where a plant grows wild. Thus the station
of a plant may be a salt-marsh, in a temperate climate, a hill-side, the bed
of the sea, or a stagnant pool. Its habitation may be in Europe, North
America, or New Holland between the tropics. . . . The terms thus de-
fined, express each a distinct class of ideas, which have been often con-
founded together, and which are equally applicable in zoology. [69]

Not much later Darwin (1859) would write, as we have seen, of the “some-
what different stations” of primroses and cowslips (49), and more generally
of “each place in the economy of nature,” some of which are “unoccupied”
and which natural selection tries to “fill up” (102; cf. 104). In turn Bates
(1862) would write of “The two sets of forms [that] seem to agree . . . in
habits, and apparently occupy the same sphere in the economy of nature in
their respective countries” (498).
But niches played no role in the ontology of species for Darwin, and
his reasons were quite simple. As Darwin put it in the Origin,

We can clearly understand why a species when once lost should never
reappear, even if the very same conditions of life, organic and inorganic,
should recur. For though the offspring of one species might be adapted
(and no doubt this has occurred in innumerable instances) to fill the exact
place of another species in the economy of nature, and thus supplant it;
yet the two forms—the old and the new—would not be identically the
same; for both would almost certainly inherit different characters from
their distinct progenitors. [315]

We can see here that Darwin thought of species as forms, though of course as
plastic, rather than simply as the occupants of a niche, but also that he
thought of them in terms of adaptations, so that even if a very different lin-
eage would come to occupy a niche vacated for whatever reasons by a former
lineage, the two populations (horizontally conceived) would not have the
same suite of adaptations because of their different evolutionary histories.10
A further reason given by Darwin is that defining species in terms of
niches, rather than adaptations, makes a mess of biogeography and the
 Not Sterility, Fertility, or Niches 123

related issues of migration and evolutionary history, that is, phylogeny in

the most inclusive sense. In the Origin Darwin makes this point with regard
to the relationship between the endemic Galapagos species and the related
South American species. As he put it,

why should the species which are supposed to have been created in the
Galapagos Archipelago, and nowhere else, bear so plain a stamp of affin-
ity to those created in America? There is nothing in the conditions of
life, in the geological nature of the islands, in their height or climate, or
in the proportions in which the several classes are associated together,
which resembles closely the conditions of the South American coast: in
fact there is a considerable dissimilarity in all these respects. On the other
hand, there is a considerable degree of resemblance in the volcanic nature
of the soil, in climate, height, and size of the islands, between the Gala-
pagos and Cape Verde Archipelagos: but what an entire and absolute dif-
ference in their inhabitants! . . . I believe this grand fact can receive no sort
of explanation on the ordinary view of independent creation; whereas on
the view here maintained, it is obvious that the Galapagos Islands would
be likely to receive colonists, whether by occasional means of transport or
by formerly continuous land, from America; and the Cape de Verde Is-
lands from Africa; and that such colonists would be liable to modifica-
tion;—the principle of inheritance betraying their original birthplace.
[398–399]

In rejecting niches as part of what a species is (although he probably

didn’t realize he was doing that), Darwin was indeed avoiding a dead end,
for not only are there at least four main niche concepts in circulation in
modern biology, but whichever concept one chooses the concept of niche
is a notoriously problematic concept, such that some have despaired of the
concept having any objectivity at all. For example, in cases of pronounced
sexual dimorphism, such as male and female woodpeckers where the males
have larger beaks and feed in different parts of the trees than females, or in
cases of ontogenetic dimorphism, such as caterpillar/butterflies where dif-
ferent ontogenetic stages occupy very different niches, if in each case one
unites the niches into a single niche, it is obvious that one is presupposing
the species and then inferring the niche, whereas if one began with the
niches and defined species in terms of niches, then one would have to di-
vide the males and females of many species into different species, as well as
different ontogenetic stages. Moreover, niche talk is a highly subjective ex-
ercise. One can specify niches ad infinitum, depending on the imagination
of the theorist and the degree of vagueness or precision desired. But that
124 DARWIN AND THE NATURE OF SPECIES

does not make niches objective. Similar problems attend the distinction
between actual and potential niches. The potential niche of a population
or species is just as conjectural and vague and indeterminable as the con-
cept of environmental niche, whereas if one confines oneself only to the
actual niche, for example, in the sense of resource utilization, then what
are clearly different species might well utilize the same resources. In short,
resolving the species problem into the niche problem does not solve the
species problem, since the niche problem has too many empirical and con-
ceptual problems to ever be resolved, such that many ecologists themselves
have despaired of the concept altogether (cf. Stamos 2003, 153–165). Dar-
win was indeed prescient to focus on forms rather than on niches when it
came to the ontology of species.
In rejecting niches, we can figure out what Darwin would say to
modern species concepts that employ niches. Likewise, we can figure out
what Darwin would say to modern species concepts that employ steril-
ity or fertility.
In the case of sterility, we know from what Darwin wrote in reply to
Huxley. In many of his post-Origin writings on Darwin’s theory of evolu-
tion, Huxley (e.g., 1860b, 555) distinguished between what he called “mor-
phological species” (species based on morphology) and “physiological
species” (species based on interbreeding) and then went on to claim that,
in accordance with the inductive standards of science, selection has proven
itself “over and over again” capable of producing morphological species,
but as for physiological species “there is no positive evidence at present that
any group of animals has, by variation and selective breeding, given rise to
another group which was even in the least degree infertile with the first”
(567–568). To be fair, though, Huxley added his belief that experiments
in a few years will “very probably obtain the desired production of mutu-
ally more or less infertile breeds from a common stock” (568), as well as his
belief that “nature does make jumps now and then” (569), summing the sit-
uation up with the claim that Darwin’s theory “is as superior to any pre-
ceding or contemporary hypothesis . . . as was the hypothesis of Copernicus
to the speculations of Ptolemy” (569).
The repeated claim by Huxley that Darwin’s theory fell short of em-
pirical evidence on the production of species was a constant thorn in Dar-
win’s side, and for a brief while Darwin acquiesced to a certain degree. For
example, in a letter to Huxley (January 14, 1862) Darwin countered with
“you never allude to the excellent evidence of varieties of Verbascum &
Nicotiana being partially sterile together,” to which he added a few lines
later “Do oblige me by reading latter half of my Primula paper in Lin. Jour-
 Not Sterility, Fertility, or Niches 125

nal for it leads me to suspect that sterility will hereafter have to be largely
viewed as an acquired or selected character” (Burkhardt et al. 1997, 19).
Nevertheless, as we have seen earlier in this chapter (note 3), Darwin shortly
after gave up the idea that selection could produce sterility and permanently
reverted to his former view in the Origin. Consequently for Darwin, Hux-
ley’s physiological species were a red herring (in fact were not necessarily
species at all), and from 1863 onward Darwin’s claim against Huxley’s view
was not only that sterility is an incidental effect of selection rather than a
product of selection, but that sterility could not possibly be a species char-
acter since in some clearly good species, for example, in Verbascum and in
his later example of Lythrum salicaria (Darwin 1865), there is sterility
within the species. Consequently in early 1863 in a letter to Huxley
(January 10) we can see Darwin’s defiance of Huxley, where he indeed uses
varieties of Verbascum along with the kind of arguments we would later see
in Variation, promising that he will “not hold my tongue” (Burkhardt et al.
1999, 29). Almost four years would go by with apparently no debate be-
tween Darwin and Huxley on the topic, but when it came up again, Hux-
ley apparently stuck to his old line, to which Darwin in a letter to Huxley
(January 7, 1867) replied “Nature never made species mutually sterile by
selection, nor will men” (Kottler 1985, 407).
Darwin’s arguments against sterility as a species criterion prove to be
highly significant when we consider the modern biological species concept.
Critics of this concept have routinely argued that it does not work well in
plants, not only that it does not at all apply to asexual organisms but that
many conspecific sexual populations have little or no gene flow between
them so that it cannot be gene flow that gives them their cohesion (e.g.,
Ehrlich and Raven 1969; Levin 1979), that between many taxonomically
good species there is significant gene flow (e.g., Van Valen 1976), and that
asexual organisms generally divide into good species as well as if not better
than sexual species (e.g., Templeton 1989). Especially interesting is Hugh
Paterson’s (1985) argument that the isolating mechanisms of the biologi-
cal species concept, in particular the premating mechanisms, could not pos-
sibly have been evolved by natural selection (a view denied by Dobzhansky
but conceded by Mayr), so that they should be considered merely as effects
rather than as mechanisms proper (22). While Paterson’s argument is clos-
est to Darwin’s, what needs to be added to all these criticisms to arguably
clinch the case is Darwin’s focus on sterility inside a species, whether be-
tween what are unequivocally varieties of the same species, as with Verbas-
cum, or within what are unequivocally varieties of the same species, as with
each of the three forms of Lythrum salicaria. Since sterility is a major kind
126 DARWIN AND THE NATURE OF SPECIES

of reproductive isolation, reproductive isolation could not possibly be, for

Darwin, the defining characteristic of species.
Does this mean that fertility must go out the window too? Fertility is
not simply the other side of the coin, since it is not affected by all the prob-
lems presented by the sterility criterion, such as Darwin’s Lythrum exam-
ple, in which the three forms of Lythrum salicaria comprise together a fertile
whole. Interestingly, Paterson (1985) thinks that his recognition species
concept, which defines a species as having a unique fertilization system, is
closest to Darwin’s, since, unlike reproductive isolation mechanisms (so-
called), fertilization mechanisms are indeed adaptations produced by nat-
ural selection (25). Indeed Paterson goes on to claim that, in the first
sentence of his orchid book, Darwin (1862a) “expresses a view in perfect
agreement with the recognition concept: ‘The object of the following work
is to show that the contrivances by which the orchids are fertilized are as
varied and almost as perfect as any of the most beautiful adaptations in the
animal kingdom; and, secondly, to show that these contrivances have for
their main object the fertilization of the flowers with pollen brought by
insects from a distant plant’” (25).
Paterson’s species concept has gained a significant following (cf. Lam-
bert and Spencer 1995), but can he truly count Darwin as a precursor? I
have to say no, for the simple reason that although fertilization mechanisms
are indeed adaptations and as we have seen adaptations are the key for Dar-
win, fertilization mechanisms are not the only adaptations for Darwin (or
for anyone else for that matter), so that it is possible for him that two pop-
ulations could have the exact same fertilization mechanisms and yet be dis-
tinct species. And indeed we see good evidence for this in Darwin’s
writings. In the Origin Darwin discusses the example of two species of
Crinum experimented on by Herbert, in which he found, in Darwin’s
words, that “some hybrids are perfectly fertile” (250). Darwin also discusses
the examples of “certain species of Lobelia” and “all the species of the genus
Hippeastrum,” which, he says, “can be far more easily fertilised by the
pollen of another and distinct species, than by their own pollen” (250). He
also gives the example of Verbascum (270–271), as we have seen earlier in
this chapter. Moreover, in his long chapter on hybridism in Natural Selec-
tion he gives numerous examples among plants and animals and with vary-
ing degrees of fertility (Stauffer 1975, 388–462).
So neither sterility, fertility, nor niches defined species according to
Darwin. Having established that conclusion, and that the key for Darwin
is adaptations, we can finally provide a definition of Darwin’s implicit
species concept, which as we have seen shows a remarkable consistency
 Not Sterility, Fertility, or Niches 127

throughout his mature writings. Accordingly I would say that in Darwin’s

view a species is a primarily horizontal similarity complex of organism mor-
phologies and instincts distinguished at any one horizontal level primarily by rel-
atively constant and distinct characters of adaptive importance from the
viewpoint of natural selection, and secondarily (though not always applicable)
by common descent and intermediate gradations.
It is interesting to notice some of the implications that follow from this
species concept for speciation. Speciation is a concept, of course, which is
dependent on a concept of species. Without a concept of species, speciation
is a vacant concept. It follows, then, that what is speciation according to one
species concept might not be speciation according to another species con-
cept. It all depends on the particular features of each species concept. At the
end of chapter 3 we have seen some of the implications for speciation that
follow from taking species to be primarily horizontal entities. In the present
chapter we have seen that sterility and fertility are not for Darwin species
characters. This has an important implication for anagenesis, speciation in
a single evolutionary line. In phylogenetic systematics, in particular cladism,
the possibility of anagenesis is rejected as a matter of principle. Instead only
cladogenesis, branching speciation, is allowed. One reason for this is a dis-
taste for the concept of similarity (e.g., Ridley 1989). The rejection of sim-
ilarity, however, as we have seen in chapter 4, creates serious problems for
phylogenetic systematics, absurdities that we have further seen at the end
of chapter 3, such that some taxonomists who generally adhere to the prin-
ciples of cladism have found it necessary to let similarity back in, through
the back door as I like to put it, rejecting what has been called “Hennigian
extinction,” for example.
We can see in Darwin’s species concept, however, in particular on the
issue of extinction, that similarity is not rejected, nor is it sheepishly or re-
luctantly accepted simply to avoid absurdities. Instead it is robustly affirmed,
not in the facile and arbitrary sense of overall similarity, but in the sense of
being objectively bounded, both by the horizontal dimension and by com-
mon descent, but most importantly by adaptive characters produced by nat-
ural selection. Darwin, for that reason alone, would not reject anagenesis.
A further reason for why anagenesis is rejected in modern phylogenetic
systematics, it seems to me, is due to the vestige of the sterility criterion in the
work of the founder of cladism, Willi Hennig. Hennig (1966, 51–52, 58) rec-
ognized the reality of species at any horizontal level in accordance with the
biological species concept of Dobzhansky and Mayr, but he also recognized
problems with this concept, which he seemed to think would dissolve away
once the horizontal dimension is extended into the vertical dimension as the
128 DARWIN AND THE NATURE OF SPECIES

defining dimension of species (52–53; cf. Stamos 2003, 256–257). The ves-
tige here is the relational nature of species according to the biological species
concept, such that a species needs other species in order to exist, reproduc-
tive isolation being meaningless if there are no other species to be reproduc-
tively isolated from. Hence Hennig defined speciation in terms of a “cleavage”
(58, 64, 66), the first cleavage inaugurating the beginning of a species’ exis-
tence, the second inaugurating its end (Hennigian extinction), so that species
are delimited “by two successive processes of speciation” (63).
What follows from Darwin’s species concept, however, in particular
his rejection of sterility and fertility, is the sensible view that speciation is
coupled to evolution, in other words, that both anagenesis and cladogene-
sis are speciation processes. Why is this sensible? It is sensible because it
avoids a major prejudice, a prejudice that is common to modern phyloge-
netic systematics. The prejudice can be uncovered as follows. Viewed from
the horizontal dimension, most will agree that a species can evolve from
being monotypic (having no subspecies or varieties) to being polytypic
(having subspecies or varieties), all the while retaining, as a species, its nu-
merical identity. In evolving thus, most will also agree that it has undergone
part of the process of speciation, namely, cladogenesis. But if the same
monotypic species should evolve by the same amount (same in the sense of
any of its subspecies or varieties in the first scenario) while still remaining
monotypic, then why should this not also be considered part of the speci-
ation process, in this case, anagenesis? To allow that the change in the first
scenario is speciation but not the change in the second scenario seems to me
nothing more than plain and simple prejudice, a prejudice that has its roots,
I suspect, in the relational aspect of the biological species concept, and that
whatever its roots is contrary to the deepest insights of Darwin, in partic-
ular his language analogy for species. Of course once the above prejudice is
removed, as it is in Darwin’s species concept, speciation is necessarily cou-
pled to evolution and the horizontal dimension for species ontology takes
its rightful place of priority.
Or rather for Darwin speciation is coupled to adaptive evolution. And
here one might argue reside the main flaws in Darwin’s species concept. For
a start, it is questionable whether natural selection is a genuine law of na-
ture (Stamos 2006). But more importantly there is an epistemological prob-
lem with Darwin’s species concept, in that the determination of which
characters are adaptive and which are not is far from an easy task, such that
it is often subjective and subject to change (cf. Stevens 1980, 344–345). For
example, one has to wonder if, on the topic of whether humanity is one
species or more (cf. chapter 5), Darwin would change to a polygenist once
 Not Sterility, Fertility, or Niches 129

he learned (as modern biologists know) that human skin color is not a neu-
tral character but an adaptive character. Would humans then become like
primroses and cowslips? And how many today would accept that?
A further problem concerns the topic of speciation. By restricting species
to adaptations, as I have shown Darwin did, and moreover by restricting
adaptations to natural selection, Darwin was resistant to the concept of in-
stantaneous speciation. Darwin, of course, did not know about polyploidy
(or genetic engineering if we want to bring that in). The problem is that
polyploidy does indeed seem to make, in one generation and commonly in
plants, good and distinct species, moreover some that are adaptive and some
that are maladaptive. Either way they are good and distinct species, even if
the latter are quickly driven to extinction. A viable modern species concept
cannot afford to ignore any of this. It therefore should not take natural se-
lection to be the only cause of speciation. Darwin granted to other causal
processes a role in species modification, but as we have seen in this chapter
he allowed none of them an exclusive role in the formation of adaptations.
My suggestion is that Darwin’s species concept needs to be weaned from its
emphasis on adaptations and natural selection, it needs to be weaned from
one vera causa to many, so that the similarity complexes that are species are
bounded both by the horizontal dimension on the one hand and by a vari-
ety of causal mechanisms (natural selection included) on the other hand.
This I have attempted to do in the final chapter of my book on the modern
species problem (Stamos 2003, ch. 5).
The above suggestion returns us, however, to a further virtue of Dar-
win’s species concept. It concerns the issue of whether a universal species
concept is today a viable idea anymore. Many biologists have become skep-
tical, embracing the idea of species pluralism, that one species concept can-
not possibly satisfy all of modern biology, so that modern biology needs a
variety of species concepts (e.g., Mishler and Donoghue 1982; Endler 1989,
625; Ehrlich 2002, 347–348 n. 21). Other biologists, however, have ex-
pressed not only the hope but the need for a universal species concept.
Melissa Luckow (1995), for example, points out that “a universal and de-
finitive species concept is most important to those systematists who actu-
ally circumscribe and name species” (589), to which she adds “the species
problem will be solved by the continued collection and analysis of data,
the clarification of issues and terms, and the application of new ideas”
(600). Joel Cracraft (1987) adds that until this happens the Modern Syn-
thesis is incomplete, since “the ‘synthetic theory of evolution’ has incorpo-
rated very disparate concepts of species, so much so that it undermines the
very notion of a ‘synthesis’” (334).
130 DARWIN AND THE NATURE OF SPECIES

How does Darwin help? First, with the help of his language analogy for
species, we have seen in chapter 3 that for Darwin the primary reality of
species is horizontal. Darwin could not have known this, but one conse-
quence of this move is that it does some serious damage to species plural-
ism, since contrary to species pluralism the priority of the horizontal
dimension for species reality privileges some species concepts over others.
The result is not that we are left with one species concept, but the door is
certainly opened to that possibility (cf. Stamos 2002, 192, 2003, 96–97).
The second way Darwin helps is by emphasizing pattern over process.
This is a further distinction by which modern species concepts can be di-
chotomously classified. Process species concepts view the causal processes
responsible for the existence of particular species as part of the ontology of
species, whereas pattern species concepts view them apart from the ontol-
ogy of species. The biological species concept, for example, is a process
species concept, while the morphological species concept is a pattern species
concept. Darwin’s species concept is also a pattern species concept, since the
process of natural selection is not for him part of what a species is. The
virtues of pattern species concepts over process species concepts are many.
One is that epistemologically they are far more accessible, since they focus
only on the product, not on the causal processes responsible for the prod-
uct (Cracraft 1989, 34), and the causal processes responsible for the prod-
uct are largely untestable (Luckow 1995, 591–592). Second, pattern
supervenes on process, in that similar products can be produced by dissim-
ilar causal processes (Levin 1979, 384). Third, pattern species concepts
tend to be neutral as to the causal processes responsible for species and
therefore do not stand or fall with those theories, which is a virtue since bi-
ologists already know that the causal processes are many and may still be
more complicated than they already think, and also because the causal
processes may well be different in different species (Cracraft 1983, 102,
1989, 34). Fourth and finally, it is only in focusing on pattern rather than
on process, along with the priority of the horizontal dimension, that hope
for a universal species concept resides. As Luckow (1995) put it, “if species
are viewed as the endpoints or results of various processes rather than as
participants, a universal species concept is still viable” (590).
Darwin’s species concept, of course, does not quite instantiate the
above virtues, since he did not cleanly separate the product, adaptations,
from the process, natural selection (as we have seen in Chapter 1 on the
topic of continuous natural selection). Nevertheless, in developing and em-
ploying what was clearly a pattern species concept, Darwin made that first
important step and once again helped show us the way.
 Chapter 7

The Varieties Problem

Given, as we have seen, that Darwin thought species are real, also that he
thought varieties are incipient species, the species problem then becomes,
when studying Darwin, the varieties problem. If species are real and vari-
eties are incipient species, then varieties would have to in some sense be
real too. In this chapter we shall compare Darwin’s view on varieties with
those of his contemporaries. The latter topic especially is important as a
preparation for the topic of the next chapter, the question of why Darwin
would repeatedly define species (both taxa and category) as not real and
yet treat them as real, and not only as real but moreover repeatedly and
consistently employ his own implicit species concept at that. In other
words, to understand Darwin’s strategy, we have to understand both
Darwin and his contemporaries on the nature of varieties.
A good place to begin is with two passages examined in chapter 1, in
which Darwin compares different species concepts and comments on the
issue. In a letter to Hooker (December 24, 1856) Darwin wrote

I have just been comparing definitions of species . . . . It is really laugh-

able to see what different ideas are prominent in various naturalists minds,
when they speak of “species” in some resemblance is everything & descent
of little weight—in some resemblance seems to go for nothing &
Creation the reigning idea—in some descent the key—in some sterility
an unfailing test, with others not worth a farthing. It all comes, I believe,
from trying to define the undefinable. [Burkhardt and Smith 1990, 309]

131
132 DARWIN AND THE NATURE OF SPECIES

The analysis presented in c hapters 2–6 should allow us to say what

Darwin really thought about each of these criteria. Although resemblance
was not everything, it was not nothing either, because of common descent.
Moreover descent for Darwin was certainly not of little weight but was
instead of great weight, because only through descent for Darwin were
adaptations acquired, and yet common descent was not the key, as we have
seen in the case of primroses and cowslips. As for sterility, it was not for
Darwin an unfailing test, nor was it not worth a farthing, but instead it
was a fairly reliable test of species distinctness, although it was no part of
what a species is. Finally, from Darwin’s practice we can see that he did in
fact have an implicit definition of “species.”
The passage quoted above, therefore, is highly disingenuous. What
makes this even more apparent is that Darwin does not mention Edward
Blyth’s criterion, which we have seen in Chapter 4 on the topic of domes-
tic cattle, and which we have seen in the previous chapter has proven to be
of great importance to Darwin, namely, constant and distinct characters. In
the next chapter we shall see how the above disingenuous passage fits with
what I shall argue was Darwin’s strategy. For the present, it is useful to
compare it with a highly similar passage from Natural Selection, found near
the beginning of Chapter IV, the first draft of which Stauffer (1975, 92)
tells us was written, according to Darwin’s Pocket Diary, from mid-
December 1856 to late January 1857:

. . . how various are the ideas, that enter into the minds of naturalists
when speaking of species. With some, resemblance is the reigning idea &
descent goes for little; with others descent is the infallible criterion; with
others resemblance goes for almost nothing, & Creation is everything;
with others sterility in crossed forms is an unfailing test, whilst with oth-
ers it is regarded of no value. At the end of this chapter, it will be seen that
according to the views, which we have to discuss in this volume, it is no
wonder that there should be difficulty in defining the difference between
a species & a variety;—there being no essential, only an arbitrary differ-
ence. [Stauffer 1975, 98]

The similarity of this passage to the other is remarkable, and indeed it

is possible that they were written on the very same day. What is especially
significant for our present purpose, however, is the difference in the end-
ings. For unlike the previous passage, Darwin in the above passage relates
the issue to the difference between species and varieties, stating that there
is no essential difference between them. The same claim would resurface in
the Origin, where Darwin says that in his view “the term species . . . does
 The Varieties Problem 133

not essentially differ from the term variety” (52), and again that “there is
no essential distinction between species and varieties” (272). The same
claim also finds echoes in his correspondence, for example, in his letter to
Lyell (September 28, 1860) in which he wrote “Is not your feeling a rem-
nant of that deeply-impressed one on all our minds, that a species is an en-
tity,—something quite distinct from variety?” (Burkhardt et al. 1993, 397).
All of the above raises the conceptual issue of what Darwin meant when
he said there is no essential distinction between a species and a variety.
According to Ghiselin (1969), Darwin “insisted on Aristotelian definition
as a criterion of reality or naturalness” (82), so that Darwin “maintained
that there are no ‘essential’ differences between species and varieties, and
that both terms designate the same basic kind of entity.” Thus, “Darwin
was denying the reality, not of taxa, but of categories” (93). Similarly,
according to Beatty (1985), “the crucial issue was that there was no way of
defining ‘species’ that distinguished species from varieties—no way of
defining the difference” (275). More specifically, “what are called ‘varieties’
of species are, in time, transmuted into what would be called ‘species’ in their
own right” (276). Thus, “Darwin’s theory of the evolution of species was,
of course, about the evolution of species taxa rather than the evolution of
the species category. So his denial of the species category did not render
his theory domainless” (278).
As we have seen in chapter 2, Ghiselin’s claim that Darwin insisted on
Aristotelian essentialism for naturalness simply does not hold up to close
scrutiny, and Beatty’s claim that Darwin simply took what his fellow nat-
uralists called “variety” and “species,” as we have seen in chapter 5, fares no
better. We shall therefore have to take a completely different approach to
the topic of Darwin on varieties.
To cut to the chase, it seems to me that Darwin, in claiming that there
is no essential difference between a species and a variety, was not claiming
that neither is real, nor was he claiming that they are the same basic kind
of entity. We shall see later in this chapter, based on the reconstruction of
Darwin’s species concept developed in previous chapters, that Darwin
thought that varieties too are real, only that their reality is partially differ-
ent than that of species. But equally important, in claiming that there is no
essential distinction between them, Darwin was merely claiming that there
is no bridgeless gap between them. That is what essentialism meant to
Darwin (and to many today). That is what Darwin meant (at least in part)
when he wrote to Lyell, quoted above, that a species is not a distinct entity.
Again, if species for Darwin are real (and we have seen that they are), and
if varieties are incipient species (unquestionably Darwin’s view), then it
134 DARWIN AND THE NATURE OF SPECIES

would have to follow that varieties too are real, but it would not have to
follow that they are real in the exact same sense as species. And indeed this is
what we shall find in this chapter. What we have found in chapter 5 is that
for Darwin the key feature of species is unique adaptations created and
fixed by natural selection. What we shall find in the present chapter, how-
ever, is not that there is a distinct difference between species and varieties,
but instead that there is a gradual difference, so that at one point we can say
that this is clearly a variety and at another point that this is clearly a species,
but that also somewhere in between we cannot say either way. What we
shall see, then, is that not only, once again, did Darwin not insist on Aris-
totelian essentialism as his criterion of reality, but also that he implicitly
denied Aristotle’s guiding principle (essential to his essentialism) that what-
ever is different in degree is not different in kind (e.g., Parts of Animals
644a16–18, Politics 1259b36–38).
What adds a wrinkle to this view, however, is that in the Origin
Darwin tells us that “every one admits that there are at least individual dif-
ferences in species under nature. But, besides such differences, all natural-
ists have admitted the existence of varieties, which they think sufficiently
distinct to be worthy of record in systematic works” (468–469). The claim
is also found earlier in Natural Selection, but with some interesting twists:
“this admission [subspecies by authors ‘of the highest authority’—Darwin
cites six] is important as sub-species fill up the gap, between species, admit-
ted by everyone & varieties admitted by everyone. Between varieties & in-
dividual differences there seems a gradual passage but to this subject we shall
recur” (Stauffer 1975, 99). Darwin’s motive behind these assertions is plain
enough, and it becomes even more obvious at the bottom of the paragraph
from which the above passage is taken: “So that between individual differ-
ences & undoubted species naturalists have made various short steps” (99).
What is plainly clear is that Darwin wanted to believe, or more pre-
cisely wanted his readers to believe, that his fellow naturalists believed
in the reality not only of variations but also of varieties and subspecies
and in fact every grade between individual variation and species, the rea-
son being one more piece of evidence in his one long argument for evo-
lution by natural selection, in particular that varieties are “incipient
species.” As Darwin put it in the Origin, “These differences blend into
each other in an insensible series; and a series impresses the mind with the
idea of an actual passage” (51). Moreover it explains why Darwin would
repeatedly focus, in both Natural Selection (Stauffer 1975, e.g., 112–113,
159–164) and the Origin (e.g., 48, 404), on the many doubtful forms
called “species” by some and “varieties” by others, a point to which he
 The Varieties Problem 135

owed much to Watson (cf. letter from Watson, November 19, 1856,
Burkhardt and Smith 1990, 276–277).
But did Darwin’s fellow naturalists actually believe in the existence of
varieties as Darwin would have us believe? This is a question that remains
to be answered. In what follows I shall argue that the answer in the main
is no, and moreover that Darwin knew it. Darwin’s knowledge that he was
misleading will prove to be especially significant when we turn to his strat-
egy examined in the next chapter.
The first point of evidence is that although naturalists gave plenty of
definitions of the concept of species (the commonality was creationism, as
we have seen in c hapter 1), they said relatively little with regard to the na-
ture of varieties. Consequently, unlike species, we have to really work at
figuring out what their concept of variety was.
Perhaps the best place to begin is with those who explicitly denied that
varieties exist. Louis Agassiz is a good case in point. In an often-cited pas-
sage, Agassiz (1860a) claimed that “varieties properly so called, have no ex-
istence, at least in the animal kingdom” (410). It might be replied that this
was simply a rearguard move on Agassiz’s part in response to the Origin.1
Mayr (1976, 258, 259), however, argues that this was not the case, that
this was a view that Agassiz held well before and in spite of the Origin.
Mayr points out that of course Agassiz recognized the existence of individ-
ual variations—who didn’t?—but he did not recognize the existence of
varieties as subspecific taxa. Agassiz avoided ascribing reality to varieties by
applying two different approaches. One was to be a splitter and to make a
species out of every definable distinction. Mayr points out that Agassiz did
this in the case of his study of the fish of the Tennessee River, published in
1854. As Mayr puts it, “His disbelief in the existence of ‘varieties’ forced
him to describe several ‘species’ from schools of single species” (258).
Agassiz’s other approach was to unite into one species forms that insen-
sibly intergrade, as he did in the case of echinoderms (258–259). In all of
this we can see his rejection of varieties as being consistent with his
Platonic/Christian approach to species. For Agassiz, as he stated in his Essay
on Classification (1857), “the individuals living at the time have alone a mate-
rial existence,” while “branch, class, order, family, genus, and species . . . .
have been instituted by the Divine Intelligence as the categories of his mode
of thinking” (8). Again: “individuals . . . do not constitute the species, they
represent it. The species is an ideal entity, as much as the genus, the
family, the order, the class, or the type” (175–176). Moreover no one
individual “exhibits at one time all the characteristics of the species” (176).
Each species, then, for Agassiz, is in reality only an idea in the mind of
136 DARWIN AND THE NATURE OF SPECIES

God, not an entity in nature. This comes out in various places in his Essay;
it also comes out clearly and succinctly in his review of the Origin (1860b).
There he says “while individuals alone have a material existence, species,
genera, families, orders, classes, and branches of the animal kingdom exist
only as categories of thought in the Supreme Intelligence, but as such have
as truly an independent existence and are as unvarying as thought itself
after it has once been expressed” (151). To this he adds that each species has
“essential” or “specific characters” which “are forever fixed,” such that “in-
dividual peculiarities” are “in no way connected with the essential features
of the species, and are therefore as transient as the individuals” (150).2
The case of Agassiz raises an issue that Darwin was certainly aware of,
namely, that most naturalists from the time of Linneaus to his own day
were splitters, not lumpers. The evidence for this is overwhelming. Caro-
lus Linnaeus, for example, the self-proclaimed Prince of Botanists, com-
plained of the varieties “which the common herd of botanists calls species,”
and for him such splitting is wrong because the variation is merely “in the
outside shell” (Ramsbottom 1938, 198 n.). Jean Baptiste de Lamarck, in
his Flore Francoise, published in 1779 and before he became an evolution-
ist, complained that “nearly all present-day botanists” are splitters, “mul-
tiplying the species, at the expense of their varieties, to infinity” (Burkhardt
1987, 163). H.C. Watson (1845b) in England wrote of the “species-
splitting monomania” (219), while Hooker and Thomson in their Flora
Indica, published in 1855, complained that “the study of systematic
botany is gradually taking a lower and lower place in our schools; it falls
into the hands of a class of naturalists, whose ideas seldom rise above
species, and who, by what has well been called hair-splitting, tend to bring
the study into disrepute” (Stevens 1997, 354). Darwin himself notes in his
Origin that “differences, however slight, between any two forms, if not
blended by intermediate gradations, are looked at by most naturalists as
sufficient to raise both forms to the rank of species” (485; cf. 424–425).
Indeed in a letter to Leonard Jenyns (April 28, 1858) Darwin complained,
in line with the complaints above, that “One chief reason why I have not
accumulated more facts of variation in a state of nature is, that naturalists
so invariably turn round & say oh they are not varieties, but species”
(Burkhardt and Smith 1991, 86). Two years later in a letter to Lyell (June
14, 1860) he would make basically the same complaint, this time specifi-
cally about Agassiz: “It is no wonder Agassiz denies varieties in animals,
when he calls even the same forms in two distant countries, two species”
(Burkhardt et al. 1993, 254). And yet “all naturalists have admitted the
existence of varieties”!
 The Varieties Problem 137

The fact is, or at least I shall argue it, most naturalists did not admit the
existence of varieties, in spite of their linguistic habits. A few were explicit
about it, but most left it implicit. Certainly it seemed to follow from their
essentialism. A good case in point is Linnaeus. Confining ourselves to his
earlier works, for which he was most famous, Linnaeus states, in for in-
stance his Fundamenta Botanica, published in 1736, that “There are as
many varieties as there are different plants produced from the seed of one
and the same species” (Leikola 1987, 49)—“Varietates tot sunt, quot dif-
ferentes plantae ex ejusdem speciei semine sunt productae” (Ramsbottom
1938, 199). This might be a simple case of what Blyth (1835) wrote about
in his attempt to classify the different ways in which naturalists used the
term “variety.” According to Blyth, “The term ‘variety’ is understood to
signify a departure from the acknowledged type of a species, either in struc-
ture, in size, or in colour; but is vague in the degree of being alike used to
denote the slightest individual variation, and the most dissimilar breeds
which have originated from one common stock” (40–41). On the other
hand, Linnaeus’ definition above might be interpreted as meaning to say
that there are as many varieties of a species as there are individuals of that
species with variations. In other words, it is a cynical treatment of varieties.
Variations are character states that individual organisms have, whereas
varieties are subspecific taxa made up of one or more individual organisms.
Linnaeus, then, it seems to me, is saying that varieties are arbitrary classifi-
cations. And indeed for Linnaeus, raised in one of the last strongholds of
Aristotelian scholasticism in Europe, the University of Uppsala in Sweden
(Leikola 1987, 45–47), species are essentialistic kinds, created from the
beginning and remaining essentially unchanged (49, 56 n. 10), defined
binomially by a genus and differentia (which was a shorthand for a full
description of essential characters), genus level characters being generative
organs, species level characters being nutritive parts such as roots, stems,
and leaves (49, 56 n. 10). Variety characters, on the other hand, were based
on characters that resulted from accidental causes such as soil and climate—
“Varietas est Planta mutata a caussa accidentali: Climate, Solo, Calore, Ven-
tis, &c., reducitur itaque in Solo mutato” (Ramsbottom 1938, 199)—and
also culture—“Naturae opus semper est Species & Genus; culturae saepius
Varietas” (199–200)—characters such as, in the case of plants, size, color,
smell, and taste—“Species varietatum sunt Magnitudo, Plenitudo, Crispa-
tio, Color, Sapor, Odor” (Ramsbottom 1938, 199). But equally important,
Linnaeus confined varieties, or more specifically the variations on which
they were based, not to the deep essence of the species, something that was
passed on in the “unity of generation” (Leikola 1987, 50), as he called it in
138 DARWIN AND THE NATURE OF SPECIES

Systema Naturae, published in 1735, but rather, as we have seen above, to

what he called in Critica Botanica, published in 1737, “the outside shell”
(Ramsbottom 1938, 198 n.).
For all of the above reasons, and more, I would agree with Larson’s
(1968) view that for Linnaeus varieties were not real (291). As we have seen
in chapter 2, for Linnaeus categories were conceived to be like boxes (Leikola
1987, 46). But Linnaean varieties do not at all fit this scheme. They have no
underlying essence, there are no bridgeless gaps between them (recall from
chapter 5 that Linnaeus was a lumper), they are based on characters that not
only are not fixed but are infinite in their possibilities (this comes out from
his cynical definition of “variety” quoted above), and finally, Linnaeus
believed in the so-called law of reversion, the belief that species can vary
within certain “fixed limits” and that when the conditions that caused them
to vary are removed, “true species,” as he put it in Critica Botanica, will
“finally revert to the original forms” (Ramsbottom 1938, 200 n.). In all of this
it is very difficult to see Linnaeus as falling under Darwin’s claim, quoted
above, that “all naturalists have admitted the existence of varieties.”
We can see the same in Linnaeus’ contemporary and nemesis George
Louis Buffon, highly influential in France and the first big name among
French naturalists. Although Buffon expressed species nominalism at the
beginning of his Histoire Naturelle, published in forty-four volumes be-
tween 1749 and 1804, he quickly took up species essentialism, arguing that
each species has what he called an internal mold (moule intérieur), which
causally preserved the form of the species from generation to generation. Of
this mold he stated that “The type of each species is cast in a mold of which
the principle features are ineffaceable and forever permanent, while all the
accessory touches vary” (Stamos 1998, 449). Granted, Buffon also devel-
oped a theory of degeneration, extending it more and more as his writings
progressed. According to this theory many species have degenerated over
the ages from a common stem (souche première), resulting in what are
wrongly thought to be species, such as the ass and zebra, which for Buffon
are merely varieties of the horse (447–448). Hence Buffon’s species are
much more inclusive, more like Linnaeus’ genera. But even so, we can see
in this what is arguably only a much more extreme view of the plasticity of
species that we find in Linnaeus and other essentialists before Darwin. For
Buffon these varieties do not have their own internal mold, each with its
essential characters, but are merely incremental perturbations, “accessory
touches” that “vary,” often cumulative and heritable but always reversible,
produced by differences in environmental conditions and domestication
(Farber 1972, 268–278).
 The Varieties Problem 139

The idea that variations in a species are perturbations from the norm
or essence of the species had quite a wide currency. In addition to Linnaeus
and Buffon, another enormously influential example is Antoine-Laurent
de Jussieu, whose natural system (which involved taking into account the
entire organization of each individual, not just characters thought to be
privileged for classification) served as the basis for plant systematics for well
over a century, and not just in France but throughout continental Europe
and England (Stevens 1994, xx). In the Introduction to his Genera Plan-
tarum, published in 1789, Jussieu defined species as
a collection of beings that are alike in the highest degree, never to be
divided, but simple by unanimous consent and simple by the first and
clearest law of Nature, which decrees that in one species are to be assembled
all vegetative beings or individuals that are alike in the highest degree in all
their parts, and that are always similar over a continued series of generations,
so that any individual whatever is the true image of the whole species,
past, present, and future. [Stevens 1994, 356–357]

This is a clear example of an essentialist species concept. Indeed a little ear-

lier in the same work Jussieu states that species are “immutable and perpet-
ual in kind” (340). And yet he allowed for varieties, but in a way that, like
Linnaeus, hardly made them real. This is because he allowed species to be
“occasionally subverted for a while by chance or human industry” (340–341),
to vary in characters such as color or size of organs. Moreover he states that
“these varieties, obeying the law of nature, and committed to a new germi-
nation of seed; they return to the primordial species, their character restored,
if other factors do not interfere” (341). Not only do we find in this the law
of reversion, but again, just like Linnaeus, we find the view that variations
are perturbations from the norm, as well as the logical consequence
(though it is implicit) that naming varieties (subtaxa based on these
variations) is ultimately arbitrary.
I will return to the law of reversion shortly. For the present I want to
continue with the idea of variations as perturbations. Among influential
sources, especially in England, James Prichard (1813) is a good example.
Prichard subscribed to the common view (as we have seen in chapter 1) that
all classificatory categories are matters of “arbitrary definition,” the excep-
tion being the species category, where “the distinction is formed by nature”
(7). Nevertheless he also states that “it is well known that considerable va-
rieties arise within the limits of one species, and such varieties often be-
come to a great degree hereditary in the race, and permanent” (8).3 To
determine whether a particular group is part of a real species “distinct from
140 DARWIN AND THE NATURE OF SPECIES

their first creation” (8) or part of a variety (which developed from a real
species), Prichard referred to what he called “indirect methods of reasoning”
(8), which for him had to be the sterility test (12–13). What is interesting
is that he then goes on to discuss “the kinds of variation in which Nature
chiefly delights” (13), and it is clear that like so many others he confused
varieties with variations. This comes out clearly when he states
When we have found that any particular deviation from the primitive
character has taken place in a number of examples, the tendency to such
variety may be laid down as a law more or less general, and accordingly
when parallel diversities are observed in instances, which do not afford us
a view of the origin and progress of the change, we may nevertheless
venture to refer the latter with a sufficient degree of probability to the
class of natural varieties, or to consider them as examples of diversified
appearance in the same individual species. [13–14]

Prichard then goes on to give the example of albinoism in mice, rats, and
crows (14), and further states that the most common kinds of change are
“varieties of form and colour” (15). We then see what one finds in the
literature of this period time and again, namely, the idea of variations,
including heritable variations, as perturbations from the norm:

We may however in general observe, that when the condition of each

species is uniform and does not differ materially from the natural and
original state, the appearances are more constant, and the phænomena of
variation, if they in any degree display themselves, are more rare and less
conspicuous, than when the race has either been brought by human art
into a state of cultivation, or domestication, or has been thrown casually
into circumstances very different from the simple and primary condition.
[15–16]

What only underscores Prichard’s confusion of variations with varieties is

when he applies his principles to the human species (Prichard was a mono-
genist), naming as varieties the albino (17–19), red or yellow hair with fair
skin (20), black hair and dark eyes with white but tanable skin (21–22),
yellow to olive skin with long black hair (“the Mongoles,” etc.) (22–23),
“Native Americans” (23), “pie-bald Negroes” (blacks with patches of black
and white skin and hair) (23–24), and finally black or dark yellowish-brown
skin (24). What is interesting about Prichard is that he does not seem to
have subscribed to the law of reversion, which as we shall shortly see was
generally accepted as a natural law in England and elsewhere.
Like Prichard, Edward Blyth (1835) also allowed for permanent vari-
 The Varieties Problem 141

eties, but he was more circumspect on how his fellow naturalists used the
term “variety” and also on the law of reversion. Dividing those uses into four
general categories, the first, as we have seen, being the confusion with indi-
vidual variation, Blyth’s second category is acquired variations either in a
single individual or in a series of individuals in which “various changes . . .
are gradually brought about by the operation of known causes” (43), while
his third category is breeds, typically produced by human agency but he
allows that they could be produced in nature as a result of “accidental iso-
lation” (45). The interesting thing about breeds, according to Blyth, is
that “if man did not keep up these breeds by regulating the sexual inter-
course, they would all naturally soon revert to the original type” (46).
Blyth’s fourth and final category, interestingly the only one to which he
thinks the term “variety” properly applies, and he knows this restriction is
“peculiar” (47), is what he calls “true varieties,” and they are what Prichard
and others would call “permanent” varieties. These true varieties, he says,
“are, in fact, a kind of deformities, or monstrous births, the peculiarities
of which, from reasons already mentioned, would very rarely, if ever, be
perpetuated in a state of nature; but which, by man’s agency, often be-
come the origin of a new race” (47). Some of the examples he gives are
ancon sheep, with their legs too short to jump over fences, tailless cats,
many kinds of domestic dogs, and fan-tailed pigeons. Of true varieties,
he says, “The deviations of this kind do not appear to have any tendency
to revert to their original form” (47).
What follows from Blyth’s analysis is that what are called “varieties” in
nature are probably not such at all, that varieties in nature probably do not
exist. Interestingly, for further reading Blyth strongly recommends Prichard
(1813), to which he adds “some sound and excellent remarks on varieties will
also be found in the second volume of Lyell’s Principles of Geology” (48).
When we turn to Lyell (1832), however, we find ideas that do not seem
congenial to Blyth’s analysis. In chapter 1 we have already seen that Lyell,
like so many others, held the premise that if species are not fixed but evolve,
then species are not real. It would be odd, then, to the point of inconsistency,
if Lyell at the same time thought that varieties are real. This is because he did
not think they are fixed, any of them. Moreover he seems to have equated
varieties merely with variations, including heritable variations. He clearly
thought of “varieties” as “the slight deviation from a common standard of
which a species is capable” (62), and he allowed for specific characters to
vary as well (25). He allows that “the organization of individuals [of a
species] is capable of being modified to a limited extent by the force of
external causes” and that “these modifications are, to a certain extent,
142 DARWIN AND THE NATURE OF SPECIES

transmissible to their offspring,” but he stresses that species have “fixed lim-
its” (23). Interestingly, Lyell thought of the “common standard” of a species,
along with its complete possible range of variation, as belonging to the on-
tology of each species: “the mutations thus superinduced are governed by
constant laws, and the capability of so varying forms part of the permanent
specific character” (64). On such a view it would therefore be impossible to
say how many varieties a particular species has, even if its full range of vari-
ation were expressed, unless one employed the cynical definition of “variety”
that we have seen in Linnaeus. To all of this, Lyell also added his vote to the
“law of reversion” (28, 33), a law that I shall discuss more fully below, to
which he does not seem to have allowed for any exceptions. Moreover, he
does not seem to have allowed that what he called “extraordinary varieties”
could ever arise in nature and let alone be permanent, for in the case of the
“extraordinary varieties” produced by horticulturalists he says they “could
seldom arise, and could never be perpetuated in a wild state for many gen-
erations, under any imaginable combination of accidents” (32).
Many of these ideas can also be found in Agassiz’s colleague in Amer-
ica, the botanist James Dwight Dana. Dana (1857) thought of each species
as having “a specific amount of concentered force, defined in the act or law of
creation” (486), to which he added that each member organism of a species
has “the specific law of force, alike in all” (487), a clear expression of species
essentialism. As for varieties, Dana argued that “there are variations in
species” and “variations have their limits, and cannot extend to the oblit-
eration of the fundamental characteristics of a species” (492). Again Dana
thought of these as perturbations from the norm caused by outside forces:
“There is then a fixed normal condition or value, and around it librations
take place. There is a central or intrinsic law which prevents a species from
being drawn off to its destruction by any external agency, while subject to
greater or less variations under extrinsic forces” (493). And indeed like Lyell
above, Dana conceived of the ontology of each species as constituting not
only its specific characters but also its full range of possible variation, the
“process of variation” being “the external revealing the internal” (493), such
that “The many like individuals that are conspecific do not properly con-
stitute the species, but each is an expression of the species in its potential-
ity under some one phase of its variables; and to understand a species, we
must know its law through all its cycle of growth [i.e., its ontogenetic stages,
cf. 487–488], and its complete series of librations” (494). In all of this it
seems impossible to find the view that varieties are real. Instead his focus is
on variations and their ranges. Again he says “For while a species has its
constants, it also has its variables” and “The variables are a necessary com-
 The Varieties Problem 143

plement to the invariables; and the complete species-idea is present to the

mind, only when the image in view is seen to be ever changing along the
lines of variables and development.” Thus he says “Whatever individualized
conception is entertained, it is evidently a conception of the species in one
of its phases,—that is, under some one specific condition as to size, form,
colour, constitution, &c., as regards each part in the structure, from among
the many variations in all these respects that are possible” (495). On such
a view, the language of varieties might be helpful, and indeed necessary for
discourse, but the varieties themselves would not be real since they are
picked from ranges of many variables.
One attempt to circumvent this kind of conclusion is to be found in
Thomas Vernon Wollaston (1856), who confined himself to his specialty,
insects. Acknowledging that it has “frequently been asserted that everything
is to be regarded as a ‘variety’ which has wandered in the smallest degree
from its normal state,” Wallace contends that this is “essentially an error”
and asserts in its place that for a variety to be real it “must have in it the
prima-facie element of stability.” Wollaston recognizes that such a variety
might well be taken for a species, and for him “even small differences should
be regarded as specific ones so long as the intermediate links have not been
detected which may enable us to refer them to their nearest types” (6).
Consequently he provides the additional criterion that a variety must have
“intermediate links (which, although rarer than the variety itself, must nev-
ertheless exist) to connect it with its parent stock” (5). For Wollaston, then,
not only must a variety be stable and connected by intermediate links with
its parental stock, but it must also have greater numbers of organisms than
the intermediate links in order to be real. This is an odd foundation on
which to base the reality of varieties, but it is exactly what Wollaston did. He
gives a general example found in many darkly colored insects, namely, that
they “vary, by slow and regular gradations, into a pallid hue, sometimes into
almost white.” For Wollaston, if “the extreme aberration” has greater num-
bers than the intermediate ones, then “there is but a single variety involved,
namely a pale one,—the gradually progressive shades which imperceptibly
affiliate it with its type not being regarded in themselves as ‘varieties’ at all”
(5). Otherwise, says Wollaston, one would have to regard as a variety “every
separate degree of colour which could possibly be found between the outer
limits” (6), and that was something Wollaston was not prepared to do.
Between the two extremes, though, Wollaston was prepared to admit more
than one variety, if, which he says sometimes occurs, “between the two
extremes, there are several nuclei, or centres of radiation, to which the names
of varieties may be legitimately applied” (6).
144 DARWIN AND THE NATURE OF SPECIES

Darwin, interestingly, found Wollaston’s suggestions intriguing, even

having “a good deal of truth” and being “very important” (Burkhardt and
Smith 1990, 171), which should not be surprising given that they lend
themselves to Darwin’s principles of natural selection and divergence. He
consequently in the same year (1856) queried Hooker, Gray, and Watson
in his correspondence to get further facts and corroboration (171, 182,
208). Hooker (July 10) replied that the rare intermediates can always be
explained away by hybridism (176). Gray (early August) replied with his
impression that the rarity of intermediates is often true but that Wollaston’s
approach went against the naming of a species by type specimen, since
the type specimen often turned out to be the rarer form (195). Watson
(December 25) replied similarly that the rarity of intermediates might be
shown by many examples, but he added that it makes the distinction be-
tween species and variety arbitrary since it entails that whatever form is the
more numerous is the species, not the variety, and this would have to be
switched if the numbers accordingly changed (310)—a point he had made
years earlier to Hooker (Burkhardt and Smith 1988, 31). Darwin took all
of this with a grain of salt, and in Natural Selection used the rarity of inter-
mediates as further though minor evidence for gradual evolution, and he
attempted to explain why on his principles they would be expected not to
last (Stauffer 1975, 268–270; cf. Darwin 1859, 176–177).
Wollaston’s position on varieties, of course, was not the norm, and the
evidence for this comes from the widespread belief in the so-called law of
reversion, which should only make us further skeptical that most natural-
ists from the time of Linnaeus to Darwin believed in the objective reality
of varieties, as a real subcategory distinct from variations. We have seen
belief in reversion explicitly in Linnaeus, Jussieu, and Lyell above, while it
is partially advocated by Blyth and is implicit in Dana. Darwin himself
often refers to this law as “well-known” (e.g., 1859, 25), and Wallace
(1859) tells us that it had “great weight with naturalists, and has led to a
very general and somewhat prejudiced belief in the stability of species” (10).
Wallace describes the law as follows: “varieties produced in a state of domes-
ticity are more or less unstable, and often have a tendency, if left to them-
selves, to return to the normal form of the parent species; and this instability
is considered to be a distinctive peculiarity of all varieties, even those occur-
ring among wild animals in a state of nature, and to constitute a provision
for preserving unchanged the originally created distinct species.” He then
adds, with regard to the difficulty the existence of permanent or true vari-
eties posed for these naturalists, that “the difficulty is overcome by assum-
ing that such varieties have strict limits, and can never again vary further
 The Varieties Problem 145

from the original type, although they may return to it, which, from the
analogy of the domesticated animals, is considered to be highly probable,
if not certainly proved” (10–11). Of course for Darwin and Wallace the law
of reversion is not a true law. But what is important for our purposes is
what it does to the supposed reality of varieties, as a legitimate subcategory
of species (which as such would have to have one or more individual organ-
isms) distinct from the category of variation (which is something an indi-
vidual organism has). It would seem to make the subcategory of variety a
mere category of linguistic convenience and nothing more.
This view becomes only strengthened when we examine what natural-
ists commonly thought was the immediate cause of reversion, the explana-
tion of reversion. This was the underlying causal essence of each species (cf.
Ellegård 1958, 209). Aristotle seemed to have this idea in mind (cf. Stamos
2003, 106–107), and we have seen evidence of it above in Linnaeus, Buf-
fon, Jussieu, Agassiz, and Dana, though among them there is of course vari-
ation in the concept. I suggest that these authors were not the exception but
that the concept of each species having an underlying essence was common
in the minds of naturalists from the time of Linnaeus to Darwin, both for
lumpers and splitters, and not simply as a matter of tradition going back to
Aristotle (or even Plato in the case of Agassiz), but more importantly be-
cause of the common belief in, supported in part by widespread evidence
for, the so-called law of reversion.
All of this once again does damage to the view that naturalists of this
period believed in the existence or reality of varieties. Instead it points toward
the view that they used the word “variety” as a mere linguistic convenience,
for the sake of communication. What further supports this view are the
adjectives they used when referring to varieties. Lyell (1832), for example,
writes of “mere varieties” (10, 23) and “a mere temporary and fortuitous
variety” (51). Even Huxley (1859c), in one of his reviews of the Origin,
referred to varieties as “accidental,” “fleeting,” and “mere” (8).
In fact Darwin himself used such terms when referring to varieties, in
particular in his correspondence with other naturalists, whom he well knew
did not think much of varieties. For example, in a letter to Asa Gray (Feb-
ruary 21, 1858) he wrote “I know what fleeting & trifling things varieties
very often are,” to which he added “but my query applies to such as have
been thought worth marking & recording” (Burkhardt and Smith 1991,
27). Moreover he knew from his main correspondent, J.D. Hooker, that
varieties are not important to most naturalists. Hooker himself expressed to
Darwin (July 5, 1845) his own view that “mutation” and “Hybridizing
&c” are “the perturbing causes of our difficulties in assigning limits to
146 DARWIN AND THE NATURE OF SPECIES

species” (Burkhardt and Smith 1987, 211), but more importantly he once
wrote to Darwin (March 14, 1858) that “The long & short of the matter
is, that Botanists do not attach that definite importance to varieties that
you suppose” (Burkhardt and Smith 1991, 49).4
This was hardly a revelation to Darwin. Even in his early notebooks we
can see him recognize the lack of importance naturalists commonly gave to
varieties. For example, in Notebook C (137) he wrote “The simple expres-
sion of such a naturalist ‘splitting up his species [‘& genera’ later added]
very finely’ show how arbitrary & optional operation it is.—show how
finely this series is graduated.—Dr Beck doubt if local varieties should be
remembered, therefore do not consider it as proved that they are varieties,
(though that would be best)” (Barrett et al. 1987, 281). Of course Darwin
needed varieties to be real so they can be incipient species, hence “that
would be best.” And yet he also knew what he was up against. Roughly one
hundred years earlier Joseph Butler, in his Analogy of Religion published in
1736, expressed the prevalent underlying philosophy when he wrote “The
only distinct meaning of the word ‘natural’ is stated, fixed, or settled,” a quo-
tation Darwin added to the frontispiece of the second edition of the
Origin (Burkhardt et al. 1993, 334 n. 11).
So what then was Darwin’s own view on varieties? In previous chap-
ters I have reconstructed his species concept. It remains now to reconstruct
his variety concept. Perhaps the first obstacle to overcome is the claim that
Darwin, like so many others, confused “variation” with “variety.” Accord-
ing to Simpson (1961, 177–178), in many naturalists of the time and
beyond it was common to use the term “variety” to denote an individual
variant, a group of such variants not forming a population, or a population
of such variants distinguishable from the norm of the species. He then goes
on to say that Darwin was guilty of this confusion and that it is “probably
the most serious logical ambiguity in the usually severely careful work of
Darwin” (178). Equally severe is Mayr (1982), who argues that Darwin’s
early association with botanists caused him to use the word “variety” am-
biguously to denote either individual variants or geographic varieties.
According to Mayr, “instead of using the term ‘variety’ consistently for
geographic races, he frequently employed it, particularly in his later writ-
ings, as a designation for a variant or aberrant individual. By this extension
of the meaning of the term ‘variety,’ Darwin confounded two rather differ-
ent modes of speciation, geographic and sympatric speciation” (268; cf.
288, 415, and Mayr 1991, 32–33).
Mayr (1982, 496) also makes the same claim about Wallace, that he
confounded “variation” and “variety,” to which Kottler (1985, 375–378)
 The Varieties Problem 147

provides detailed support. In the case of Darwin, however, I just don’t see
it. For a start, there is his connection with Blyth. We know that Darwin
had some considerable correspondence with Blyth in 1855, and also that he
thought very highly of Blyth, as he stated in the Origin (18). But most of all,
Darwin had an annotated copy of Blyth’s article on varieties in his library
(Burkhardt and Smith 1989, 468 n. 11), so we know he had read Blyth
(1835). From those facts alone it would be remarkable if Darwin went on
to confuse “variation” with “variety.” And indeed when we turn to the
Origin, we do not see Darwin making this mistake (except perhaps for some
minor lapses here and there). Instead he distinguished the two concepts fairly
well. For example, when writing on what it is that natural (and by analogy
artificial) selection is supposed to work on, Darwin repeatedly says that it is
“variations,” never that it is “varieties” (e.g., 30, 61, 80, 84, 102, 108, 169,
454–455, 467, 469). Darwin also says that it is “varieties” that are incipient
species, never that it is “variations” or “variants” (e.g., 52, 54, 59, 111, 133,
169, 176, 325, 404, 469–470). Moreover near the beginning of this chap-
ter we have seen Darwin make an explicit distinction, in both Natural
Selection and the Origin, between individual differences and varieties.
Simpson’s claim and Mayr’s following it just do not hold up.
As for Darwin on speciation, he was a pluralist, much like many biol-
ogists today (including Mayr himself). Speciation by geographic isolation
always remained important for him. As he stated in his concluding chap-
ter in the Origin, “Widely ranging species vary most, and varieties are often
at first local,—both causes rendering the discovery of intermediate links
less likely. Local varieties will not spread into other and distant regions until
they are considerably modified and improved; and when they do spread, if
discovered in a geological formation, they will appear as if suddenly created
there, and will be simply classed as new species” (464–465; cf. 301). As for
sympatric speciation, we have seen in chapter 3 that Darwin in the Origin
favored this more than geographic speciation (isolation). But this was not
because of any conceptual confusion between “variation” and “variety.” In-
stead it was because of the importance of the role he allotted to his princi-
ple of divergence.
Further support for the view that Darwin did not confuse the two con-
cepts becomes evident when we look closely at how he distinguished vari-
eties from species.
To begin, in the concluding chapter of the Origin he says “Hereafter we
shall be compelled to acknowledge that the only distinction between species
and well-marked varieties is, that the latter are known, or believed, to be
connected at the present day by intermediate gradations, whereas species
148 DARWIN AND THE NATURE OF SPECIES

were formerly thus connected” (485). In his barnacle work Darwin put this
criterion into practice. For example, he says (1854b) “In determining what
forms to call varieties, I have followed one common rule; namely, the dis-
covery of such closely allied, intermediate forms, that the application of a
specific name to any one step in the series was obviously impossible; or,
when such intermediate forms have not actually been found, the knowledge
that the differences of structure in question were such as, in several allied
forms, certainly arose from variation” (156; cf. 197).
But this was not the only difference between species and varieties that
we can find in Darwin’s mature writings. The fact is, he advocated other
criteria as well. Again in the Origin he says “Undoubtedly there is one most
important point of difference between varieties and species; namely, that
the amount of difference between varieties, when compared with each other
or with their parent-species, is much less than that between the species of
the same genus” (57). The phrase “amount of difference” should ring a
bell, for in chapter 5, in particular in the case of primroses and cowslips,
we have seen that the intergrading criterion was overruled by the amount
of difference criterion. But we also saw in that chapter that in addition to
amount of difference was the criterion, for species, of constant and distinct
characters, and also that behind that criterion was the criterion of adapta-
tions. This, it turns out, is the real distinction for Darwin between un-
equivocal varieties and unequivocal species, and it is time to bring it out
more clearly.
First, any character upon which a variety is based must for Darwin be
heritable. As he puts it in the Origin, “Any variation which is not inherited
is unimportant for us” (12). Moreover, the variations upon which varieties
are based must not be constrained by barriers such as reversion but must in
principle be capable of varying to the point of specific difference. Again in
the Origin Darwin says “It has often been asserted, but the assertion is quite
incapable of proof, that the amount of variation under nature is a strictly
limited quantity” (468). All of this, which is necessary if varieties are to be
“incipient species,” is enough to make Darwin’s concept of variety remark-
ably different from those of his contemporaries.
But of course there is more. Darwin distinguished between individual
variants and varieties, the difference being that the latter is a subtaxon or
the beginning of one, and moreover that natural selection is in operation
toward making it a species. This comes out rather clearly in Natural Selec-
tion (Stauffer 1975), where Darwin distinguishes between species and
varieties, the former in his view “being much more constant in all their
characters,” this constancy being “partly due to the several causes of vari-
 The Varieties Problem 149

ability having acted less energetically on the two species under comparison
than on the one species yielding the two or more varieties; and partly to the
characters of the two species having been long inherited, & by this cause
having become more fixed.” Hence, he says a few lines later, “as a general
rule, species may be looked at as the result of variation at a former period; &
varieties, as the result of contemporaneous variation” (165). This is from his
chapter IV titled “Variation Under Nature,” which comes two chapters be-
fore his chapter VI titled “On Natural Selection.” When Darwin does get to
that chapter, he connects this constancy of characters with natural selection:
In regard to the difference between varieties and species, I may add that
varieties differ from each other & their parents, chiefly in what naturalists
call unimportant respects, as size, colour, proportions &c; but species dif-
fer from each other in what naturalists consider more important respects.
But we have seen in Chapter IV, that varieties do occasionally, though
rarely, very [sic, vary] slightly in such important respects; and in so far as
differences in important physiological characters generally stand in direct
relation to different habits of life, modifications however slight in such
characters would be very apt to be picked out by natural selection & so
augmented, thus to fit the modified descendants from the same parent to
fill as many & as widely different places in nature as possible. [244]

As for the origin of a variety, in particular a variety “in some degree per-
manent,” back in Chapter IV Darwin says “whether it has originated in a
single accidental variation, or by the addition of several such successive vari-
ations through natural selection, or through the direct & gradual action of
external conditions, as of climate, its first origin is even of less importance
to it, than its preservation” (137). Its preservation, of course, as well as its
modification into a species, will be effected by natural selection. This is what
makes varieties incipient species. And naturally Darwin was ready to admit
that he was “far from supposing that all varieties become converted into
what are called species: extinction may equally well annihilate varieties, as it
has so infinitely many species” (159).
In all of this we can see that Darwin had a clear distinction in mind be-
tween variation and variety. This comes out, for instance, later in chapter VI:
It should always, be borne in mind that there is a wide distinction be-
tween mere variations & the formation of permanent varieties. Variation
is due to the action of external or internal causes on the generative sys-
tems, causing the child to be in some respects unlike its parent; & the dif-
ferences thus produced may be advantageous or disadvantageous to the
child. The formation of a permanent variety, implies not only that the
150 DARWIN AND THE NATURE OF SPECIES

modifications are inherited, but that they are not disadvantageous, gen-
erally that they are in some degree advantageous to the variety, otherwise
it could not compete with its parent when inhabiting the same area. The
formation of a permanent variety must be effected by natural selection;
or it may be the result, generally in unimportant respects, of the direct
action of peculiar external conditions on all the individuals & their
off-spring exposed to such conditions. [240]

All of this was prefigured by Darwin in one of his many letters to

Hooker (November 23, 1856), written a few weeks before he began the
first draft of c hapter IV of Natural Selection and roughly five months
before he completed the first draft of chapter VI (Stauffer 1975, 213):

. . . my conclusion is that external conditions do extremely little, except

in causing mere variability. This mere variability (causing the child not
closely to resemble its parent) I look at as very different from the forma-
tion of a marked variety or new species.—(No doubt the variability is
governed by laws, some of which I am endeavoring very obscurely to
trace).—The formation of a strong variety or species, I look at as almost
wholly due to the selection of what may be incorrectly called chance vari-
ations and variability. [Burkhardt and Smith 1990, 282]

In all of this, we can see not only that Darwin, contra Simpson and Mayr,
did not confuse “variation” with “variety,” but that he developed a concept
of variety that was revolutionarily different from anything that had been used
before. By raising the importance of variations, which hitherto were thought
to be unimportant, by raising the reality of varieties and redefining them as
incipient species, which hitherto were thought to be neither, and by focus-
ing on the production of adaptations by natural selection, Darwin accom-
plished, or attempted to accomplish, an enormous paradigm shift.
But in saying that between species and varieties there is no essential
difference Darwin was not saying that species and varieties are not real. In-
stead he was simply saying that there is no barrier preventing a real variety
from becoming a real species. To say that there is an essential difference is
to say that there is such a barrier. For species essentialists in Darwin’s day,
there was such a barrier. As Watson wrote in a letter to Darwin (May 10,
1860), “Until a faith in certain impassable barrier between existent species
becomes thoroughly shaken, naturalists will resist your views, & hail diffi-
culties as if conclusive arguments on the contra side. Differently as these un-
seen barriers are traced or placed, they are believed in about as strongly by
almost all” (Burkhardt et al. 1993, 203). What Darwin was doing was
 The Varieties Problem 151

arguing against the existence of that barrier. In a very important way, then,
ironically, his concept of variety, not his concept of species, was the
Archimedean fulcrum with which he would shake and eventually move
the world.
The fact remained, however, that Darwin’s contemporary naturalists
did not think of varieties, and consequently of species, as he himself did. In
order to shake their faith in what Watson called the “certain impassable
barrier,” more than straight exposition must have seemed to Darwin
required. We are thus led to Darwin’s strategy.
This page intentionally left blank.
 Chapter 8

Darwin’s Strategy

If the previous analysis is basically correct, it raises the obvious question of

why Darwin would define “species” nominalistically, both taxa and cate-
gory. If he was indeed a realist, it seems extremely odd that he would do
this, especially since it is duplicitous and he was, after all, a man of honor.
In this chapter I will develop and defend a novel answer to this problem.
Since Darwin, as far as is known, never provided an explicit answer to this
question, and since it is obviously too late to get inside his head in the
manner of a Schilpp volume, the best that we can do, once again, is to
apply the principle of charity and make the best of the evidence that we can.
I believe that the answer I develop has considerable evidence in its support.
But first I want to look at the answers given by Ghiselin, Mayr, Beatty,
Sulloway, Hodge, McOuat, and my former self. The failings of their an-
swers will further justify the search for a new solution.
Ghiselin (1969), as we have seen in chapter 2, argued that Darwin
was a species taxa realist, going as far as to characterize him as a precursor
of the species-as-individuals view. When it came to the species category,
however, Ghiselin characterized Darwin as a nominalist. Ghiselin’s justi-
fication for this conclusion hinges on his claim that Darwin had an Aris-
totelian concept of category, a concept characterized by strict essentialism.1
Since varieties, in Darwin’s view, gradually evolve into species if they do
not first go extinct, then contrary to the creationists there cannot be an es-
sential distinction between species and varieties. So this is why the term

153
154 DARWIN AND THE NATURE OF SPECIES

“species” cannot on Darwin’s view be defined. As Ghiselin (1969) put it,

Darwin “insisted on Aristotelian definition as a criterion of reality or nat-
uralness. To Darwin, as to many other taxonomists, an inability to give rig-
orous definitions for the names of taxonomic groups led to a belief that
somehow such assemblages were artificial” (82). Ghiselin claims further
that Darwin “maintained that there are no ‘essential’ differences between
species and varieties, and that both terms designate the same basic kind of
entity.” Thus, says Ghiselin, “Darwin was denying the reality, not of taxa,
but of categories” (93).
There are a number of problems with this view. First, as we have seen,
Darwin did not think that species and varieties are basically the same kind
of entity. Although they have a lot in common in Darwin’s view, from
the perspective of the horizontal dimension (and that as we have seen is
the dimension for Darwin in which species have their primary reality)
there is a fundamental distinction, most notably natural selection and the
fixity of adaptations.
Second, there is no reason to believe that Darwin subscribed to Aris-
totelian essentialism, with the corresponding concept of definition (genos
and differentiae), as “a criterion of reality or naturalness.” For a start, Dar-
win never read Aristotle (cf. Burkhardt and Smith 1988, 498 n. 9). If he
subscribed to Aristotelian essentialism for the reality of categories, then, his
knowledge of Aristotle must have been secondhand. This knowledge was,
of course, part of the worldview that Darwin inherited. Roughly a century
before Darwin’s Beagle voyage, Linnaeus wedded the Aristotelian paradigm
to Christian creationism. This would be the received view for over the next
hundred years (Stamos 2005). It was a view, as we have seen in chapter 1,
that involved essentialism for membership in categories and bridgeless gaps
between those categories, conceptualized as boxes, with accidental
(nonessential) characters being irrelevant.
Although Darwin had indeed read Linnaeus, and of course the latter’s
worldview was (roughly speaking) the established view among Darwin’s
scientific contemporaries, at least for species, there is nevertheless nowhere
in Darwin’s writings where he explicitly subscribes to Aristotelian or Lin-
nean criteria for the reality of categories. Granted, he makes statements
which could be interpreted in that way. For example, in Natural Selection
he says “it is no wonder that there should be difficulty in defining the dif-
ference between a species & a variety;—there being no essential, only an
arbitrary difference” (Stauffer 1975, 98). Moreover, as we have seen earlier,
near the end of the Origin he says “we shall at least be freed from the vain
search for the undiscovered and undiscoverable essence of the term species”
 Darwin’s Strategy 155

(485). But these statements are no more to be taken at face value than his
nominalistic claims about species taxa. The fact is that Darwin, as we have
seen, in spite of his evolutionary gradualism, thought that species taxa are
real. And it is especially important here to pay attention to his language. As
we saw in chapter 3, Darwin in the Origin expressed his belief that “species
come to be tolerably well-defined objects, and do not at any one period pre-
sent an inextricable chaos of varying and intermediate links” (177, italics
mine). Moreover near the end of the Origin he concluded that “Systema-
tists will have only to decide (not that this will be easy) whether any form
be sufficiently constant and distinct from other forms, to be capable of
definition; and if definable, whether the differences be sufficiently important
to deserve a specific name” (484, italics mine). And again, as we have seen
in Descent (1871 I), he says “Every naturalist who has had the misfortune
to undertake the description of a group of highly varying organisms, has
encountered cases . . . precisely like that of man; and if of a cautious dispo-
sition, he will end up by uniting all the forms which graduate into each
other as a single species; for he will say to himself that he has no right to give
names to objects which he cannot define” (226–227, italics mine).
In all of this there is no evidence that Darwin was using the term “de-
finition” in an Aristotelian sense. Quite the contrary, all he meant was
description. Hence his use in the Origin of the phrase “good and distinct
species” (47, 61, 259). All he was referring to were species that are distinct
enough to be described, as in a species monograph. And Darwin, of course,
had plenty of experience doing that, given his eight years of work on bar-
nacles. Moreover there was nothing idiosyncratic in his use of the term “de-
finition.” For example, Hugh Strickland, in his first attempt at rules for
zoological nomenclature (Strickland 1837), wrote in explanation of his
Rule 7 (“A name may be expunged which has never been clearly defined”)
that “Unless a group is defined by description or figures when the name is
given, it cannot be recognised by others; and the signification of the name
is consequently lost. . . . Many collectors of shells and fossils are in the habit
of labelling those species which they do not find described, with names of
their own invention; but, unless they publish descriptions of these new
species, they cannot expect these names to stand” (174).
Adding to all of this the evidence from chapters 2 and 3 that Darwin
was a species taxa realist, it becomes evident that Darwin’s concept of def-
inition for individual taxa allowed for the reality of taxa even though his
theory of gradual evolution precluded essentialism for those taxa. We can
only conclude, then, that when it came to taxa Darwin had a realist though
non-Aristotelian concept of definition. It seems odd, then, that Darwin
156 DARWIN AND THE NATURE OF SPECIES

would not extend this non-Aristotelian concept of definition to categories

as well. But there is no reason to conclude that he did not. In fact the prin-
ciple of charity dictates that we ascribe to Darwin (even if only at an intu-
itive level) a non-essentialistic concept of definition for the objective reality
of categories, consistent with his non-essentialistic concept of definition
for the objective reality of taxa. And if we do this, we place Darwin on per-
fectly logical ground. Strictly essentialistic categories, categories that imply
bridgeless gaps among other things, are not the only kind of realist cate-
gories. Night grades into day and day into night, and not only are partic-
ular days and nights real but so are the two categories of night and day, yet
neither category is strictly essentialistic.
By ascribing to Darwin a non-Aristotelian concept of definition and
categories, we also place Darwin closer in a conceptual sense to the mod-
ern day. This is because it is common today among biologists who proffer
or subscribe to a realist species concept to acknowledge that because of
gradual evolution there are going to be “messy situations” in the applica-
tion of their particular species concepts. A good example is Ernst Mayr. As
we saw in chapter 2, Mayr argued that the species category, as defined by
the biological species concept, is extra-mentally real. And yet he repeatedly
acknowledged the existence in the biological world of what he called “messy
situations.” For example, Mayr (1982) states, in response to the often-
repeated criticism that the biological species concept does not work well
with plants (which we have seen in chapter 6), that “To be sure there are
‘messy’ situations . . . but I am far more impressed by the clear distinction
of most ‘kinds’ of plants I encounter in nature than by the occasional
messes” (285). Moreover Mayr later conducted a study on a local flora in
the township of Concord, Massachusetts (Mayr 1992), in which he con-
cluded that hybridization accounts for messy situations in 9.2% of the Con-
cord flora (⫽ 90.8% “good” species), which he thought might be reduced
to 6.6% following further study (236). This analysis is in close accordance
with McDade’s (1995) survey of 104 recent botanical monographs, cover-
ing 1,790 species (mostly sexual), in which she concludes from her data
that “something on the order of 15% of species are, at this horizontal time
slice, involved in one or more biological processes that blur species bound-
aries or lead to infraspecific (and subsequently specific?) differentiation”
(614). Moreover she concludes that “Variation was reported as sufficient to
cause problems in delimiting about 7% of species treated” (613). Darwin
was therefore justified, from a modern point of view, in stating his belief in
the Origin that “species come to be tolerably well-defined objects, and do
not at any one period present an inextricable chaos of varying and interme-
 Darwin’s Strategy 157

diate links.” All things considered, his belief in the relatively low propor-
tion of messy situations at any one horizontal level did not preclude species
category realism on his part any more than it does for modern biologists
with their various definitions of “species.”
Another strategy theory is provided by Mayr (1982). Mayr takes Dar-
win’s nominalistic statements on species, examined in chapter 1, pretty much
at face value, so that for Darwin species are “purely arbitrary designations”
(269). Moreover Mayr points the finger at botanists (the bane of Mayr’s own
species concept) for influencing Darwin toward species nominalism, espe-
cially William Herbert, whom he quotes as having written “there is no real
or natural line of difference between species and permanent or discernible
[‘descendible’ in the original] variety” (268). But the main thrust of Mayr’s
view is that Darwin had a “strong, even though perhaps unconscious, moti-
vation . . . to demonstrate that species lack the constancy and distinctiveness
claimed for them by the creationists” (269). On the prevailing creationist
view, species must have distinct boundaries, even if difficult to determine by
humans, for God in his perfection would surely not create a world with, or
with the potential for, a confusion of forms. Indeed as Darwin put it in the
Origin, “The view generally entertained by naturalists is that species, when in-
tercrossed, have been specially endowed with the quality of sterility, in order
to prevent the confusion of all organic forms” (245). Of course such bound-
aries would make no sense from the viewpoint of gradual evolution by nat-
ural selection. So Darwin, according to Mayr, had to deny the boundaries in
order to make a case for his view of evolution. “Hence,” he says, “it was good
strategy to deny the distinctness of species” (269). This is why Mayr thinks
Darwin focused on degree of difference rather than on reproductive isola-
tion. By doing so he could make a more convincing case for species being
“purely arbitrary designations” and thus for evolution.
There are a number of problems with Mayr’s claims. First, there is a
tension between his claim that Darwin was a species nominalist and his
further claim that for Darwin “species continue to evolve” (269). One won-
ders how species can do this if they are “purely arbitrary designations.”
Second, as we have seen in detail in chapter 1, in spite of William Her-
bert virtually all botanists in Darwin’s time (and not just botanists) were
species realists.
Finally, there is equally no reason to accept Mayr’s claim that Darwin
focused on degree of difference rather than on reproductive isolation as a
matter of strategy. As we have seen in Chapter 6, Darwin rejected repro-
ductive isolation, principally in the form of sterility, because he did not think
that such a character could be formed by natural selection. Accordingly,
158 DARWIN AND THE NATURE OF SPECIES

species defined in terms of reproductive isolation would not on his view be

part of a vera causa system of classification.
The most influential strategy theory has been provided by John Beatty
(1985),2 which is an improved version of his previous work (Beatty 1982).
Because of its influence, it is important to examine it in some detail. To
begin, Beatty (1985) acknowledges “Frank Sulloway’s suggestion that Dar-
win’s choice of species concept was guided by ‘tactical’ considerations.
Among those tactical considerations was the decision to employ his fellow
naturalists’ species concept, in order to speak to them ‘in their own lan-
guage’” (265). According to Sulloway (1979), most of Darwin’s fellow nat-
uralists were “morphologists” and “typologists.” Darwin therefore adopted
a “morphological species definition” (38) as part of his tactic. In stressing
the arbitrary distinction between species and varieties, “Darwin spoke to
them in their own language and tailored his argument so as to exploit the
inevitable sense of frustration that virtually all naturalists had at one time
or another experienced in their taxonomic work” (37).
Beatty also acknowledges his debt to Ghiselin’s (1969) interpretation of
Darwin as a species taxa though not a species category realist. Beatty explic-
itly accepts this aspect of Ghiselin’s thesis. In his own words, “Darwin’s ref-
erences to the arbitrariness and unreality of species pertained only to the
species category, not to species taxa. . . . he did not deny the reality of the var-
ious species taxa like the cabbage and the radish” (277). As for species taxa,
he also agrees with Ghiselin that such taxa were for Darwin “chunks with[in]
the genealogical nexus of life” (278). Thus, for Beatty, Darwin’s theory was
not rendered domainless; in spite of its apparent nominalism, it still had
referents, the individual species taxa. In going so far with Ghiselin, Beatty
did not recognize, or at least acknowledge, what I claimed in chapter 2,
namely, that such a view ascribes to Darwin a fundamental contradiction:
if no species category, then “species taxa” becomes a contradiction in terms.
Although Beatty does not recognize a contradiction, he does nevertheless
recognize a fundamental tension. Near the end of his paper he says

The suggestion that natural history could really get by without defini-
tions of the categories of classification—especially a definition of
“species”—is admittedly hard to swallow. Of course, it should be ac-
knowledged that natural history was only temporarily without a defini-
tion of “species.” The non-evolutionary definitions rejected by Darwin
have since been replaced. Definitions such as Ernst Mayr’s “biological
species concept” and George Gaylord Simpson’s “evolutionary species
concept” are already so well entrenched as to be considered traditional.
[280]
 Darwin’s Strategy 159

So far there is nothing really new in Beatty’s theory. Where he departs

from Ghiselin is in the referential use of the term “species” that he ascribes
to Darwin and in the motive behind that use. We have already examined
this aspect of Beatty’s theory in chapter 5, but it is useful to look at it again.
In Beatty’s view, Darwin knew that his theory of evolution by natural
selection was going to be a hard sell. According to Beatty, in writing the
Origin Darwin was not trying to sell his theory to the general public. No
Robert Chambers was he. Instead, he “addressed his theory” to the “com-
munity of ‘naturalists,’ of which Darwin considered himself a member”
(266). The problem was that the species concept that these naturalists used
was so theory-laden with non-evolutionary connotations that he could not
possibly use their definition. But to replace it with an evolutionary species
concept would have only made it more difficult, from a psychological point
of view, to get his theory across to these naturalists. So he took the more
subtle route of denying any reality to the species category, all the while
throughout the Origin employing the word “species” in such a way so that
it “agreed with his fellow naturalists’ actual referential uses of the term”
(269). As Beatty more succinctly put it,

Darwin indeed perceived the difficulty posed by definitional language rules

that undermined the theory he wished to communicate. He tried to get
around this difficulty by distinguishing between what his fellow naturalists
called “species” and the non-evolutionary beliefs in terms of which they
defined “species.” Regardless of their definitions, he argued, what they called
“species” evolved. His species concept was therefore interestingly minimal:
species were, for Darwin, just what expert naturalists called “species.” By try-
ing to talk about the same things that his contemporaries were talking
about, he hoped that his language would conform satisfactorily enough for
him to communicate his position to them. [266]3

But Beatty goes even further than this. Accepting the quotations examined
in chapter 1, where Darwin states that the term “species” is undefinable, Beatty
claims that Darwin did not merely feign species category nominalism but ac-
tually meant it. In rejecting the species concepts of his fellow naturalists and
employing only the referential use of those naturalists, says Beatty, it “allowed
Darwin to communicate the position that the term ‘species’ was undefinable.
In other words, Darwin not only rejected non-evolutionary definitions of
‘species,’ he also rejected the idea that the term could be defined at all” (274).
Since Beatty clearly thinks that the term “species” can indeed be defined from
an evolutionary perspective, given his reference to Mayr and Simpson above,
it follows that he thinks that Darwin was just plain wrong.
160 DARWIN AND THE NATURE OF SPECIES

I have argued in previous chapters that Darwin did indeed have an

evolutionary species concept, albeit implicit. This saves Darwin from being
wrong, not necessarily in his particular species concept, but in thinking, as
Beatty suggests he thought, that gradual evolution precludes a realist species
concept. At any rate, given that Beatty with his strategy theory goes against the
(older) received view so far as to accept that Darwin really was a species taxa
realist, one has to wonder why he does not extend his strategy theory further
and argue that Darwin only feigned species category nominalism, that he was
a species category realist after all. No doubt part of the reason is that Beatty,
like everyone else, has failed to discern a species concept in Darwin’s mature
writings. But there is a further reason, much more overt, which, however, I
must save for later in this chapter when I examine Hooker’s conversion.
There is another aspect of Beatty’s theory that I must save for later. His
theory is clearly part of the larger incommensurability thesis made famous
by Thomas Kuhn (1970), according to which adherents of old and new the-
ories during a scientific revolution can barely if at all communicate their
ideas between each other because of the theory-laden nature of their respec-
tive terminologies. I shall explore this issue, including Beatty’s solution with
regard to Darwin, in the next chapter. For the present, I want to go through
the many reasons why I think Beatty’s strategy theory should be rejected.
This is necessary in order to clear the path for my own strategy theory.
Beginning with problems with Beatty’s strategy theory, then, it is ques-
tionable whether Darwin wrote the Origin exclusively or even primarily for
the community of his fellow naturalists. There is evidence, both within and
outside the Origin, that he intended from the beginning a much wider
audience. For a start, Darwin defines the term “endemic” in “endemic
species” (390), something his fellow naturalists would surely not need done
for them. Indeed the book is remarkably reader friendly all round. More-
over the Origin is filled with euphemisms designed, it would seem, for the
greater sensibilities of the general public. For example, to play down the
shivering coldness and utter horror of natural selection in the wild, Darwin
wrote “When we reflect on this struggle, we may console ourselves with
the full belief, that the war of nature is not incessant, that no fear is felt, that
death is generally prompt, and that the vigorous, the healthy, and the happy
survive and multiply” (79). Elsewhere he wrote of “the beautiful and har-
monious diversity of nature” (169). Comforting words indeed, but Dar-
win’s fellow naturalists would have known better. Even many amateur
naturalists would have known better. Darwin himself, of course, knew
better. In a letter to Hooker (July 13, 1856), for example, he wrote “What
a book a Devil’s chaplain might write on the clumsy, wasteful, blundering
 Darwin’s Strategy 161

low & horridly cruel works of nature!” (Burkhardt and Smith 1990, 178).
Similarly in a letter to Asa Gray (May 22, 1860) he wrote “There seems to
me too much misery in the world. I cannot persuade myself that a benefi-
cent & omnipotent God would have designedly created the Ichneumonidæ
with the express intention of their feeding within the living bodies of cater-
pillars, or that a cat should play with mice” (Burkhardt et al. 1993, 224).
There is further evidence of an intention for a wider audience that is even
more direct. Interesting is Darwin’s correspondence with his publisher,
John Murray. Although Darwin admitted in one of his letters to Murray
(April 2, 1859) that “My volume cannot be mere light reading, & some
parts must be dry & some rather abstruse,” he immediately added that “yet
as far I can judge perhaps very falsely, it will be interesting to all (& they are
many) who care for the curious problem of the origin of all animate forms”
(Burkhardt and Smith 1991, 277; cf. 278, 440, and Burkhardt et al. 1993,
240). Given the high sales through many editions of Robert Chambers’
anonymous Vestiges of the Natural History of Creation, beginning in 1844,
Darwin could not have helped but know that there would indeed be
“many” among the general public who would read his book.
A second and more important problem with Beatty’s theory concerns
his claim about Darwin’s referential use of the term “species.” I have argued
extensively in chapters 4, 5, and 6 that Darwin did not simply follow the
referential use of his fellow expert naturalists with regard to this term. In
fact, most instructive are the cases where Darwin went against that referen-
tial use. Indeed it was those cases that largely helped form my reconstruc-
tion of Darwin’s species concept.
Third, again in reference to Darwin’s referential use of the term
“species,” it must be stressed that Darwin’s fellow naturalists often did not
provide him with a general consensus of referential use of species names. In
many cases they themselves could not agree. Nowhere was this more evident
to Darwin than in the case of barnacles. Characteristic of his frustration
was what he wrote in a letter to J.S. Henslow (July 2, 1848):

I am anxious to make out the distribution of the British species—And

new species may turn up, for the group has been made out most super-
ficially,—for instance under Balanus punctatus (which must be made a
distinct genus) three or four varieties have been called distinct species;
whereas one form, which has not been called even a variety, is not only
a distinct genus, but a distinct sub-family.—Yesterday I found four or 5
named genera are all the closest species of one genus: this will give you
a specimen of the utter confusion my poor dear Barnacles are in.
[Burkhardt and Smith 1988, 156]
162 DARWIN AND THE NATURE OF SPECIES

The situation was more widely reflected in the reality of species lumpers ver-
sus splitters. Even lumpers and splitters each among themselves often could
not agree. As H.C. Watson put it in a letter to Darwin (March 24?, 1858),
after tabulating the number of species variously allocated to six genera by
eight botanists, “Thus, whether we compare together different Authors
publishing at nearly the same dates, or the same Author publishing at dif-
ferent dates, much discrepancy appears in their ideas of species. . . . they
[species] are far indeed from being fixed and certain in books” (Burkhardt
and Smith 1991, 55–56). Far from providing Darwin with a referential con-
sensus, then, Darwin’s fellow naturalists often provided him with the oppo-
site. Moreover they even lacked a consensus on the definition of the term.
Indeed this is a theme that is repeated in the Origin. For example, Darwin
says “No one definition has as yet satisfied all naturalists; yet every natural-
ist knows vaguely what he means when he speaks of a species” (44). Again,
with regard to forms that naturalists could not agree on whether they are va-
rieties or species, he says “to discuss whether they are rightly called species
or varieties, before any definition of these terms has been generally accepted,
is vainly to beat the air” (49). The theme would later be repeated in Descent
(Darwin 1871 I), wherein, on the issue of whether the human races consti-
tute one or several species, he says “it is a hopeless endeavour to decide this
point on solid grounds, until some definition of the term ‘species’ is gener-
ally accepted; . . . We might as well attempt without any definition to de-
cide whether a certain number of houses should be called a village, or town,
or city” (228). Given, then, the already prevalent disagreements on the ref-
erential use of the term “species,” and even on its definition, it would not nec-
essarily have been a great impediment to the acceptance of Darwin’s
evolutionary views were he to redefine the concept.
A fourth problem with Beatty’s theory, and the final one that I shall
focus on, is his suggestion that Darwin not only learned of the problem of
the theory-ladenness of species concepts from the writings of Watson
(1845a, 1845b), but that Watson’s writings “might also have suggested to
Darwin a means of dealing with that difficulty” (271), which is the same
means that Beatty attributes to Darwin. Much of Beatty’s discussion on
Watson cannot be contested. There is an error nevertheless, and the crux
of it is in his use of the following quotation from Watson (1845a). Beatty
writes that Watson “proposed to ‘write of “species” as commonly under-
stood by botanists, without attempting any rigorous definition of the term,
which may hereafter be found to represent only a fiction of the human
mind’ (1845a, p. 142)” (273). In spite of the drift of the last part of this pas-
sage, it is important to emphasize that nowhere in Watson’s writings, in
 Darwin’s Strategy 163

spite of his conversion to evolutionism apparently as a result of Chambers’

Vestiges, does he clearly and unequivocally advocate species category nom-
inalism or species taxa nominalism. To the quotation above he prefixes it
with the words “For the present I must write of ‘species’ as . . . .” Moreover
he uses the word “may” when he writes of the word “species” representing
a fiction of the mind. He does not actually say that it is a fiction. In fact,
as we have already seen in chapter 1, although Watson came close at one
point to embracing species nominalism, he never went all the way, and
continued to think of real species in nature. He did indeed think of cate-
gories of higher taxa as indeed fictions of the mind (which was a common
view), but he did not think the same for the species category. Moreover
this was made clear to Darwin not only in Watson’s writings but also in
Watson’s correspondence to Darwin. There remains no reason, then, to
think that Darwin was following a strategy laid out earlier by Watson.
There was no such strategy, and what Darwin did was entirely different
from what Beatty suggests.
Parasitic on Beatty’s theory is the theory of M.J.S. Hodge (1987).
According to Hodge, “only for some twelve months (late 1837 to late 1838)
did he [Darwin] articulate and uphold a definition of species. Before that
and afterwards he was consistently content not to attempt to contribute
one more to the growing number of definitional proposals in the litera-
ture” (233). During the period mentioned by Hodge, Darwin explicitly
endorsed, as we have seen in chapter 1, a species concept based on repro-
ductive isolation. According to Hodge, this ceased once Darwin formu-
lated his theory of natural selection, which Hodge places in 1838 “in late
November and early December (not, contrary to legend, in September)”
(240).4 Hodge explicitly follows Ghiselin in ascribing to Darwin, after this
period, species category nominalism combined with species taxa realism,
and he explicitly follows Beatty in ascribing to Darwin no change in the ref-
erence (denotation, extension) of the term “species” from his fellow natu-
ralists. But Hodge adds to Beatty’s theory a new twist. Beatty (1985) argues
that with regard to the non-evolutionary theory and corresponding defin-
ition of “species” of his fellow naturalists, Darwin had “at least two op-
tions”: (i) he might respect the language rules of his fellow naturalists and
agree that the term “species” is to be used only for non-evolutionary enti-
ties but “object that there is nothing in the world that actually satisfies the
definition of the term,” or (ii) he might respect not the language rules but
the practices of his fellow naturalists, the “examples of his fellow community
members’ use of the term ‘species’,” but object to the use of the term for
only non-evolutionary entities, such that “the old theory about species, and
164 DARWIN AND THE NATURE OF SPECIES

the theory-laden definition of the term, are substantial mischaracterizations

of things that the community members have actually called ‘species’” (268).
According to Beatty, Darwin chose the second option because he thought
it would be better for communicating his theory of evolution and for con-
verting his fellow naturalists to his view. According to Hodge, Darwin did
not even have a choice. As Hodge puts it, “Darwin was not setting out
merely to find causes for those entities that were included in the reference
of the term species. He was setting out to find causes for those properties
that those entities had by virtue of which properties they counted, accord-
ing to accepted criteria, as species and not as varieties. So Darwin’s prob-
lem situation presupposed acceptance of the way the meaning or intension
of the term species, as given by those criteria, had settled, determined, the
reference of the term as it had done” (249).
Since Darwin did not, as I have shown, simply follow the referential use
of his fellow naturalists with regard to species designations, his motive could
not have been, contrary to what Hodge thinks, merely to determine the
cause or causes of the specific characters as determined by his fellow natu-
ralists. As we saw in chapters 4, 5, and 6, Darwin did not always follow
species designations but often revised in accordance with his own evolution-
ary species concept. Moreover some of the characters that his fellow natu-
ralists thought to be specific were accepted by Darwin if they had been
fixed by natural selection. But other characters were rejected, such as steril-
ity. Ghiselin (1969) points out that with Darwin “Classification ceased to
be merely descriptive and became explanatory” (83). To my mind it be-
came more, much more, to the point of actually altering some of the clas-
sifications of his contemporaries.
A further twist on Beatty’s theory is provided by Gordon McOuat.
McOuat (1996, 475, 2001, 3 n. 9) explicitly relies on Beatty’s theory. More-
over McOuat makes much of what he (McOuat) calls, following Kitcher,
the “cynical definition of species,” which he says “was Darwin’s only ‘defin-
ition’ of species” (1996, 515), the “only one clear definition of species in
any of his work” (2001, 4 n. 10), explicitly given in Natural Selection and
echoed in the Origin, viz., “In the following pages I mean by species, those
collections of individuals, which have commonly been so designated by
naturalists” (Stauffer 1975, 98). (Interestingly, when quoting this passage
McOuat left out the word “commonly.”) For McOuat, Darwin was not try-
ing to bypass the theory-laden definition of “species” held by the majority
of his fellow naturalists. For one thing, McOuat is skeptical that there was
any consensus there at all. As he puts it (1996), “against the usual historical
account, it is unclear whether there really was a monolithic definition of species
 Darwin’s Strategy 165

accepted by Darwin’s fellow naturalists, or even the hope of one. Moreover,

science may not work like that anyway. Definitions are not at the core”
(475). Instead of definitions, according to McOuat, what was important to
Darwin’s contemporary naturalists was “property rights,” in particular the
issue of whose nomenclature for taxa got to stick in the language commu-
nity of the naturalists. This was especially important given the different sys-
tems of classification and nomenclature that were competing for acceptance
at the time. Hence the importance of Strickland’s (1837) initial rules for
zoological nomenclature and later for the rules of a committee drawn to-
gether and reported by Strickland (1842). Of this committee, says McOuat
(1996), “it was Darwin who convinced Strickland to co-opt the new mem-
bers, from his own circle of contacts. Darwin soon became the main conduit
between members, . . . Strickland was the organizational force behind the
new commission, Darwin the fulcrum” (507). Of these rules, the law of pri-
ority would be “the one prime directive” (510). But more than that, says
McOuat, in this publication of rules “the species category itself is not de-
fined. This was not an ‘essentialist’ species concept, species with real ‘defi-
nitions,’ ‘creationist’ species, ‘permanent’ species, or whatever our histories
have led us to believe. Rather, the solid core of conventional names marked
something unsaid, yet something ‘agreed’ upon on trust: species are nodes
in the communication of a network of ‘competent’ naturalists” (511). Thus
for McOuat the stability of species came not from nature but from the need
“to maintain the stability of naturalist discourse” (511). More specifically,
“It was the restraint of language—the necessity of keeping names and their
referents while all else, including definitions, remained ‘up in the air’” (515).
And it was this consideration, says McOuat, which explains Darwin’s con-
servatism in accepting the species designations of his fellow naturalists. As
he puts it, “Darwin, ever cautious, ever the conservative of revolutionaries,
signed the document, knowing that soon he was to invoke the greatest
change of ‘meaning’ while at the same time faithfully retaining the inherited
names and stabilities of natural history and its objects” (514).
This is an interesting strategy theory indeed. Unfortunately it fails on
at least two counts. First, in Strickland (1842) “the true system of nature”
is assumed and the rules are meant to apply only “when naturalists are
agreed as to the characters and limits of an individual group or species, [and
when] they still disagree in the appellations by which they distinguish it”
(106). Moreover there is a repeated appeal to follow the Linnean binomial
system, going so far as to say “Two things are necessary before a zoological
term can acquire any authority, viz. definition and publication. Definition
properly implies a distinct exposition of essential characters, and in all cases
166 DARWIN AND THE NATURE OF SPECIES

we conceive this to be indispensable, . . . publication, nothing short of the

insertion of the above particulars in a printed book can be held sufficient”
(113–114). This points to what we have seen in Chapter 1, that in spite of
the variety of species definitions competing at the time there was a common
core, a core involving realism and what Wollaston called an “axiom” involv-
ing independent creation and non-transition. The different definitions in-
volved, instead, different diagnostic criteria. Moreover I have argued in the
previous chapter and elsewhere (Stamos 2005, 92–93) that there was a
common core involving an underlying causal essence. J.D. Dana (1857), for
example, expressed this view when he compared biological species to chem-
ical elements, the main difference being that individuals of a biological
species reproduce, such that “when individuals multiply from generation to
generation, it is but a repetition of the primordial type-idea; and the true
notion of the species is not in the resulting group, but in the idea or poten-
tial element which is at the basis of every individual of the group” (487).
Ellegård (1958, 210) calls this the traditional view, and I would concur.
Secondly and equally important, McOuat makes the same mistake as
the others, and that it is to assume that Darwin simply followed the species
designations of his fellow naturalists. But as we have seen in previous chap-
ters, this turns out not to be true. The square peg does not fit the round
hole. Another theory is therefore needed.
Finally, there is my strategy theory published in Stamos (1996). Un-
like Ghiselin and Beatty before me, I felt guided by the principle of char-
ity to reject their view that Darwin was a species taxa though not a
species category realist. This seemed to me to attribute to Darwin a fun-
damental confusion: “No species category, no species taxa” (133, italics
added). To say there are species taxa but no species category is to make
out of the phrase “species taxa” a contradiction in terms. It didn’t seem
fair to attribute to Darwin such a confusion. Nevertheless instead of try-
ing to reconstruct Darwin’s realist though implicit definition of the
species category, I concluded that “there appears no good reason (con-
tra Beatty) to believe that he [Darwin] would have denied the reality of
the species category, although he may not have had a sufficiently clear
idea in order to define it. But, of course, it need only be remembered that
to define it would have been for Darwin to contradict and defeat what
needed to have been a lifetime strategy” (144). “Needed to have been a
lifetime strategy”? I still think so, though not for the same reasons. “May
not have had a sufficiently clear idea in order to define it”? I now sin-
cerely think otherwise. If a theorist inconsistently uses grouping and
ranking criteria, then perhaps a good case can be made for that person
 Darwin’s Strategy 167

not having a clear idea. But when a theorist, especially of Darwin’s cal-
iber, repeatedly and consistently uses the same grouping and ranking
criteria, then a strong case can and should be made for that person hav-
ing a clear idea. In the case of the species category, to say that Darwin
did not believe that the species category is real is to say that he did not
believe that there are objective and universally valid criteria for delimit-
ing species (objective and universal in the sense of the four criteria dis-
cussed in chapter 2). Since, as I believe, I have in previous chapters
reconstructed to a reasonable degree what those criteria were for Darwin,
a strategy theory, in particular one that fits the facts, is needed in order
to complete the picture.
But if all of the foregoing strategy theories are unacceptable, then
what is left? If Darwin was not only a species taxa realist but also a species
category realist, with an actual though implicit definition of the term
“species” fully in accordance with his theory of gradual evolution by nat-
ural selection, why then would he explicitly maintain the undefinability,
and by implication the unreality, of the species category? This cries out
for an explanation. Moreover, it cries out for an explanation that fits as
much of the evidence as possible and that extends to Darwin the princi-
ple of charity to the greatest possible degree.
Such an explanation cannot possibly hope to ignore the fact that the
received view in Darwin’s time, prior of course to the Origin, was that
species were created by a First Cause (God), either once or in a series of
successive creations, and that varieties were created by secondary causes
(natural laws). On this popular view the reality of species depended on
this essential distinction. If not only varieties but also species were cre-
ated by secondary causes, so that varieties are incipient species, this en-
tailed not only that species are indefinitely mutable but that species are
no more real than varieties, which in turn impugns the reality of both
species taxa and the species category. And indeed we have seen this in
chapter 1. Lyell (1832), Whewell (1837), Watson (1845b), and Wol-
laston (1860) are prominent examples of the view—what may be taken
to have been during that time the prevalent view—that either species
are fixed, in which case they are real, or they are not fixed, in which case
they are not real.
In defining both species taxa and the species category nominalisti-
cally, then, it seems to me—building on my theory first presented in
Stamos (1996, 133–137)—that Darwin was not so much trying to com-
municate his theory of evolution as he was applying to his opponents a
consistency argument:
168 DARWIN AND THE NATURE OF SPECIES

P1: You already employ common descent in grouping males and females
and larvae and adults and monsters and varieties as conspecific, in spite of
often great dissimilarity.5
P2: You believe that species are real because they are fixed by divine fiat, but
that varieties are not real because of their variability from secondary causes,
so that what are called “varieties” are merely arbitrary groupings made for
the sake of convenience.6
P3: New evidence and arguments strongly suggest, contrary to the received
view, that what are often called “varieties” are incipient species (indeed
this is the main theme of the Origin).7
C1: You therefore must conclude that, just like varieties, species are merely
arbitrary groupings made for the sake of convenience, that they are not real
either.

Given that almost all of Darwin’s contemporaries were creationists of

some sort (cf. chapter 1), this would have been a clever argument against
them. It is an example of the reductio ad absurdum form of argument, not
in the technical sense that one finds in books on formal logic, but in the in-
formal sense. Whether Darwin knew of the name of this type of argument,
I do not know, but certainly it is an intuitive form of argument that has
been around for thousands of years. As Gorovitz et al. (1979) characterize
this informal technique, “we show a position to be unacceptable by show-
ing that it leads to—or can be reduced to—something absurd or clearly
unacceptable for some other reason” (141). Having accomplished that, the
technique forces the opponent to reject one of the original premises. In the
above example, the conclusion that species are not real follows logically
from the premises. And if the premises are true, then the argument is sound.
But the conclusion that species are not real would not be acceptable to most
of Darwin’s contemporaries. Better to have species that evolve than no
species at all. The paradigm shift would be difficult, but it would be better
than to accept the conclusion that species are not real. The latter involved
a taxonomic chaos that few could stomach. Lyell, for example, stated in a
letter to Hooker (July 25, 1856) that as long as those who multiply species
“feared that a species might turn out to be a separate and independent cre-
ation, they might feel checked; but once abandon this article of faith, and
every man becomes his own infallible Pope” (Burkhardt and Smith 1990,
194 n. 6).8 Moreover there was the religious difficulty. How could species
be not real if God exists? Does God think not of species? Does not the Bible
itself, in Genesis, speak of God creating different kinds of plants and ani-
mals? The only way out is to keep God and accept evolution, with species
being in some different way real and their evolution being directed by the
 Darwin’s Strategy 169

will of God. One of the above premises, then, had to go. And the obvious
candidate for rejection would be P2. P1 was followed by all parties, so it could
not go. P3 could go if one accepted that Darwin had not made a strong case
for the fact of evolution. Since Darwin had thought he made a strong case
(and this was indeed borne out over the decade following 1859), the only
premise remaining, and the one that must be rejected and replaced, was P2.
If this was indeed Darwin’s strategy, why then, one must wonder, did
he not come out with his species concept once the Origin had done its job
and converted most of the educated world in the West from a static con-
cept of the biological world to an evolutionary one? The answer, I suggest,
if my line of reasoning is correct thus far, is because of the central element
of natural selection in his species concept, reconstructed in c hapter 5.
Although Darwin’s Origin had converted most biologists in Great Britain
(many of whom were Anglican clergymen), many other scientists, and
much of the lay public to evolutionism in the space of a decade (Ellegård
1958; Hull et al. 1978), it never converted that world to natural selection
as the main mechanism of species evolution. In fact, Darwin’s theory of
natural selection remained largely unaccepted among biologists (never mind
the general public) until well after his death, beginning in the 1930s (Ruse
1979, 228–233). Darwin, then, may have perceived it as hazardous to prof-
fer his own species concept, given that it was based on natural selection,
until natural selection itself was generally accepted. Since natural selection
was always far from being generally accepted during the remainder of his
lifetime, he never bothered to make explicit his mature species concept.
Moreover, given the already contentious nature of the definition of
“species” from within the creationist framework, Darwin may also have
thought (in a prescient way) that even if evolution by natural selection is
eventually accepted by most biologists, they still will not be able to agree
on a definition of “species.” This is all, of course, pure speculation, emi-
nently reasonable though lacking positive evidence, and I won’t try to take
it any further.
A further problem raised by my theory on Darwin’s strategy concerns
the concept of variety, what Darwin’s fellow naturalists thought about the
nature of varieties, whether they’re real or arbitrary, and what Darwin him-
self thought. Clearly much of my theory hinges on this issue, so much so
that I venture here to affirm that if my analysis in the previous chapter ul-
timately fails, then so too does my strategy theory developed in the present
chapter. For the sake of presenting the theory, however, I shall assume that
my analysis in the previous chapter is basically sound. Onward, then, with
the theory.
170 DARWIN AND THE NATURE OF SPECIES

Although there is an obvious problem with the record, since much of

Darwin’s correspondence is missing,9 and we obviously do not have record-
ings of his conversations, a particular trend is nevertheless sufficiently per-
ceivable to the attentive eye while moving up the strata of Darwin’s
correspondence. In case after case, it seems that Darwin employed a stance
of species nominalism when dealing with an outstanding naturalist whom
he thought he had a chance of converting to his views, which he then re-
placed with a stance of species realism once the conversion had become to
his mind sufficiently complete. It is also significant that with each of these
naturalists he never so much as even hinted at species nominalism prior to
his confession of evolutionism.
A prime example is Asa Gray, Darwin’s main exponent of his views in
America. Prior to his conversion by Darwin, Gray had been for a long
time a creationist and essentialist. According to Dupree (1959), Gray
“took his main arguments” from Augustin-Pyramus De Candolle’s Théorie
Élémentaire de la Botanique, first published in 1813, in which a species is
defined as “the collection of all the individuals who resemble one another
more than they resemble others; who are able, by reciprocal fecundation,
to produce fertile individuals; and who reproduce by generation, such kind
as one may by analogy suppose that all came down originally from one
single individual” (54). The species concept defined here, although it al-
lows for variation and varieties, is clearly essentialist and precludes the evo-
lution of one species from another (54). Gray first read De Candolle’s
Théorie in 1834 (40) and defended his species concept in his (Gray’s) El-
ements of Botany, published in 1836. This species concept Gray retained
and defended for many years, until he was converted to evolutionism by
Darwin (217).
In a letter to Gray (July 20, 1857), Darwin first confessed to Gray his
evolutionism, providing a very brief précis of his theory. In this letter Darwin
did not touch on the subject of whether species should now be thought of as
real or not. The most he says is “I have come to the heterodox conclusion that
there are no such things as independently created species—that species are
only strongly defined varieties” (Burkhardt and Smith 1990, 432). A little
later in another letter to Gray (September 5, 1857) Darwin provided a more
detailed précis of his theory, which was later read to the Linnean Society on
July 1, 1858, along with Wallace’s “On the Tendency of Varieties to depart
indefinitely from the Original Type” (Barrett 1977 II, 3–19), which together
were published in the Society’s Journal in early 1859. But Gray was still by
no means convinced. A little less than three months later than the previous
letter Darwin would write to Gray (November 29, 1857) the following:
 Darwin’s Strategy 171

When I was at systematic work, I know I longed to have no other diffi-

culty (great enough) than deciding whether the form was distinct enough
to deserve a name, & not to be haunted with undefined and unanswer-
able question whether it was a true species. What a jump it is from a well
marked variety, produced by natural cause, to a species produced by the
separate act of the Hand of God. . . . By the way I met the other day
Phillips, the Palæontologist, & he asked me “how do you define a
species?”—I answered “I cannot” Whereupon he said “at last I have found
out the only true definition,—‘any form which has ever had a specific
name’”! [Burkhardt and Smith 1990, 492–493]

In this letter Darwin clearly takes a stance of species nominalism. In quot-

ing, apparently favorably, the paleontologist John Phillips, one might think
that Darwin was only advocating species nominalism over time, only ver-
tical species nominalism. But what he wrote immediately prior to this pas-
sage suggests otherwise. Darwin clearly states that he thinks the term
“species” is not only “undefined” but that its definition is an “unanswerable
question.” Combined with what he says about the difficulty in deciding
whether a particular form is a species or a variety, the gist of this letter is
nominalism for both species taxa and the species category.
Gray remained unconvinced. Interestingly in a letter written over a
month after the publication of the Origin, Darwin seems to repeat his
stance of species nominalism. In a letter to Gray (December 21, 1859) he
expressed his view that intelligent readers of his book who are not natural-
ists “will drag after them those naturalists, who have too firmly fixed in
their heads that a species is an entity” (Burkhardt and Smith 1991, 440).
According to Dupree (1959, 249–269) it was not until Gray had read
Darwin’s Origin, primed for it mostly by Hooker’s correspondence through-
out 1859 and by his own earlier and recent work on the highly similar flora
of Japan and eastern North America, that Gray became in the main con-
vinced. I am not quite so sure, however. At any rate, what matters for my
strategy theory is Darwin’s perception of Gray’s conversion. Gray had fin-
ished reading Darwin’s Origin on January 1, 1860. He was duly impressed,
writing to Hooker on January 5 that “It is done in a masterly manner”
(Burkhardt et al. 1993, 16). However, in a letter to Darwin (January 23),
with regard to his first review of Darwin’s book, he wrote that it is better for
Darwin’s theory that he take a neutral position rather than announce him-
self as a “convert,” to which he immediately added “nor could I say the lat-
ter with truth” (47). Accordingly, Darwin did not include Gray in his list of
converts. In a letter to Hooker (March 3) he listed Gray as a convert only
“to some extent” (116). However, Gray’s extensive efforts on Darwin’s
172 DARWIN AND THE NATURE OF SPECIES

behalf (including three favorable reviews and public debates with Agassiz)
were not lost on Darwin. In a letter to Lyell (July 5) he wrote “Huxley,
Hooker & J. Lubbock (as I am pleased to hear) seem to have stuck up for
modification of Species like Trojans. Asa Gray, as I hear today, also goes on
fighting well” (281). Indeed in a letter to J.D. Dana (July 30) Darwin stated
that “No one person understands my views & has defended them so well as
A. Gray” (303). Similarly in a letter to Lyell, written on the same day, Dar-
win wrote “No one I think understands whole case better than Asa Gray, &
he has been fighting nobly” (306). In the letter to Dana, Darwin immedi-
ately added the disclaimer “though he does not by any means go all the way
with me.” Gray’s reservations were mainly theological. He could not accept
that variations were random with respect to the environment. Instead he
thought that favorable variations must be directed by God. Nevertheless
Gray was enough of a convert, accepting the probability of both species evo-
lution and natural selection as the main mechanism, and Darwin clearly
thought highly of his efforts. Accordingly Darwin changed his language with
respect to species reality in his correspondence with Gray. As we have seen
in chapter 1, in reply to Agassiz’s (1860b) quip “If species do not exist at all,
as the supporters of the transmutation theory maintain, how can they vary?”
(143), Darwin wrote to Gray (August 11, 1860) “How absurd that logical
quibble;—‘if species do not exist how can they vary?’ As if anyone doubted
their temporary existence” (317). In chapter 3 I attempted to elucidate Dar-
win’s meaning of the word “temporary.” At this point, however, we need not
be concerned with that matter. Existence, however temporary, is still real
existence, and Gray could hardly have exacted any other meaning from Dar-
win’s letter (cf. chapter 1 note 8). Darwin’s language here to his new con-
vert, unlike his earlier letters written when Gray was still relatively orthodox,
is clearly the language of species realism.
An equally striking though more complicated example is Darwin’s cor-
respondence with J.D. Hooker, not only for many years Darwin’s closest
friend but also his main correspondent. Darwin first confessed his evolu-
tionism to Hooker in one of his letters (January 11, 1844), in which he
stated “I am almost convinced (quite contrary to opinion I started with)
that species are not (it is like confessing a murder) immutable” (Burkhardt
and Smith 1987, 2). Roughly three years later, as indicated in one of his let-
ters to Hooker (February 8, 1847), Hooker read Darwin’s Essay of 1844
(Darwin 1909). However, it is by no means easy to determine when
Hooker became convinced. We know that by mid-1850 he was still not a
convert. In a letter to Darwin (April 6 and 7, 1850) he wrote “I remember
once dreaming that you were too prone to theoretical considerations about
 Darwin’s Strategy 173

species & unaware of certain difficulties in your way, which I thought a

more intimate acquaintance with species practically might clear up. Hence
I rejoiced at your taking up a difficult genus [barnacles] & in a manner the
best calculated to throw light on specific characters their value &c. Since
then your own theories, have possessed me, without however converting
me” (Burkhardt and Smith 1988, 328). As with Darwin’s Natura non facit
saltum claim for species, Hooker’s conversion appears to have been exceed-
ingly gradual. Peter Stevens (1997, 346) suggests that Hooker became a
convert early on, already by the time that he was traveling in India (Hooker
set sail for India on November 11, 1847, botanized for almost 3½ years
there, and returned home on March 25, 1851). Desmond and Moore
(1992, 416, 713 n. 28) suggest that Hooker’s conversion was done by July
1855 (cf. Hooker’s letter to Darwin, July 8, 1855, Burkhardt and Smith
1989, 372). I believe that Hooker’s conversion was not settled until three
years later. At any rate, what really matters for my strategy theory is Dar-
win’s perception of Hooker’s conversion. To begin, in a letter to Hooker
(December 24, 1856) that we have seen previously, Darwin wrote
I have just been comparing definitions of species, . . . It is really laugh-
able to see what different ideas are prominent in various naturalists minds,
when they speak of “species” in some resemblance is everything & descent
of little weight—in some resemblance seems to go for nothing & Cre-
ation the reigning idea—in some descent the key—in some sterility an
unfailing test, with others not worth a farthing. It all comes, I believe,
from trying to define the undefinable. [ Burkhardt and Smith 1990, 309]

Here we find Darwin clearly advocating species nominalism, at least for

the species category. As we have seen in the previous chapter as well as in
chapter 1, an almost identical passage was written at nearly the same time
for the manuscript Natural Selection, perhaps even on the same day. When
we take a closer look at the species concepts that Darwin is referring to in
both of these passages, however, the duplicitous nature of both of these
passages becomes quite apparent when viewed in the light of the preceding
chapters. What is interesting is what is missing. In Natural Selection (Stauf-
fer 1975, 95–97) Darwin discusses a number of species concepts. He dis-
cusses at length the species concept of Alphonse de Candolle, who
considered resemblance more important than descent. Darwin rejects this
using the case of primroses and cowslips. He then discusses the species con-
cept of Louis Agassiz, who allowed for perfectly similar forms to be differ-
ent species if found in widely separate localities. It is obvious how Darwin
will reject this, when in his manuscript he gets to the topic of dispersion.
174 DARWIN AND THE NATURE OF SPECIES

Finally, he discusses the species concept of Joseph Gottlieb Kölreuter, who

took sterility between forms as the sole test. Darwin will reject this when
he gets to the topic of sterility. What is missing in all of this is the kind of
species concept closest to Darwin’s own, as I reconstructed from his usage
(e.g., dogs, cattle, primroses and cowslips, pigeons, humans). Specifically,
what is missing is the kind of species concept advocated by Edward Blyth
on domestic cattle and added to by Darwin in terms of descent. Indeed, it
allows us to say how Darwin would have rated each of the criteria discussed
in the passage quoted above, as I have done in the previous chapter.
At any rate, because this nominalistic passage is in a letter rather than
a published work, Beatty (1985, 274–275) apparently took this passage in
league with the others like it, as revealing Darwin’s true beliefs, so that he
did not feel a need to attempt a strategy theory for Darwin’s apparent
species category nominalism. I strongly suggest, however, that this passage
from Darwin’s letter to Hooker should be taken no more seriously than
any comparable passage in his published works. For it seems to me that
this letter may have been occasioned by the apparent fact that Hooker was
still orthodox with respect to species. Only a month and a half prior to this
letter, after having read Darwin’s manuscript chapter on geological distri-
bution (chapter XI in Natural Selection), Hooker wrote to Darwin (No-
vember 9, 1856) “Your case is a most strong one & gives me a much higher
idea of change than I had previously entertained; &, though, as you know,
never very stubborn about unalterability of specific type, I never felt so
shaky about species before” (Burkhardt and Smith 1990, 259).
Although shaken, Hooker was still nevertheless not converted. At any
rate, what matters here is Darwin’s perception. Interesting is what Darwin
wrote about Hooker in chapter IV of Natural Selection: “Dr. Hooker ob-
jects to my whole manner of treating the present subject because varieties
are so ill defined; had he added that species were likewise ill defined, I
should have entirely agreed with him; for my belief is that both are liable
to this imputation” (Stauffer 1975, 159–160). Here Darwin clearly refers
specifically to species taxa. It is obvious that he thinks that Hooker thought
of species taxa as being real. But instead of agreeing (which we know he
would have), he impugns their reality, taking a stance of species taxa nom-
inalism. Darwin’s reference to Hooker is either to a verbal communication
or to a letter now lost.10 But the significance is not lost. The implication is
that Darwin felt that Hooker still believed to some degree in the definabil-
ity of the term “species,” in other words, the reality of the species category
in some sense traditionally conceived. According to Stauffer (1975, 94–95),
the chapter section from which this passage is taken was begun on March
 Darwin’s Strategy 175

10, 1858, and completed on May 14 of the same year. This means that it
was completed roughly 16½ months after Darwin’s letter to Hooker above
(December 24, 1856), in which he claimed that the term “species” is un-
definable. To this we may add Darwin’s later reflection written in his
autobiography (1876a):

It has sometimes been said that the success of the Origin proved “that
the subject was in the air,” or “that men’s minds were prepared for it.” I
do not think that this is strictly true, for I occasionally sounded not a few
naturalists, and never happened to come across a single one who seemed
to doubt about the permanence of species. Even Lyell and Hooker,
though they would listen with interest to me, never seemed to agree.
[123–124]

As his closest friend and confidant, Darwin was certainly in a position

to be privy to Hooker’s thoughts and feelings on the issue. Thus, although
the above fits my strategy theory, the problem concerning Hooker’s con-
version needs to be resolved. Hooker would certainly not have withheld it
from Darwin. Although Hooker may have teetered back and forth for many
years, there must have come a point when he teetered back no more but
rested on evolutionism. So was it as early as Stevens suggests, or a little later
as Desmond and Moore suggest, or was it closer to the publication of Dar-
win’s Origin? Hooker’s first public statement of his conversion to evolu-
tionism, and specifically that of the Darwin and Wallace variety, occurred
in the introductory essay of his Flora Tasmaniæ, begun in mid-December
1857 (Burkhardt and Smith 1990, 508) and completed and sent to press
in mid-March 1859 (Burkhardt and Smith 1991, 258), though its publi-
cation was delayed until mid-December 1859. According to Burkhardt and
Smith (1991), “it was in the course of writing up this essay that Hooker be-
came convinced of the value of Darwin’s theory” (xx).11 Interestingly, when
Hooker began writing this essay he had no intention of expressing any con-
version to evolutionism. Instead, as he wrote in a letter to Darwin (Decem-
ber 17–23, 1857), he intended “to confine it to a clear exposition of all the
main features of the Flora of Australia & leave all conclusion drawing to
others” (Burkhardt and Smith 1990, 508). In reply, Darwin wrote to
Hooker (December 25) “I am very sorry to hear that you do not intend to
give generalisations in your Tasmanian Introduction, but I do not believe
you will be able to resist: what is in the spirit must come out” (516). And
of course it did eventually come out, in December of 1859. But at what
point during the writing of his introductory essay did Hooker become com-
pletely convinced?
176 DARWIN AND THE NATURE OF SPECIES

The answer is apparently after reading Darwin’s manuscript on large

and small genera, which would become the second section of chapter IV in
Darwin’s Natural Selection (Stauffer 1975, 134–171), in which Darwin ar-
gued that large genera tend to have more varieties per species than small
genera, the upshot being the strong support this generalization provides
for his view that varieties are incipient species (163–164). Upon reading the
manuscript, Hooker wrote to Darwin (July 13–15, 1858) “I was at Mr
Smith’s of Combe Hurst last Sunday . . . . I there went deep into your Mss
on variable species in big & small genera . . . . After very full deliberation
I cordially concur in your view & accept it with all its consequences”
(Burkhardt and Smith 1991, 131). As if to confirm Hooker’s conversion,
Darwin would write a few months later to Hooker (October 20) “indeed I
thought, until quite lately, that my M.S. had produced no effect on you &
this has often staggered me” (174). Certain that Hooker’s conversion was
set, we then find Darwin write to his friends, starting in 1859, that “Dr.
Hooker has become almost as heterodox as you [Wallace] or I” (January
25), that “I have had the great satisfaction of converting Hooker” (Febru-
ary 12), that Hooker “is a full convert” (April 5), “goes the whole length”
(August 9), and “gives up species in grand style” (September 2), which is
to say “has completely given up species as immutable creations” (Novem-
ber 11) (Burkhardt and Smith 1991, 241, 247, 279, 324, 330, 372).12
Having believed from Hooker’s letter (July 13–15, 1858) that Hooker
was now a complete convert, Darwin no longer used the language of species
nominalism on him. For example, in a letter to Hooker (December 24,
1858) he states “what an important datum, it would be if one knew aver-
age comparative rate of specific change in Mammals & plants” (Burkhardt
and Smith 1991, 222). This was in reply to Hooker’s claim in his letter to
Darwin (December 22) that “I also still regard plant types as older things
than animal types” (219), in which he remained neutral on whether his
fossil and living plant examples, “identical to all appearances,” were specif-
ically identical. Darwin would still use in his correspondence with Hooker
during this post-conversion period the phrase “give up species,” as for ex-
ample when he wrote to Hooker (November 14, 1858) that it has long
troubled him that he interfered with Hooker’s own work by belaboring
him with so many letters and questions and ideas, and added “My only
comfort is, that without you were prepared to give up species” (199). But
by this he only meant the immutability of species.
Another striking case is Darwin’s correspondence with Charles Lyell.
As we have seen in chapter 1, Lyell, as Darwin well knew, claimed in his
Principles of Geology (Lyell 1832) that if species evolve, then they aren’t real
 Darwin’s Strategy 177

(cf. Coleman 1962). Moreover, Lyell retained this view for a considerable
amount of time. For example, in a letter to Darwin (May 1–2, 1856), in
reference to the weekend in late April 1856 during which Hooker, Huxley,
and Wollaston stayed at Darwin’s home, Lyell wrote “I hear that when you
& Hooker & Huxley & Wollaston got together you made light of all
species & grew more and more unorthodox” (Burkhardt and Smith 1990,
89). Moreover in a letter to Hooker, Lyell wrote that if naturalists believed
that species are “artificial, or mere human inventions” rather than separate
creations, then “once abandon this article of faith, and every man becomes
his own infallible Pope” (194 n. 6). Interestingly, earlier in the month of
April, the month that Hooker, Huxley, and Wollaston visited Darwin,
Lyell himself had visited Darwin for a weekend. According to Burkhardt
and Smith (1990, 91 n. 10), all the evidence, including Lyell’s own notes,
strongly suggests that it was at this time that Darwin revealed his evolu-
tionism to Lyell, including his theory of natural selection. One would es-
pecially expect, then, given Lyell’s ontological dichotomy on the species
question, that Darwin would employ against Lyell the nominalism/realism
strategy that I ascribe to him.
And this is indeed what one finds. In fact, one finds it well before Dar-
win confessed his evolutionism to Lyell (assuming Burkhardt and Smith are
correct about when that was). In a letter to Lyell (July 30, 1837) Darwin
wrote “Has your late work at shells startled you about the existence of
species? I have been attending a very little to species of birds, & the passage
of forms, do appear frightful—every thing is arbitrary; no two naturalists
agree on any fundamental idea that I can see” (Burkhardt and Smith 1986,
32). This was written during Darwin’s brief monadic theory of evolution,
which makes Darwin’s taunt even more interesting. Many years later,
shortly after Lyell learned about Darwin’s evolutionism (again, assuming
Burkhardt and Smith are correct about when that was), Lyell strongly en-
couraged Darwin to publish his views and was instrumental in arranging
the reading of the Darwin and Wallace papers to the Linnean Society on
July 1, 1858. Nevertheless, he was far from being a convert. Of course the
two had long been good friends, and Darwin respected Lyell’s feedback to
the extent that he had Lyell read drafts of chapters of the Origin before
going to press. It is significant that in a letter to Lyell (September 2, 1859),
in which Darwin briefly discussed the chapters he was having sent to Lyell,
he wrote “I have read some of Hookers Introduction to Australian Flora,
& he gives up species in grand style” (Burkhardt and Smith 1991, 330).
Darwin had to have known that Lyell would have taken this as referring to
species nominalism, a position that would be a logical consequence of
178 DARWIN AND THE NATURE OF SPECIES

Lyell’s dichotomy on species, that either species are fixed and real or not
fixed and unreal.
Lyell’s acceptance of the main theme of the Origin, however, is tricky
to nail down. But again, what really only matters here is Darwin’s percep-
tion of Lyell’s conversion. In a letter to Wallace (November 13, 1859) Dar-
win wrote “No one has read it [the ms. of the Origin] except Lyell, with
whom I have had much correspondence. Hooker thinks him a complete
convert; but he does not seem so in his letters to me” (Burkhardt and Smith
1991, 375).13 However, shortly after, the day before the official publication
of the Origin, Darwin seems to have thought that Lyell had become a con-
vert. In a letter to Lyell himself (November 23, 1859) he wrote:
I rejoice profoundly that you intend admitting doctrine of modification
in your new Edition [the next edition of Lyell’s Principles of Geology].
Nothing, I am convinced, could be more important for its success. I
honour you most sincerely:—to have maintained, in the position of a
master, one side of a question for 30 years & then deliberately give it up,
is a fact, to which I much doubt whether the records of science offer a
parallel. [391–392]

Moreover throughout the month of December Darwin proudly proclaimed

Lyell a convert to a number of his correspondents. In a letter to J.L.A. de
Quatrefages de Bréau (December 5) he wrote “Sir C. Lyell, who has been
our chief maintainer of the immutability of species, has become an entire
convert” (Burkhardt and Smith 1991, 415). In a letter to Gray (December
24) he wrote “Lyell is a complete convert” (446). In a letter to his cousin
W.D. Fox (December 25) he wrote “The Book has already made a few en-
thusiastic & first-rate converts, viz. Lyell, Hooker, Huxley, Carpenter &c.”
(449). And this continued on for quite some time. In a letter to Wallace
(May 18, 1860) Darwin wrote “Lyell keeps as firm as a tower, & this
autumn will publish on Geological History of Man, & will there declare his
conversion, which now is universally known” (Burkhardt et al. 1993, 220).
Accordingly, Darwin now uses with Lyell the language of species realism.
For example, in a letter to Lyell (December 22, 1859) he wrote “I have
heard from Sir W. Jardine: his criticisms are quite unimportant—some of
the Galapagos so-called species ought to be called varieties, which I fully
expected—Some of the sub-genera thought to be wholly endemic have
been found on Continent (not that he gives his authority) but I do not
make out that the species are the same” (Burkhardt and Smith 1991, 442).
Lyell, however, was in reality fence sitting, working ideas out in his
species notebooks (Wilson 1970), and Darwin was misled. In fact, Lyell
 Darwin’s Strategy 179

did not discuss Darwin’s theory in print until The Antiquity of Man (Lyell
1863), which deeply disturbed Darwin because of Lyell’s inconclusive
discussion on the evolution of species, to the point of rejecting the evolu-
tion of man from another species. In fact, Darwin was shocked that Lyell
did not come out for evolution, as he was convinced (“at times”) that Lyell
was in fact a complete convert (cf. Darwin’s letters to Lyell, March 6 and
12–13, 1863, and to Gray, Hooker, and Huxley in the month before;
Burkhardt et al. 1999, 207–209, 222–223, 166, 173–174, 181). In fact it
was not until the tenth edition of his Principles of Geology, issued in two
volumes in 1867–1868, that Lyell finally publicly accepted Darwin’s view
(Hull et al. 1978, 52)—although a few years earlier on the evening of
November 30, 1864, in an after-dinner speech in honor of Darwin having
been awarded the Copley Medal of the Royal Society, he gave what he
called a “confession of faith as to the Origin” (Darwin 1892, 274).
What is interesting is that throughout the years 1860, 1861, 1862, and
up until the publication of Lyell’s Antiquity, Darwin sometimes suspected
Lyell’s fence sitting but never fully caught on. We can see this especially in
the correspondence between Lyell and Darwin in 1860 (Burkhardt et al.
1993). In a letter to Hooker (March 3), he listed Lyell as a convert among
geologists (116). In July he was looking forward to Lyell publishing on the
subject of evolution (299), and then a little later he writes to Lyell of “our
side” (306). By September, however, Lyell was talking about “Most species
immutable & true to the death, as I maintained in the Principles that all
were, but a few of the whole plastic & becoming the centers of new generic
& ultimately higher groups as you maintain” (348), which Darwin took as
a hint of “the creation ‘of distinct successive types,’” such that “You cut
my own throat, & your own throat” (356), to which Lyell replied “You
need not be afraid of my starting any theory of successive creation of types”
(366). Darwin then wrote to Gray (September 26) “I can perceive in my
immense correspondence with Lyell, who objected to much at first, that he
has, perhaps unconsciously to himself converted himself very much during
last six months” (390). A day later, however, Lyell wrote to Darwin (Sep-
tember 27) saying he is “haunted with a kind of misgiving” (393) over Dar-
win’s thesis of the multiple origin of domestic dogs. Darwin then replied
(September 28) with a kind of chastisement, saying “I cannot see yet how
multiple origin of dog can be properly brought as argument for multiple
origin of man. Is not your feeling a remnant of that deeply-impressed one
on all our minds, that a species is an entity,—something quite distinct from
variety?” (397). In this passage there is a hint of the language of species
nominalism, but it would not come out fully quite yet.14 Two days later
180 DARWIN AND THE NATURE OF SPECIES

Lyell in a letter to Darwin (September 30) reiterated his theme of im-

mutable species, calling them “a thousand to one” compared to the muta-
ble kind, which he wrote “accords with my old notion that species as a rule,
or the majority of them, are immutable, & have always been so” (399).
Consequently we find Darwin (October 8) hint to Lyell that he is still a cre-
ationist (421). And then in a further letter to Lyell (December 4) Darwin
clearly returns to the language of species nominalism: “How far to lump &
split species is indeed a hopeless problem.—It must in the end, I think, be
determined by mere convenience” (512).
Turning to 1861 and 1862, Lyell had been working on the sixth edi-
tion of his Manual of Elementary Geology (which would not be published
until 1865), and in a letter to Darwin (before August 21, 1861) he ap-
pended a note that he intended to add to the Manual, which was more
conciliatory to Darwin’s views (Burkhardt et al. 1994, 239 n. 2). Lyell’s
letter (not extant) was apparently in agreement with Gray’s view that God
guides favorable mutations in the evolution of species, and Darwin in his
reply letter to Lyell (August 21) apparently took the letter and the note to
mean that Lyell was “one of the wretches” (237), meaning a believer in
evolution. Accordingly we find Darwin writing to Gray (September 17) “I
have been lately corresponding with Lyell, who, I think, adopts your idea
of the stream of variation having been led or designed” (267). In a letter to
Lyell (August 22, 1862) Darwin expresses his hope that Lyell’s Antiquity
will be out in October (Burkhardt et al. 1997, 378). Darwin, of course,
would have to wait six months to get his shock. There is little correspon-
dence between Lyell and Darwin in 1862, or until Lyell’s Antiquity ap-
peared in early February 1863, but at any rate the trend is discernible that
so long as Darwin thought Lyell was a convert there was no species nomi-
nalism talk on his part.
To complete the examination of Darwin’s inner circle, we must exam-
ine Darwin’s correspondence with T.H. Huxley. Huxley does not seem to
have come around to evolutionary thinking until shortly before the publi-
cation of the Origin. Years before this, Darwin confessed his evolutionism
in a letter to Huxley (September 2, 1854), stating “I am almost as unortho-
dox about species as the Vestiges itself, though I hope not quite so unphilo-
sophical” (Burkhardt and Smith 1989, 213). This was just after he had
read Huxley’s savage review of the tenth edition of Chambers’ Vestiges,
which Huxley later recalled as having set him strongly against evolution
(Burkhardt and Smith 1989, 213 n. 5). Adrian Desmond (1997) claims
that Huxley did an “about-face” (226) in favor of evolution shortly after the
last week of April 1856, during which Darwin had Huxley, Hooker, and
 Darwin’s Strategy 181

Wollaston stay at his home for a weekend. Although Desmond claims that
at this stay Darwin did not show his hand to his guests (I don’t know how
Desmond knows this, but at any rate Hooker and Huxley already knew
Darwin’s view), the issue of the limits of species mutability was certainly the
main topic of discussion, and Desmond argues (in typical fashion) that
Huxley converted to evolutionism a few weeks after mainly for political
reasons rather than genuinely scientific ones. As Desmond puts it,
Huxley realized he had been wrong-footed. He was turning, knowing
that his secular needs could be well served by transmutation (which could
be opposed to supernatural creation), even if it had no jot of “demonstra-
tive evidence in its favour.” Could individuals be successively modified?
After the Downe weekend he saw no reason why not. After all pigeon
breeds differ “as widely from one another as do many species.” Darwin
had taught him that. [226]15

Whatever Huxley’s main motive, the best evidence suggests that he did
not convert until the summer before the publication of the Origin. On June
3, 1859, Huxley read a paper to the Royal Institution, with the title “On
the Persistent Types of Animal Life.” At the end of this paper Huxley
(1859a) stated, with regard to the “hypothesis which supposes the species
of living beings living at any time to be the result of the gradual modifica-
tion of pre-existing species,” but without any explicit reference to Darwin,
that the “hypothesis which though unproven, and sadly damaged by some
of its supporters, is yet the only one to which physiology lends any coun-
tenance” (153). According to Burkhardt and Smith (1991), “The paper
was Huxley’s first public defence of CD’s theory” (302 n. 3). Prior to the
reading of this paper, there is no good evidence (at least that I am aware of )
that Huxley thought evolution probable. Instead, he still retained his em-
phasis on archetypes and the Cuverian purpose of classification, thinking
evolution unimportant for classification.16
At any rate, what matters, again, is Darwin’s perception of Huxley’s
conversion. While working on the Origin, Darwin queried Huxley on some
points on anatomy. Having received Huxley’s reply, Darwin in a letter
(March 13, 1859) pointed out to him that “the ideal morphologies of nat-
uralists, are on my notions, real changes in the course of time from one
part or organ into another” (Burkhardt and Smith 1991, 262). He then
added that he expected Huxley “will be just as savage with me” as he was
with Agassiz’s Essay on Classification, which Darwin himself characterized
as “utterly impracticable rubbish.” Later that year, close to the publication
of the Origin, Darwin in a letter to Wallace (November 13) wrote “If I can
182 DARWIN AND THE NATURE OF SPECIES

convert Huxley I shall be content” (375). It should be obvious, then, that

prior to the publication of the Origin Darwin did not think that Huxley was
a convert. Accordingly for my view, prior to Darwin’s perception of Hux-
ley’s conversion he used the language of species nominalism on him. For
example, in response to reading Huxley’s “Introductory Essay” in his man-
uscript later published as A Catalogue of the Collection of Fossils in the
Museum of Practical Geology, Darwin wrote in a letter to Huxley (Decem-
ber 16, 1857) “I do not understand how you can say that if only fossils
existed there would be no difficulty in practically species—as there is vari-
ation amongst fossils, as with recent, there seems to be same difficulty in
grouping” (Burkhardt and Smith 1990, 506). This example, of course, is
not as clear as one would like (and we must remember again that not all of
Darwin’s correspondence is extant), but it certainly is in the same spirit as
Darwin’s nominalistic statements with other correspondents. Moreover,
we must remember that Darwin knew that Huxley would soon be reading
his Natural Selection, in which he defines species nominalistically.
Once Darwin was sure that Huxley was a convert, however, his lan-
guage with regard to species changed. Huxley, having received his presen-
tation copy prior to the official publication date of the Origin (November
24, 1859) and having quickly read it, wrote to Darwin (November 23,
1859) “I think you have demonstrated a true cause for the production of
species & have thrown the onus probandi that species did not arise in the
way you suppose on your adversaries.” He also dedicated himself to de-
fending Darwin’s views, writing “I am sharpening up my claws and beak
in readiness” (391). Accordingly, Darwin no longer hints of species nom-
inalism in his correspondence with Huxley. The problem now, however, is
that there is barely a hint of species realism either. There is, of course, the
problem of lost letters. At any rate, shortly after reading the Origin Huxley
had asked Darwin to supply him with sources on domestic breeding for a
lecture he was preparing in defense of Darwin. In reply, Darwin in a letter
(November 27) stated that he knew of no one book and that he relied on
direct acquaintance with breeders. A couple of weeks later Darwin supplied
in a letter to Huxley (December 13) an enclosure on pigeon breeding. In
that letter he included some paraphrases of quotations from the writings of
John Matthews Eaton. Although Eaton preferred to refer to domestic
pigeons as “species,” Darwin in his accompanying letter to Huxley referred
to the “Breeds” of pigeons, remarking that if Huxley could see the draw-
ings that Darwin has, he “would have grand display of extremes of diver-
sity” (Burkhardt and Smith 1991, 428). Why would Darwin group all the
breeds of pigeons together as one species (as he did in the Origin, 20–28),
 Darwin’s Strategy 183

contrary to the breeder Eaton, if he thought that the designation of

“species” and “varieties” was arbitrary? Indeed we have seen the answer in
chapter 5. Darwin did not designate the different breeds of pigeons as dif-
ferent species—although it would have helped him in his argument with
Huxley, as we have seen in chapter 6—because the differences between
them were produced by artificial selection and therefore were not really
adaptations, which for Darwin was what distinguished real species. Dar-
win’s correspondence with Huxley throughout 1860 is preoccupied with re-
views and replies to reviews, and there are no more examples to be found
for my theory. If Darwin used the language of species realism more explic-
itly than the example I have provided above, the letters in which he did so
are, lamentably, no longer to be found. As such, then, the case of Huxley
does not strongly confirm my theory, although there is nothing there to
disconfirm it either.
So much for the above four cases of Gray, Hooker, Lyell, and Huxley.
I should add that my examination of them is not meant to hide anomalies
against my theory. None of the anomalies, however, seem to me terribly dif-
ficult. For example, as we have seen in chapter 1, in a letter to Hooker (Sep-
tember 23, 1853) Darwin wrote “Farewell, good luck to your work [on
Himalayan species], whether you make the species hold up their heads or
hang them down, as long as you don’t quite annihilate them or make them
quite permanent; it will all be nuts to me” (Burkhardt and Smith 1989,
156). This expression of species realism conflicts with my claim that
Hooker did not become a convert until mid-July 1858. However, none of
Hooker’s letters to Darwin have survived during the almost two years pre-
vious to this letter, the previous extant letter being from November 1851
(Burkhardt and Smith 1989, 627). Given the regular correspondence be-
tween Hooker and Darwin, then, it is quite possible that in a letter now lost
(or in a conversation) Hooker said something to make Darwin think that
he had become a convert. Hence, we do not find species nominalism in
the letter quoted above.
Another interesting case is a letter written by Darwin to Hooker (April
23, 1861) well after Darwin clearly and rightly considered Hooker a con-
vert. In that letter Darwin wrote “I have been much interested by Ben-
tham’s paper in N.H.R; . . . I liked the whole,—all the facts on the nature
of close & varying species. . . . I was, also, pleased at his remarks on classi-
fication, because it showed me that I wrote truly on this subject in the Ori-
gin” (Burkhardt et al. 1994, 99). According to Burkhardt et al. (1994),
“Bentham held the view, as did CD, that classification was a matter of con-
vention” (101 n. 7). Thus in this letter Darwin would seem to use the
184 DARWIN AND THE NATURE OF SPECIES

language of species nominalism to Hooker, which goes flat against my the-

ory given the dating of the letter. But if one turns to Bentham’s paper
(1861)—which was originally read before the publication of Darwin’s
Origin—the apparent anomaly quickly disappears. In the very first para-
graph Bentham points out that “the whole system of classification depends,
in the first instance, on a right understanding of what is meant by species”
(133). He then begins by considering common descent in the meaning of
“species,” and points out some difficulties with that criterion. He then con-
siders an alternative suggestion, which is to eliminate descent and employ
similarity (either a common character or set of characters). But he rejects
this, because “The species or collection of individuals thus defined, be-
comes, therefore, as arbitrary as the genus or collection of species, and re-
duces the rules of classification in the one case, as in the other, to little more
than rules of convenience.” Bentham then, believing he says “that there
exist in nature a certain number of groups of individuals, the limits to
whose powers of variation are, under present circumstances, fixed and per-
manent,” proceeds to give his own realist definition of “species” (for
botany), which is “the whole of the individual plants which resemble each
other sufficiently to make us conclude that they are all, or MAY HAVE BEEN
all, descended from a common parent” (133). Given Bentham’s species
realism, then, Darwin’s letter to Hooker cannot properly be read as an
endorsement of species nominalism.
A more problematic case is Darwin’s correspondence with H.C. Wat-
son. Although not normally thought of as part of Darwin’s inner circle,
Watson was an important correspondent of Darwin’s, and Darwin thought
very highly of him (cf. Darwin 1859, 48). Interestingly, Watson had be-
come a transmutationist apparently after reading Chambers’ Vestiges. And
although, as we have seen in chapter 1, he shortly afterward came close to
embracing species nominalism, he never went the whole way, and contin-
ued to make a distinction between book species and real species in nature.
We should suspect, then, if my strategy theory is true, that Darwin would
never have any reason to employ it against Watson. And yet we find Dar-
win write in one of his post-Origin letters to Watson (July 17, 1861) “The
difficulty to know what to call vars & what species,—hopeless—But your
suggested plan to take list of forms which some call vars & some species
? print list?” (Burkhardt et al. 1994, 207). And we cannot say this is because
Watson did not yet accept natural selection as the main mechanism of evo-
lution, because he did. In a letter to Darwin (November 21, 1859) he wrote
“Your leading idea will assuredly become recognized as an established truth
in science, i.e. ‘natural selection’” (Burkhardt and Smith 1991, 385). The
 Darwin’s Strategy 185

problem becomes greatly diminished, however, by two further pieces of

evidence. First, the letter that Darwin was replying to is lost. Second and
more importantly, the letter by Darwin is not complete, and an important
part of it has survived in a copy of that part made by Watson (Burkhardt
et al. 1994, 208 n. 5). Evidently the issue was only over “very closely allied
forms,” more specifically “how often do such allied forms (held species by
some, vars. by other botanists) live mingled in same spot,—how often in
same district, but in different stations,—and how often in different geo-
graphical districts.” Since, as we have seen in chapters 1 and 3, Darwin did
not believe that messy situations were the norm, this letter, then, should not
be read as an expression of species nominalism.
Another interesting case, and it is the final one I shall examine, is Dar-
win’s correspondence with A.R. Wallace. In what is his earliest extant letter
to Wallace (May 1, 1857), Darwin wrote “This summer will make the 20th
year (!) since I opened my first-note-book, on the question how & in what
way species & varieties differ from each other” (Burkhardt and Smith 1990,
387). In this letter there is no hint of species nominalism. And yet Darwin
did not yet think of Wallace as an evolutionist. Wallace had written a paper
on geographical distribution (Wallace 1855), in which he claimed it a law
that “Every species has come into existence coincident both in space and
time with a pre-existing closely allied species” (186, 196). In the margin of
his copy, Darwin wrote “nothing very new,” “Uses my simile of tree—It
seems all creation with him,” “I shd. state that put generation for creation &
I quite agree” (Burkhardt and Smith 1989, 522 n. 1). It is not known when
Darwin wrote these words in the margin, but Wallace’s article was favorably
referred to by Edward Blyth in a letter to Darwin of the same year (Decem-
ber 8, 1855) (Burkhardt and Smith 1989, 519, 520). In fact, two years later,
after an exchange of a number of letters with Wallace (Burkhardt and Smith
1990, 388 n. 2, 457), Darwin still thought of Wallace as a creationist. In a
letter to Wallace (December 22, 1857) he says “But you must not suppose
that your paper [Wallace 1855] has not been attended to: two very good
men, Sir C. Lyell & Mr E. Blyth at Calcutta specially called my attention to
it. Though agreeing with you on your conclusion in that paper, I believe I
go much further than you” (Burkhardt and Smith 1990, 514).17 Wallace,
unknown to Darwin, had become an evolutionist in 1845 shortly after read-
ing Chambers’ Vestiges. And of course many years later, in February of 1858,
while ill with a malarial fever in the Malay archipelago, Wallace indepen-
dently conceived of evolution by the mechanism of natural selection. In the
same month he wrote his seminal paper (Wallace 1858) and sent it to Dar-
win in the hope that Darwin would send it to Lyell, which unknown to
186 DARWIN AND THE NATURE OF SPECIES

Wallace forestalled Darwin (Kottler 1985, 368–371). Darwin received Wal-

lace’s letter (which is no longer extant) with the accompanying article on
June 18, 1858 (Burkhardt and Smith 1991, 107, cf. xvii–xviii) and was of
course shocked. What is important for my purposes is that Darwin could no
longer think of Wallace as a creationist. So why then, if my theory is correct,
did Darwin not employ his nominalism/realism strategy on Wallace? The
answer, it seems to me, is because prior to late June 1858—which is a little
after Darwin got his shock and it was quickly resolved that papers by Dar-
win and Wallace would be read to the Linnean Society on July 1; cf.
Burkhardt and Smith (1991, 121 n. 3)—Darwin was very selective about
who he revealed his evolutionism to. In fact, the list does not go much be-
yond Hooker, Lyell, Gray, Huxley, and Watson, in which cases, as we have
seen, my strategy theory applies very well. Wallace, being principally a spec-
imen collector and in Darwin’s eyes (prior to June 18, 1858) an amateur the-
orist, simply didn’t rate (cf. Endersby 2003, 397–398). Therefore we should
not be surprised to find that my strategy theory does not apply to the cor-
respondence between Darwin and Wallace.
In sum, I think it should be evident from the above examples, including
even the difficult ones, that there is statistically significant evidence support-
ing my theory that Darwin in his correspondence took a stance of species
nominalism only to replace it with a stance of species realism once the desired
conversion, in a minimal sense, had been accomplished. Contrary to Beatty’s
thesis, there is no reason to believe that Darwin did this in order to better
communicate his theory of evolution. On the contrary, it is abundantly evi-
dent with the group that would eventually become his inner circle—namely,
Hooker, Lyell, Huxley, and Gray—that they knew perfectly well what he was
talking about, even though they might have disagreed with him at the time.
Instead, Darwin’s strategy was a further part of his argumentation. In league
with his arguments about artificial selection, biogeography, classification, and
so forth, his reductio ad absurdum strategy was designed to loosen his corre-
spondents from their orthodox position on species. And it would serve the
same function in his published writings. In short, if they believe that varieties
are not real and if Darwin can demonstrate to them that varieties are incipi-
ent species, then they must conclude that species are not real either. Otherwise
they must change their concept of both varieties and species.
With all of this in mind, it is now time to widen our scope and look at
theories on the nature of concept change in scientific revolutions. The case
of Darwin on species and varieties, as now newly understood from this and
previous chapters, will prove to be an interesting case study.
 Chapter 9

Concept Change in Scientific Revolutions

The case of Darwin on the nature of species presents a major example of

concept change during a scientific revolution. It becomes even more inter-
esting if we add to it the species problem from the time of Darwin to the
present. At any rate, if we compare scientific revolutions to mass extinc-
tions, the Darwinian revolution must rank among the top five, possibly as
number one, comparable to the Permian extinction. Of course, just as with
mass extinctions, it would be a mistake to generalize from one case, no mat-
ter how important that case. Nevertheless, it is interesting and instructive
to compare the case of species, both in the case of Darwin and in the case
of biology from the time of Darwin to the present, with a number of promi-
nent theories on concept change in scientific revolutions. In particular, it
will be interesting to see how preconceived theories, drawn from other ex-
amples in science, have been applied to the case of Darwin, or how they
would apply, and how they must be revised because of the analysis pre-
sented in the previous chapters.
Perhaps a good place to begin is with the pragmatist philosopher John
Dewey’s 1909 essay on the influence of Darwin on philosophy, published
in Dewey (1910). Dewey argues that the concept of eidos, developed by the
ancient Greeks and termed species by the scholastics, was a concept of knowl-
edge right up to the time of Darwin. Species were “the sacred ark of perma-
nency” (1) in the “flux” of change (5), each of them a “fixed form and final
cause, . . . the central principle of knowledge as well as of nature” (6).

187
188 DARWIN AND THE NATURE OF SPECIES

Natural selection, on the other hand, “cut straight under this philosophy”
(11), such that the Origin constituted an “intellectual face-about” (3), a
“protest” against “the long-dominant idea” (4), and as such “precipitated a
crisis” (2).
What is especially interesting about Dewey’s (1910) argument is his
implied claim about the future of the concept of eidos/species. He says “Old
ideas give way slowly . . . . We do not solve them [‘old questions’]; we get
over them. . . . Doubtless the greatest dissolvent in contemporary thought
of old questions . . . is the one effected by the scientific revolution that
found its climax in the ‘Origin of Species’” (19).
Part of what would make Dewey’s argument attractive is the occur-
rence of overt species nominalism in Darwin’s Origin, as we have seen in
chapter 1 and which Dewey was surely aware of. But another part is con-
temporary with Dewey, namely, the surge in species nominalism in biol-
ogy around the time of his writing (cf. Morgan 1903, 33; Bessey 1908,
218; Arthur 1908, 244, 248; Cowles 1908, 267; Coulter 1908, 272).
Subsequent history, however, has not been kind to Dewey’s argument.
There are some today, of course, who advocate species nominalism, but mod-
ern biology has experienced nothing like the surge in species nominalism
around the time of Dewey (1910). Instead, since that time, species nominal-
ism has constituted a minority view compared with species monism and
species pluralism. Indeed biology since the forging of the Modern Synthesis
has experienced a bush pattern proliferation of species concepts (Stamos 2003).
To be sure, there are some today, such as Robert O’Hara (1993), who
have urged that we should try “not to solve the species problem, but rather
to get over it” (232). For one, I don’t see this happening. The species prob-
lem is not like a failed marriage. We cannot hope to get over it by trying
to forget about it and get on with our lives. For a start, biology needs a
common currency, where species constitute the “trade of ideas” in biology
(Paterson 1985, 137; cf. Maynard Smith 1975, 217–218). When we en-
counter an organism, whether old or new, we want to know “What is it?”
This is not just of theoretical but of practical importance, as in biological
control and conservation biology. “Correct identification,” as David Rosen
(1978) puts it, “provides the key to any available information about the
species, its distribution, biology, habits, and possible means of control,
which would otherwise have to be independently investigated at a consid-
erable expense of time and effort” (24).
Accordingly, most subdisciplines in biology require a species concept in
order to convey much of their information. Moreover, species are the basal
units in taxonomy, and it is difficult to imagine taxonomy without them.
 Concept Change in Scientific Revolutions 189

The problem then remains, as Cracraft (1989) puts it, that “Different species
concepts organize the world differently and conflictingly” (40). Ignoring this
problem will not solve it or make it go away. On the contrary, with the grow-
ing environmental crisis the pressure on biology to solve the species problem
has become greater than ever (Cracraft 1997, 332–337). The past 95 years,
since the time of Dewey’s writing (1910), has borne this need out.
In the case of species, then, Dewey did not get it right on the nature
of concept change during and after a revolution. Much more to the point
would seem to be Thomas Kuhn’s position, beginning with his now clas-
sic The Structure of Scientific Revolutions (1970). Against the attempt to in-
terpret his work as entailing kind nominalism, Kuhn later (1993) affirmed
that he had always held that a scientific revolution will require not simply
individuals but “both kind-concepts and their names” (316). Back in Struc-
ture (1970), Kuhn claimed that during normal science scientists learn to
group objects into “similarity sets.” During and as a result of a scientific rev-
olution, however, “some of the similarity relations change. Objects that
were grouped in the same set before are grouped in different ones after-
ward and vice versa” (200). The classic example Kuhn gives is that of the
concept of planet during and after the Copernican revolution, where the
concept did not become nominalistic but instead the extension of the con-
cept was changed to include the earth and exclude the sun. One can also
think of the concepts of sun and star, for which Giordano Bruno was
burned at the stake for maintaining that each star is a sun and our sun is a
star. Kuhn goes on to say that “Since most objects within even the altered
sets continue to be grouped together, the names of the sets are usually pre-
served. Nevertheless, the transfer of a subset is ordinarily part of a critical
change in the network of interrelations between them” (200).
I shall return to Kuhn’s last sentence above shortly. As for what he says
before it, he is pretty much on when it comes to the case of Darwin. As we
have seen, Darwin did not really attempt to eradicate the concept of species.
But he did not keep the extension of species names the same either. In-
stead, in a significant number of cases, but certainly not most, he changed
the extension of species names in accordance with his implicit evolution-
ary species concept, which was in accordance with the vera causa ideal of his
fellow naturalists for both scientific explanation and natural classification.
Moreover, in line with what Kuhn says above, Darwin generally preserved
the names of species. He did not invent new species names for already
known organisms. (This was true even in the case of the taxonomic mess
that confronted him in his work on barnacles.) Instead, in a number of
cases what were called “varieties” of one species were now called “species”
190 DARWIN AND THE NATURE OF SPECIES

by Darwin (as with primroses and cowslips), while in a number of cases

what were singularly called “species” were broken into a number of differ-
ent species (as with domestic dogs and cattle).
In all of this we can see at work what Jagdish Hattiangadi (1987) calls
the progressive and conservative forces in language change. The progressive
force is the increase in knowledge. Keeping to what is relevant to the case
of Darwin and species, kind concepts and categories are theories about
the world, and they necessarily need to change if they become untenable in
the light of new knowledge. In the case of Darwin, the progressive force was
the evidence for evolution by natural selection and all that this involved.
The conservative force, on the other hand, is the need for communication.
As Hattiangadi puts it, “So long as we try to communicate a new idea, it is
necessary to be modest in our linguistic demands—otherwise we will be
misunderstood” (172). For Darwin this meant that he could not afford to
get rid of species talk. His fellow language users were his fellow naturalists
and the public at large. Consequently he kept the term “species” and retained
in most cases, but not all, the use of that term.
Returning to Kuhn (1970), what is problematic in his claim above is
the last sentence that I said I would return to. It refers to his famous the-
ory of the “incommensurability” (incomparability) of competing para-
digms during a scientific revolution. According to Kuhn, “when such
redistributions [in the sets or kinds discussed above] occur, two men whose
discourse had previously proceeded with apparently full understanding
may suddenly find themselves responding to the same stimulus with in-
compatible descriptions and generalizations.” For Kuhn, such problems
are not merely verbal, so that “they cannot be resolved simply by stipulat-
ing the definitions of troublesome terms” (201). Kuhn would later (1993)
call incommensurability “the central innovation introduced by the book
[Structure]” (315), but to understand it better we need to fill it out with
some of Kuhn’s other key concepts. For Kuhn (1970), the history of a
science is characterized mainly by normal science—comparable, I might
add, to the period of stasis for the history of a species in the theory of
punctuated equilibria—during which puzzles are solved within the frame-
work of the existing paradigm and it is the abilities of the scientists them-
selves that are constantly tested, not the paradigm itself. If failures or
anomalies start accumulating, however, the situation might eventually
reach a threshold or crisis point, where the abilities of the scientists work-
ing under the paradigm cease to be questioned and the paradigm itself
becomes questioned. This period of crisis “generally” (67) precipitates a
scientific revolution1—comparable again to the punctuation in punctuated
 Concept Change in Scientific Revolutions 191

equilibria—where “The novel theory seems a direct response to crisis”

(75). Kuhn added to this his theory of incommensurability, which he char-
acterizes as involving (i) different kinds, definitions, and problems (148),
(ii) misunderstanding, “communication breakdown” (149, 201), even
“talking through each other (132), and (iii) “different worlds” (111) in
the sense of radically different worldviews, such that the shift in an indi-
vidual scientist from one paradigm to another during a scientific revolu-
tion, if that shift occurs at all (Kuhn subscribed to the view that quite
often the older scientists simply die off), “cannot be made a step at a time”
but is instead sudden, being much like the gestalt switch in psychology
(150), as with the famous duck/rabbit (111), or like a religious conver-
sion experience (151), involving “faith” that the new paradigm will suc-
ceed in solving problems and is on the right track (158).2 Granted, Kuhn
does qualify his meaning such that he is not to be taken as saying “no ar-
guments are relevant or that scientists cannot be persuaded to change their
minds” (152). Nevertheless, he argues (199–200), especially in Kuhn
(1977), that the underlying reasons will be largely psychological and so-
cial (139, 325), the values that guide conversion often shifting and con-
flicting (322–325), and he never calls them epistemic. Moreover, he
explicitly (1970) disavows that successive scientific revolutions approach
nearer to the truth (170–173, 205–207), although he allows for progress
in an instrumentalist sense within a paradigm (166).3 In either case, gestalt
switch or conversion, what is involved, says Kuhn, is “neural reprogram-
ming” (204). He also characterizes incommensurablity as a matter of
speaking different languages (this became his exclusive meaning in later
writings), so that translation, which for him is never more than partial, is
therefore required, which in turn serves a powerful role in conversion
(202). Although Kuhn can be found claiming that a newly established par-
adigm is not only “incompatible” with the one it replaced but “often ac-
tually incommensurable” (103), where the keyword is “often,” he can later
be found saying (Kuhn 1977) that “communication between proponents
of different theories is inevitably partial,” such that these limits “make it
difficult or, more likely, impossible for an individual to hold both theo-
ries in mind together and compare them point by point with each other
and with nature” (338). Moreover, if a scientist makes the change it is like
the change from being a native speaker of one language to being like a
native speaker of another (339). Thus, all things considered, for Kuhn “an
individual’s transfer of allegiance from theory to theory is often better
described as conversion than as choice” (338), the word “choice” being
inappropriate partly because it depends on comparison (339).
192 DARWIN AND THE NATURE OF SPECIES

Kuhn in his analysis of scientific revolutions notoriously underrepre-

sented biology (cf. Hoyningen-Huene 1993, 5). What is especially interest-
ing about the Darwinian revolution, in particular Darwin and the concept
of species, is how poorly it fits Kuhn’s model.
For a start, Frank Sulloway (1996, 16–19, 346–348) argues that the
Darwinian revolution does not seem to have been precipitated by a crisis.
Indeed, says Sulloway, “the more radical the revolution, the less it seems to
accord with Kuhn’s formula” (347). Interestingly, Kuhn (1977) tried to
make out a crisis in the decades before the Origin, vaguely referring to

fields like stratigraphy and paleontology, the geographical study of plant

and animal distribution, and the increasing success of classificatory sys-
tems which substituted morphological resemblances for Linnaeus’s par-
allelisms of function. The men who, in developing natural systems of
classification, first spoke of tendrils as “aborted” leaves or who accounted
for the different number of ovaries in closely related plant species by re-
ferring to the “adherence” in one species of organs separate in the other
were not evolutionists by any means. But without their work, Darwin’s
Origin could not have achieved either its final form or its impact on the
scientific and the lay public. [139–140]

In all of this it is hard to see a crisis. The use of metaphorical language,

such as “aborted” leaves, hardly gives Kuhn what he wants. Sulloway would
seem to be right when he argues that there was no buildup in anomalies or
failures in puzzle solving. Seemingly anomalous findings, such as new fos-
sils found in the fossil record or Darwin’s Galapagos discoveries, were rather
easily accommodated into the existing explanatory framework. Moreover,
from the time before Lamarck to the Origin, “evolutionary thinking was
considered a sign of scientific incompetence rather than a solution to wide-
spread problems in interpreting the existing data” (17). I might add that,
as we have seen in Chapter 1 in the quotation from Wollaston, special cre-
ation was considered almost an axiom, while, as we have seen in chapters
2 and 7, limited variation and reversion were considered to be laws. This
did not really begin to change until the Origin was published. Indeed, as
Sulloway further remarks, “Chambers, Darwin, and Wallace jointly created
a crisis where none had been before” (347).
Turning now to incommensurability, it might appear that Darwin
himself was aware of the problem of communication breakdown and com-
plained of it in his efforts to gain converts. For example, in a letter to David
Thomas Ansted (October 27, 1860), in particular with respect to review-
ers of the Origin, Darwin wrote “I declare that the majority of readers seem
 Concept Change in Scientific Revolutions 193

utterly incapable of comprehending my long argument. . . . I am often in

despair in making the generality of naturalists even comprehend me. Intel-
ligent men who are not naturalists and have not a bigoted idea of the term
species, show more clearness of mind” (Burkhardt et al. 1993, 446). And
yet, as Sulloway (1996) argues,

Darwin himself seems to have considered evolution as fully commensurable

with creationism. His Origin of Species was “one long argument” compar-
ing how well the available biological evidence could be interpreted by cre-
ationism and evolution. He sought to demonstrate, point by point, that
rational criteria consistently favored the evolutionary alternative. . . .
Darwin was so successful in showing how commensurable evolution and
creationism were that he forced his readers to make a direct choice be-
tween the two theories. Most reasonable people had to admit that evolu-
tion was the superior theory. After Darwin’s careful dissection of the
Creator’s handiwork, the Creator began to look so mindless that many
creationists wisely sought to move Him into the biological background as
a “first cause.” [349–350]

I would add to this that Darwin’s “one long argument” is an excellent

example of what has come to be known in philosophy of science as infer-
ence to the best explanation. This is a model of scientific explanation that in
many ways is superior to the traditional deductive-nomological model, one
reason being that in biology explanation is not, if ever, in terms of laws of
nature (cf. Mayr 1982, 37; Stamos 2006). At any rate, at the very heart of
inference to the best explanation is contrastive explanation. To use a simple
example given by Peter Lipton (1990), it is not enough to explain that Kate
won the essay prize because her essay was excellent. A full explanation
would have to include why it was better than the essay of the other final-
ist, Frank. As Lipton puts it, “One reason that explaining a contrast is some-
times harder than explaining the fact alone is that explaining a contrast
requires giving causal information that distinguishes the fact from the foil,
and information that we accept as an explanation of the fact alone may not
do this” (252).
Sulloway goes on to argue that firstborns, those who were actually the
oldest among their siblings or functionally took their role, would be more
likely to claim incommensurability, since firstborns tend to be more conser-
vative and less open to new ideas than laterborns, who tend to be more flex-
ible, even rebels. As Sulloway (1996) puts it, “Psychologically speaking,
incommensurability is a problem for inflexible people, not for those who
are open to experience” (349). This is of a piece with Sulloway’s enormous
194 DARWIN AND THE NATURE OF SPECIES

statistical analysis, 26 years in the making, based on the biographies of sci-

entists involved in scientific revolutions, in particular the Copernican,
Newtonian, Darwinian, and Einsteinian revolutions. Sulloway found, re-
markably, that those scientists who defended the traditional paradigm
tended to be firstborns, while those who defended the emerging new para-
digm tended to be laterborns. Rationality, thus, says Sulloway, is “typically
subject to a threshold effect” (341), laterborns having a lower threshold and
therefore tending more quickly to see the better theory for what it is, and
firstborns having a higher threshold. Indeed as empirical evidence and argu-
ment accumulate, and the revolution comes to a close, birth order differences
tend to diminish (337), the firstborns either dying in their stubbornness or
converting to the new science (34–36).4
What is magnificent about Sulloway’s (1996) work is that he literally
spares no effort, not only in developing theories but in subjecting them to
rigorous testing. This latter part is terribly missing in the humanities. For
example, feminists and sociologists seem averse to the point of hostility in
subjecting their theories to tests, whether to the statistics of rape (which
point away from culture and environment being the sole cause), to the sta-
tistics of homosexuality (which in males increasingly points to a partly evo-
lutionary/genetic cause), or to the statistics of behavioral differences between
men and women (which fit an increasingly evolutionary view). Marxists too
seem bent on believing, no matter how much evidence to the contrary, that
human nature is completely plastic. Social constructionist history of science
(which I shall consider in the next chapter) fares no better.
At any rate, returning to the topic of incommensurability, a major part
of the problem created by Kuhn stems from his adherence to the tradi-
tional theory of meaning (cf. Kuhn 1970, 198), according to which the in-
tension (concept, definition, description, sense, connotation) of a term
determines its extension (reference, denotation), either by a conjunction
of defining predicates (e.g., Mill, Frege, Russell) or by a cluster of defining
predicates (e.g., Wittgenstein, Searle). This was a view that Kuhn main-
tained throughout his career and refused to change (cf. Kuhn 1993, 316).
On this view of meaning, given that a revolutionary theory (the new para-
digm threatening to displace the old) typically employs radically different
meanings for at least some of the key terms, it would happen that scientists
on opposite sides of the revolutionary divide would be referring to differ-
ent kinds of things even though they would be using the same terms. Con-
sequently they would be having a communication breakdown. Moreover it
would be impossible to say that knowledge of kinds progressed over time,
from the traditional paradigm to the revolutionary paradigm that displaced
 Concept Change in Scientific Revolutions 195

it. Instead there would simply be different theories of kinds, with an overlap
in terminology but not an overlap in reference.
Many philosophers (cf. LaPorte 2004, 117–118 for references) have
found that the incommensurability thesis of Kuhn is undermined by the
new causal theory of reference. This theory, connected principally with
Saul Kripke, Hilary Putnam, and Keith Donnellan (cf. the Introduction
and papers in Schwartz 1977), is arguably superior to the traditional the-
ory as both a theory of meaning for proper names and for natural kinds.
Beginning with proper names, for which the causal theory was originally
designed, it is fundamental to the traditional theory of meaning that a set
of defining predicates determines the reference of, say, “Aristotle,” such as
“son of Nicomachus and Phaestis,” “student of Plato,” and “teacher of
Alexander the Great.” However, if these defining predicates (which the
proper name represents) turn out to be historically false, then the name
“Aristotle” has no historical reference and we are talking about no one. This
is a paradoxical result that the causal theory avoids. On this theory, what
fixes the reference of the name “Aristotle” is an initial baptism, an osten-
sion, typically accompanied by a description that aids in the ostension, not
in the sense of listing attributes but in fixing the reference, such as, “This
baby is ‘Aristotle.’” Consequently, references to Aristotle by later genera-
tions can correctly refer to the historical person, Aristotle, even if every-
thing they believe about him happens to be historically false, just so long
as their use of the name “Aristotle” is part of a causal/historical chain lead-
ing back to the initial baptism. For the creators of this new theory of mean-
ing, the reference of natural kind terms (e.g., “gold,” “water,” and “tiger”)
are fixed in basically the same way, by referring to a sample of the kind and
typically using a reference-fixing description.5
The way in which this theory of meaning undermines incommensura-
bility is that scientists on opposite sides of the revolutionary divide can be
talking about the very same kinds even though their theories about those
kinds are radically different. Their meaning, in the sense of reference, can
be exactly the same. Realism about kinds can therefore be maintained.6
Moreover, progress in knowledge about the kinds can be accomplished
because the reference between theories remains the same.7
Indeed it is this theory of meaning that forms the background to Beatty’s
(1985) theory about Darwin on “species.”8 Again, as we have seen in previous
chapters, according to Beatty (1985) Darwin “tried to get beyond definitions
to referents” (269), “he used the term [‘species’] in accordance with examples
of its referential use by members of his naturalist community” (277), “By try-
ing to talk about the same things that his contemporaries were talking
196 DARWIN AND THE NATURE OF SPECIES

about, he hoped that his language would conform satisfactorily enough for
him to communicate his position to them” (266). Here we can see Beatty
claim, in other words, that Darwin surmounted the incommensurability prob-
lem, that he bridged the revolutionary divide and facilitated communication,
by retaining the reference to species made by his fellow naturalists. And indeed
Philip Kitcher (1993), explicitly following Beatty (1985)—ironically whose
work he himself influenced (cf. note 8 above) with his (Kitcher 1978) discus-
sion on and modification of the causal theory of reference—makes explicit
what Beatty left more or less implicit: “the term [‘species’] retained its old ref-
erent (the set of species taxa after Darwin is the same). Similarly, there is con-
tinuity of reference of the term ‘homology’ (and its correlative, ‘analogy’)”
(32). Moreover, “Partly because they were talking about the same groups of
species and the same traits as homologous, partly because the loci of disagree-
ment were so readily identifiable, Darwin and his critics had not the slightest
difficulty in communicating their ideas to one another” (32 n. 46).
Interestingly, the case of Darwin on species actually fits much better,
though not perfectly, Kitcher’s own modification of the causal theory of ref-
erence, which has at its core what he calls “reference potential.” As Kitcher
(1993) puts it, the reference potential of a scientific term is “the com-
pendium of modes of reference for a term (type) . . . . reference potentials
are typically heterogeneous: that the linguistic community to which a sci-
entist belongs allows a number of distinct ways of fixing the reference of to-
kens of terms” (78). Kitcher (1978) is where Kitcher first developed this
theory, and his focus there is on the concept of phlogiston. Although the
reference for this term was first fixed in a particular way, namely, the ref-
erence-fixing description (possibly by Stahl) that phlogiston is “that which
is emitted in all cases of combustion,” the reference later expanded, in a
way that all subscribers to the theory of phlogiston agreed upon (540), to
include also “dephlogisticated air,” what they called the gas obtained by
heating the red calx of mercury, and which Lavoisier later came to call “oxy-
gen.” This meant that even though “phlogiston” failed in fact to refer, its
related concept “dephlogisticated air” did not. Equally important, it meant
that the event types responsible for each token of the word “phlogiston” by
the community that subscribed to phlogiston theory became a set of two
types, or reference-fixing descriptions. (I am ignoring a third type, “inflam-
mable air,” for the sake of simplicity.) In this way, according to Kitcher, the
reference potential of a term shifts, sometimes expanding in terms of the set
of event types or reference-fixing descriptions, sometimes contracting. This
allows us to avoid “conceptual relativism” (535), to “allow for radical con-
ceptual revision without conceptual discontinuity” (544), and to “clarify the
 Concept Change in Scientific Revolutions 197

idea that theoretical concepts must absorb theoretical hypotheses and so

enhance our understanding of conceptual change in science” (543).
What is interesting about this theory is how well it applies to Darwin,
in the case of “species” but even more importantly in the case of “variety.”
Although it is virtually impossible (as far as I know) to determine who began
the first reference-fixing description for the term “variety,” by the time of
Darwin the term clearly had a heterogeneous reference potential, as we have
seen in Chapter 7. The difference between the case of “phlogiston” and the
case of “variety” is that not all naturalists at the time of Darwin, or from
the time of Linnaeus to Darwin, agreed upon the total set of reference-
fixing descriptions, or token-initiating events, for the term “variety.”
(Indeed the reference of the term “variety” was no more fixed than the ref-
erence for the term “species.”) And yet this does not seem to matter. The im-
portant fact is that the heterogeneous reference potential was there. What
Darwin did, again as we have seen in chapter 7, was attempt to contract that
reference potential, limiting the use of the term to only what would qualify
as an “incipient species,” which had to be a group of organisms character-
ized by a heritable trait that either is being evolved or could be evolved by
natural selection into an adaptation. The heterogeneous set of reference-
fixing potentials for the term “variety” was therefore in an important
respect similar to the set for the term “phlogiston,” in that, much like
Lavoisier with what would later be called “oxygen,” Darwin established con-
tinuity with one item in that set, namely, those varieties called as such by his
fellow naturalists that qualified as incipient species. This also explains why
and how Darwin attempted to shift the reference potential of the term
“species.” Even though his fellow naturalists were mistaken in their theory-
laden heterogeneous set of reference-fixing descriptions, many of the entities
they referred to as “species” were in fact species in accordance with Darwin’s
new reference-fixing description, namely, his implicit species concept defined
near the end of chapter 6. Of course, unlike Lavoisier and the reference of the
term “oxygen,” the shift in the reference potential of the term “species” that
Darwin tried to effect did not become fixed. Instead, the term is still to this
very day shifting between different reference-fixing descriptions, with no
agreement or consensus among the expert users of the term (cf. Stamos 2003),
even as to whether species are evolutionary (89, 323). The situation in mod-
ern biology is thus very different than the situation in modern chemistry.
Kitcher, of course, missed all of this, apparently for the simple reason that
he followed Beatty (1985) in thinking that Darwin retained the reference of
the term “species” of his fellow naturalists. He consequently thought that there
was no incommensurability problem. For others who subscribe to the causal
198 DARWIN AND THE NATURE OF SPECIES

theory of reference, however, and also to Beatty’s theory on Darwin, the mat-
ter is not so simple. Joseph LaPorte (2004), for example, accepts a modified
version of the causal theory (5–7), and he also explicitly accepts Beatty’s (1985)
claim that Darwin retained the reference of the word “species” of his fellow
naturalists (192 n. 13). But unlike Kitcher, LaPorte argues that when applied
to the case of the Darwinian revolution Kuhn was partly right about incom-
mensurability. He refers to evidence such as Darwin’s letter to Ansted quoted
earlier in this chapter, in which Darwin seems to complain about communi-
cation breakdown, as well as Wollaston’s (1860) claim in his review of the
Origin that “it is no sign of metaphysical clearness when our author [Darwin]
refuses to acknowledge any kind of difference between ‘genera,’ ‘species,’ and
‘varieties,’ except one of degree” (133), in which Wollaston seems to reject
Darwin’s view as being too paradoxical. Thus, for LaPorte, Kuhn’s incom-
mensurability thesis when applied to the concept of species in the Darwinian
revolution “turns out to enjoy more merit than a first glance suggests” (122).
For LaPorte, Kuhn is partly right not because of the traditional theory of
meaning but because the reference-fixing descriptions of Darwin’s fellow nat-
uralists were not only individually vague but jointly conflicting. For example,
for one (William Hopkins) evolution destroys the reality of species, while for
another (Henry Fawcett) evolution means that the whole tree of life is one or
a few species (123–126). What Darwin did, then, according to LaPorte, is
retain their reference for “species” but replace their vague and conflicting
reference-fixing descriptions with a new one that was clearer and more knowl-
edgeable about what they were in fact referring to. Indeed, “Darwin under-
stood more than did speakers before him about both what he called ‘species’
and what speakers before him called ‘species’” (129), so that his new use of
“species” constituted progress over the previous uses of “species,” progress in
the sense of “replacing vague statements that are neither clearly true nor clearly
false with straightforwardly true statements” (131).
Not surprisingly LaPorte is quite vague on the content of Darwin’s
“newly refined use of ‘species’” (123), except to say that “For Darwin the
supposition that there are lots of ‘species’ is retained, and the supposition
that all blood relatives of a lineage’s original population belong to that species
is discarded” (125). Of course another part of Darwin’s “newly refined use
of ‘species’” is that statements such as “New species arise by evolution” are
true (131).
Failing to get even close to what Darwin really meant by “species,”
however, is not the main problem with LaPorte’s account, although it is a
big one. Much deeper is that he follows Beatty (1985) on Darwin’s refer-
ential use of the word “species.” The problem is enormous because, as we
 Concept Change in Scientific Revolutions 199

have seen mainly in chapter 5, Beatty’s thesis is highly inaccurate. Indeed,

it would appear that Beatty took the causal theory of reference and at-
tempted to impose it on the evidence, rather than test the theory against
the evidence. Remarkably this quickly and easily became the received view.
Rather than look at Beatty’s claim as a theory that needed to be tested
against the evidence, it was simply accepted as a fact. At any rate, Darwin,
as we have seen, did not in fact simply retain the reference of the term
“species” of his fellow naturalists. Instead, he quite often changed that ref-
erence. Because Beatty missed this, he failed to see what Darwin was really
doing. Those who followed Beatty not surprisingly failed to see it too. In
not seeing that Darwin did not always follow the referential use of the term
“species” of his fellow naturalists, they saw no reason to inquire any further.
But once one sees that Darwin often changed the reference of the term
“species” of his fellow naturalists, one is then naturally led to inquire as to
why. That question led to the previous chapters culminating in the recon-
struction of Darwin’s species concept in chapter 6, the implicit but highly
developed species concept that Darwin had in the back of his mind every
single time he changed the reference of the term “species.” All of this re-
quires us to look anew at the incommensurability problem as applied to
the Darwinian revolution.
LaPorte indeed tries to retain some of Kuhn’s incommensurability the-
sis as applied to Darwin and the Darwinian revolution, but because he fol-
lows Beatty, he has failed to see what the real situation was, and consequently
what the real problem was. For a start, it is surely significant that Darwin
held back his true meaning of “species” and defined the term instead nom-
inalistically (for both category and taxa). This certainly adds an interesting
twist to communication breakdown. Indeed, the breakdown on this topic
has continued to the present day! But this is not to say that Darwin’s fellow
naturalists would not have understood what he meant by “species” had he
taken the time to define it (assuming for the sake of argument that my re-
construction in previous chapters is essentially correct). The problem was not
simply one of language or meaning or communication. Nor was it that the
species concept of Darwin’s fellow naturalists was “theory laden,” as Beatty
(1985, 271, 279) has claimed. That is to put it much too mildly. The prob-
lem was not theories but deep-seated beliefs. Darwin, in the letter to Ansted
quoted above, refers to the “bigoted idea of the term species” (italics mine).
This compares favorably (in spite of my analysis in chapter 8) with what
Darwin wrote in a letter to Asa Gray (December 21, 1859) with regard to
an American reprint of the Origin, that “I have made up my mind to be
well abused; but I think it of importance that my notions shd. be read by
200 DARWIN AND THE NATURE OF SPECIES

intelligent men, accustomed to scientific argument though not naturalists.

It may seem absurd but I think such men will drag after them those natu-
ralists, who have too firmly fixed in their heads that a species is an entity”
(Burkhardt and Smith 1991, 440, latter italics mine). The problem was not
meaning or theories but beliefs, deep-seated beliefs shared by naturalists but
not by other intelligent men, namely, the “axiom” of special creation as we
have seen Wollaston call it in chapter 1, the “assumption” or “faith in cer-
tain impassable barrier between existent species” as we have seen Watson
call it also in chapter 1, the belief in the sterility barrier as we have seen in
chapter 6, and the belief in the laws of limited variation and reversion as we
have seen in chapters 2 and 7.
These and other beliefs did not pose a barrier to understanding and
therefore to choice. For example, not everyone believed in the sterility bar-
rier, and yet those who did believe in it knew what those who did not were
talking about and vice versa. Similarly, lumpers knew what splitters were
talking about and vice versa. In each case the problem was not understand-
ing but agreement.
The real problem, then, was not one of content or meaning, but of
prejudice and dogma. What further added to this were the religious and
moral dimensions. Adam Sedgwick, for example, in a letter to Darwin
(November 24, 1859), was surely not speaking only for himself when he
wrote “Passages in your book . . . greatly shocked my moral taste”
(Burkhardt and Smith 1991, 397). In line with this, Darwin was surely not
simply joking when he referred to the conversion to his views as “perver-
sion” (e.g., Burkhardt and Smith 1991, 241, 347), a usage originally given
to him by Lyell (349).
In the case of Darwin, then, we have every reason to believe that his fel-
low naturalists would have sufficiently understood his real species concept
had he defined it, and therefore would have been able to make compar-
isons between his concept and other species concepts. This is because every
element in Darwin’s species concept would have been perfectly familiar to
them. As Loren Eiseley (1958) in a different context put it, “The complex
of ideas which later went to make up Darwinism was widely enough dif-
fused in the eighteenth century” (119). Moreover, no mental acrobatics
would have been needed to understand Darwin’s unique combination of
the elements, so very much unlike Einstein’s concept of curved space (one
of Kuhn’s favorite examples of conceptual incommensurability; cf. Kuhn
1970, 149). For a start, the importance of adaptations was shared by Dar-
win’s fellow naturalists, as was the importance of common descent (which
they limited to within a species). The horizontal dimension would have
 Concept Change in Scientific Revolutions 201

been understood from Darwin’s language analogy. The process that Dar-
win claimed makes species, namely, natural selection, would have been well
understood because they knew of artificial selection and it was well under-
stood. They also would have appreciated the fact that Darwin’s species con-
cept was fully in accord with their vera causa ideal for both scientific
explanation and classification, as we have seen in chapter 5. Moreover,
many of the species of Darwin’s fellow naturalists coincided in terms of
reference (restricted of course to contemporary organisms, whether living
or fossil) with the species determined by Darwin’s species concept, since the
species of the former would have had to have at least one unique adapta-
tion (as they often did) if there was to be overlap. And of course no one
doubted that species (their members) have adaptations. That Darwin did
not provide his definition of “species,” then, was not because of a concern
for communication. Indeed this was not much of a problem in itself and
Darwin knew it, which is revealed in the fact that he was not afraid to
change the reference of what his fellow naturalists called “species,” again as
we have seen in chapter 5. Instead Darwin’s concern was for conversion,
and it was because of this concern that he employed not only the nature
of his “one long argument” in the Origin but also the nominalistic strategy
revealed in chapter 8.
The subject of conversion presents a further problem for Kuhn’s
model. Kuhn’s claim about conversion is a psychological one, that conver-
sion in an individual scientist during a revolution is saltational not gradual.
This follows from his connection of conversion with a gestalt switch, irre-
gardless of whether he further thought of conversion in the religious sense
(cf. note 2 above). This is a theory that is testable. The evidence that con-
stitutes the testing is, of course, indirect, both because we are talking about
the minds of scientists and because we are dealing with history, where most
of the principals are dead. In many ways the situation is analogous to the
theory of punctuated equilibria. In both cases (confining Kuhn’s thesis to
the Darwinian revolution) the organisms in question are all dead and we are
dealing exclusively with historical records, so that the evidence is indirect.
Moreover, the debate is between saltation versus gradualism (of course with
paleontology the “saltation” is in terms of geological time, which accord-
ing to punctuated equilibria is 5,000–50,000 years). Finally, the issue is
one of frequency (cf. Stamos 2003, 222–223). Interestingly, in the case of
the theory of punctuated equilibria the evidence does not seem to be in its
favor (cf. Erwin and Anstey 1995). In the case of Kuhn’s theory of conver-
sion when applied to the Darwinian revolution, the evidence against seems
even stronger.
202 DARWIN AND THE NATURE OF SPECIES

To begin with, there is certainly indirect evidence that supports Kuhn’s

theory. For example, in his autobiography, when looking back at the rev-
olution he inaugurated, Darwin (1876a) appears to view conversion as a
kind of gestalt switch. In an often-quoted passage he says “What I believe
was strictly true is that innumerable well-observed facts were stored in the
minds of naturalists, ready to take their proper places as soon as any theory
which would receive them was sufficiently explained” (124).
When we look more closely at the Darwinian revolution, however, we
find time after time that among key cases the “conversion”—as we have
seen in chapter 8 the term was used quite frequently by Darwin and his
contemporaries—was not saltational but gradual, no matter whether the
conversion was full or partial. We have seen this in chapter 8 in the case of
Lyell, in the case of Gray, in the case of Huxley (which was even longer if
Di Gregorio 1982 is correct), and in the case of Hooker. Indeed Hooker’s
conversion, as we have seen, was extremely gradual, and this is borne out
by Henry Fawcett (1860) in his review of the Origin, in which he quotes
from a speech Hooker gave at a then recent meeting of the British Associ-
ation: “I knew of this theory [Darwin’s] fifteen years ago; I was then entirely
opposed to it; I argued against it again and again; but since then I have de-
voted myself unremittingly to natural history; in its pursuit I have travelled
around the world. Facts in this science which before were inexplicable to
me became one by one explained by this theory, and conviction has been
thus gradually forced upon an unwilling convert” (92).
Moreover, Darwin’s own conversion was not saltational but gradual.
The evidence for this is meticulously examined by Sulloway (1982), in which
he concludes that Darwin’s “conversion to the theory of evolution did not
spring full-blown as the result of his voyage, but emerged gradually in inti-
mate cooperation with the numerous systematists who helped to correct
many of his voyage misclassifications” (388). What is not examined by Sul-
loway is the evidence from Darwin’s own recollections on the matter years
later, repeated again and again, recollections for which there is no good rea-
son for doubt or disbelief. For example, in a letter to Leonard Jenyns (Oc-
tober 12, 1844) he wrote “The general conclusion at which I have slowly
been driven from a directly opposite conviction is that species are mutable
& that allied species are co-descendants of common stocks” (Burkhardt and
Smith 1987, 67). Around the time of the publication of the Origin Darwin
emphasized this even more. For example, in a letter to James Dwight Dana
(November 11, 1859) he wrote “It took me many long years before I wholly
gave up the common view of the separate creation of each species”
(Burkhardt and Smith 1991, 368). In a letter to John Stevens Henslow
 Concept Change in Scientific Revolutions 203

(November 11) he calls it a “process” (370), while in a letter to Jenyns

(November 13) he says “It took long years to convert me” (374). In a letter
to John Gwyn Jeffreys (December 29) the language is identical: “It took me
long years before I gave up the creation view of each species” (460). In a
letter to Charles Bunbury written the next year (February 9, 1860) he
stressed the point further, stating “I changed so slowly myself that I am in-
deed surprised at anyone becoming a convert” (Burkhardt et al. 1993, 76).
Moreover, late in his life Darwin’s memory of his conversion remained the
same. For example, in a letter to Dr. Otto Zacharias (February 24, 1877) he
wrote “in July, 1837, I opened a note-book to record any facts which might
bear on the question. But I did not become convinced that species were mu-
table until, I think, two or three years had elapsed” (Darwin 1892, 175).9
Darwin’s perception of the conversion of others was that it was also
often gradual. In a letter to Cuthbert Collingwood (March 14, 1861), for
example, he expresses “surprise” that he has “been successful in converting
some few eminent Botanists, Zoologists, & Geologists,” to which he adds
“In several cases the conversion has been very slow & that is the only sort
of conversion which I respect” (Burkhardt et al. 1994, 53). Again, in a let-
ter to Alphonse de Candolle (January 14, 1863), he wrote “I remember
well how many years it cost me to go round from old beliefs. It is encour-
aging to me to observe that everyone who has gone an inch with me; after
a period goes a few more inches or even feet” (Burkhardt et al. 1999,
39–40). And in a letter to Dana (February 20, 1863) he repeats again that
he values only gradual conversion: “Indeed I should not much value any
sudden conversion; for I remember well how many long years I fought
against my present belief” (155).
Perhaps a much better comparison for personal conversion in a scien-
tific revolution, then, at least as indicated by the Darwin revolution, is not
religious conversion but religious apostasy, the loss of faith, which tends to
be gradual (cf. the collection of autobiographical essays in Babinski 1995,
one of which is by yours truly). Often in such cases one does not simply
give up one’s former worldview; one also replaces it with another, a natu-
ralistic one, a worldview that one thinks is both best and true. This is very
different than the typical conversion from one religion to another. The
latter is typically saltational, the former typically gradual. Of course in
many cases the transition or conversion away from religious belief is for
non-intellectual reasons: psychological, social, and in many cases even
economic. But not all. Many cases are primarily for reasons of logic and
evidence. Darwin’s own loss of faith is a case in point (cf. Darwin 1876a,
85–96), and it was anything but saltational. Significantly, it seems to
204 DARWIN AND THE NATURE OF SPECIES

mirror his own conversion (and that of many others) to evolution, as being
mainly a matter of the slow sifting of evidence and argument in spite of
deeply-entrenched prior beliefs.
The case of religious conversion, on the other hand, where the conver-
sion is from one religion to another, usually does not seem at all to fit this
picture. The conversions of Pascal and Paul are but two of many cases in
point. Here the reasons seem to be deeply psychological and emotional,
not evidential and rational, which would explain why they tend to be salta-
tional. In the case of Paul, for example, named Saul at the time, his con-
version appears to have been the result of a buildup of guilt resulting from
his zealous persecution of Christians, a buildup that finally peaked and
erupted into a mental breakdown while on the road to Damascus, accom-
panied by hallucinations and temporary blindness. This is a gestalt switch
par excellence, like a duck/rabbit or Necker cube, but it bears little if any
similarity to the scientific conversions of Darwin, Hooker, Huxley, Gray,
or Lyell, or of many others for that matter.
Indeed, what is of further interest here is that it would seem that it was
not really the evidence per se that Darwin examined in his early years that
converted him to evolution, but his genius in reworking and interpreting
that evidence. On this view it would be his further genius, then, in develop-
ing his argument over the next 20 years, which played the main role in the
conversion of other naturalists such as Hooker and Huxley. Sulloway (1982)
points to “the fact that Darwin subsequently published this [early] evolution-
ary evidence in his Journal of Researches (1839) and in the Zoology of the Voy-
age of the H.M.S. Beagle (1841), fully two decades before the Origin of Species
(1859). And yet not one naturalist appears to have been converted to a be-
lief in evolution by these earlier works” (389). The real source for Darwin’s
conversion, then, claims Sulloway, was “Darwin himself,” his “gifted indi-
vidualism,” and “not the evidence per se, that ultimately imposed the un-
orthodox interpretations that led him to embrace the theory of evolution”
(389). Sulloway argues further that Darwin’s genius did not stop there but
continued in his later work, in which “he continued to do highly original
work and to make important discoveries missed by his peers, in almost every
branch of natural history to which he turned” (389). Interestingly, Sul-
loway’s view (which I share), with respect to the 20 year period leading up
to the Origin, seems to have been corroborated by one of Darwin’s contem-
poraries, namely, H.C. Watson, who in a letter to Darwin (November 21,
1859) remarked “How could Sir C. Lyell, for instance, for thirty years read,
write, & think, on the subject of species & their succession, & yet constantly
look down the wrong road!” (Burkhardt and Smith 1991, 385).
 Concept Change in Scientific Revolutions 205

What is interesting in all of this is the role of logic and evidence and
individualism in concept change in a scientific revolution, something seri-
ously ignored or at least underemphasized by Kuhn and by all those who
follow him. But it is also often missed by those who try to reply to Kuhn
by taking the route of the causal theory of reference. We have seen in the
case of Beatty (1985) especially, the attempt to apply this theory to the case
of Darwin on species. He took a modern philosophical theory on meaning
and concept change and attempted to force it on the evidence, like trying
to force a square peg into a round hole. And surprisingly everyone interested
in the subject, including professional Darwin scholars who should have
known better, followed him as if he had been discovering new facts. As re-
markable as this all is, the fact remains that what Beatty did was revision-
ist history in the worst sense of the term. And it is that kind of history,
though done in a very different way, that brings us to the next chapter.
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 Chapter 10

Darwin and the New Historiography

In this chapter I wish to take the analysis presented in the first eight chap-
ters and use it to deal in a particular way with a small number of represen-
tatives of what may rightly be called in professional history of science “the
new historiography.” This is definitely not meant to include all or possibly
even most professional historians of science flourishing in the past few
decades, including those who focus on Darwin and topics Darwinian. It
does not include, for example, those committed to what has been called
the “Darwin industry,” namely, the enormous amount of labor that has
been put into publishing, and not just publishing but providing minute
background details to, Darwin’s unpublished manuscripts (Stauffer 1975),
his articles (Barrett 1977), his notebooks (Barrett et al. 1987), his margin-
alia (Di Gregario and Gill 1990), and—the biggest job of all—his massive
correspondence (which began with Burkhardt and Smith 1985 and which
still, at the time of this writing, has the last seventeen years of Darwin’s life
to go). The Darwin industry also includes numerous interpretative essays
largely based on these writings (e.g., Kohn 1985a), as well as the very fine
biography of Darwin by Janet Browne (1995, 2002). Historians in this
field, as David Kohn put it in his Introduction to Kohn (1985a), feel a
“pressing need to place Darwin in the context of Victorian science,” but
they also share “the implicit premise . . . that Charles Darwin was a thinker
of profound intellect and influence” (1).

207
208 DARWIN AND THE NATURE OF SPECIES

Instead, it is the other end of the spectrum of modern history of sci-

ence that I am mostly concerned with here, and it is what I mean by “the
new historiography.” It is usually called “externalist history” or “contextual
history of science.” In his biography of Huxley, Adrian Desmond (1997)
uses the second of these phrases (xiv), telling us that his book is an example
of “The newer approaches to science, emphasizing its class and social under-
pinnings” (xvi), that it “looks at evolution’s use in order to understand the
class, religious or political interests involved,” that it is “a story of Class,
Power, and Propaganda” (xiv). As such, it is, as he says near the end of his
book, “a reaction to the old history of ideas, which displaced the person,
made him or her a disembodied ghost, a flash of transcendent genius” (617).
Similarly, in their biography of Darwin, Desmond and Moore (1992) state
that “Our Darwin sets out to be different—to pose the awkward questions,
to portray the scientific expert as a product of his time; to depict a man grap-
pling with immensities in a society undergoing reform” (xviii), or in other
words “We want to understand how his theories and strategies were embed-
ded in a reforming Whig society” (xix).
At its most extreme, the new historiography, in league with the “strong
program” in the sociology of science, sees science as a mere social construc-
tion, its evidence and facts and theories no more objective or capturing re-
ality than the stories of the Homeric gods, so that the distinction between
externalist and internalist history is denied—denied because the internal
development of scientific ideas (the ideational, intellectual, epistemic view
of history) is itself denied (cf. Moore 1997, 290). In a sense, the new his-
toriography is the exorcised ghost of behaviorism, with its emphasis on the
environment, reincarnated into an entirely different discipline. But unlike
behaviorism, which considered itself a genuine science, the new historiog-
raphy is varyingly postmodern in its orientation, which denies that there is
any real distinction between fact and fiction, truth and falsity, objectivity
and subjectivity. Instead, looking to Nietzsche as their guide (which, inci-
dentally, he would have despised), who proclaimed that the essence of all
life is will to power, postmoderns dutifully denounce any claim to objec-
tive evidence and rationality as a disguise, as a mask hiding will to power,
as Michel Foucault did against Noam Chomsky in his debate with him
(Smith 1999, 181). Forget scientists testing theories against evidence, or
their theories progressing toward truth; everything ultimately is a matter of
will to power.1
Arguably there is much of this in Desmond’s (1997) biography of Hux-
ley, where in the very Introduction he tells us that “At the dawn of the
twenty-first century ‘reason’ seems a precarious, value-laden yardstick, and
 Darwin and the New Historiography 209

one which has an infuriating habit of changing allegiance” (xv). Indeed he

sees Huxley, in his effort to secularize and professionalize science and take
it out of the hands of the clergy, as “establishing a rival evolutionary priest-
hood” (xvi), such that his biography of Huxley is “the human story behind
these sea changes” (xix). In all of this, Huxley the scientist is noticeably
missing (cf. Lyons 1999). Given that Desmond’s book is an attempt “To
understand . . . the making of our modern Darwinian world” (xx), if we
connect the dots he seems to be saying that our modern Darwinian world,
including modern evolutionary biology, is merely a “sea change,” subject to
“changing allegiance.” Kuhnian stuff at its most extreme (which Kuhn, in
spite of the way he wrote, would always denounce as a misunderstanding
of his view). Is this what is really behind Desmond and Moore’s (1992)
biography of Darwin? Michael Ruse (1993b) calls it “social constructivist”
(229). In chapter 8 (note 15) we have already seen one example of the way
in which their social constructionism operates. Later in the present chap-
ter we shall see some further examples.
It only needs to be noted at this point how utterly alien social con-
structionism, along with its epistemological relativism, is to the modern
scientific mind, for whom, as Dobzhansky (1973) approvingly put it,
“nothing in biology makes sense except in the light of evolution.” This
quotation, not accidentally, is repeated and emphasized near the beginning
of all three textbooks on evolution that I just happen to have at my home
(Ridley 1993, 5; Freeman and Herron 1998, 51; Futuyma 1998, 3–4).
Most of the historians that I shall deal with in this chapter would, I
strongly suspect, deny that they are part of what I call “the new historiog-
raphy.” Perhaps they all would. It does not matter. What we shall see is
that, whether they rightly belong or not, they share in common—at least
on the topic of Darwin on species—the following of a trend in their pro-
fession, namely, the desire to put scientists in general and Darwin in par-
ticular in his place, to embed him in his Victorian context, to keep him
there, and to either diminish or deny his importance in what (typically by
others) has often been called the most important (and still ongoing as I
shall argue) scientific revolution of all time. It is all a matter of degree. Some
professional historians do this more, some less. In this chapter I hope to try
to counterbalance this trend, keeping of course my focus on Darwin and
the nature of species.
Perhaps a good place to begin, and it will surprise many, is Peter
Bowler’s (1988) book, controversially titled The Non-Darwinian Revolution.
To the naive browser at the local bookstore or library, this title might sug-
gest that Darwin never started a revolution at all. Of course, creationists can
210 DARWIN AND THE NATURE OF SPECIES

put a lid on their hallelujahs, for once one reads the book one finds that
what the title might suggest is not the reality at all. Bowler does indeed rec-
ognize that there was, or is, a Darwinian revolution, but he does not think
it was started by Darwin’s Origin. At the very heart of Darwinism, “the true
essence of Darwinism” as he calls it (7), is not evolution but evolution by
natural selection (in the form of adaptation), and it is natural selection,
Bowler argues convincingly, that was rejected by the great majority of biol-
ogists from the time of the Origin right through to the forging of the Mod-
ern Synthesis, which began in the 1920s. (And even that, says Bowler, was
not so much of a Darwinian revolution, since it in large part incorporated
the genetics started by Mendel.) Thus for Bowler, the Origin did not start
a Darwinian revolution; instead it merely served as a “catalyst” (5) for non-
Darwinian evolutionary views, including mainly Lamarckism, orthogene-
sis, saltationism, and theistic evolution, all of them teleological to the core
and all of them coming to an end in biology with the advent of the Mod-
ern Synthesis. Natural selection throughout this intervening period, when
it was accepted, was typically seen as a minor mechanism in the pageantry
of evolution. Hence the non-Darwinian revolution, the received view that
the Origin began the Darwinian revolution being a “historical myth,” a fur-
ther example of “Whig history” (16).
There are two points I want to make about this, and they each have to
do with the topic of Darwin on the nature of species. First, I am inclined
to agree with Ernst Mayr (1985) that Darwin did not have one theory but
five, and that together they are properly to be understood as what Bowler
calls “the true essence of Darwinism.” According to Mayr, these are evolu-
tion as such, common descent (that every living species is descended from
an ancestral species), the multiplication of species (the branching concep-
tion of life), gradualism, and natural selection. Mayr argues that the first
three of these theories were accepted by the great majority of naturalists
within fifteen years of the publication of the Origin (1859) and that the
remaining two theories became accepted as a result of the Modern Synthe-
sis. Later, Mayr (1991) called the acceptance of the first three theories the
“first Darwinian revolution” (12, 25, 107) and the acceptance of the latter
two theories the “second Darwinian revolution” (89, 132). In other words,
the Darwinian revolution is not a misnomer; it was neither a one-theory nor
a one-episode affair.
That Darwin’s Origin would start a revolution, and not be a mere cat-
alyst for a non-Darwinian one, was perceived at the outset, and I don’t
think these testimonials should be disregarded. Watson, for example, upon
reading the Origin, wrote to Darwin (November 21, 1859) “Your leading
 Darwin and the New Historiography 211

idea will assuredly become recognized as an established truth in science,

i.e. ‘natural selection’.—(It has the characteristics of all great natural truths,
clarifying what was obscure, simplifying what was intricate, adding greatly
to previous knowledge. You are the greatest Revolutionist in natural history
of this century, if not of all centuries” (Burkhardt and Smith 1991, 385).
Similarly, Wallace wrote in a letter to George Silk (September 1, 1860) “I
have read it [the Origin] through five or six times, each time with increas-
ing admiration. It will live as long as the ‘Principia’ of Newton. . . . Mr.
Darwin has given the world a new science, and his name should, in my opin-
ion, stand above that of every philosopher of ancient or modern times. The
force of admiration can go no further!!!” (Burkhardt et al. 1993, 221 n. 1).
Darwin himself, of course, also recognized the revolutionary impact
that his work would have, and he continually looked to posterity. In letters
to Hooker, for example, he wrote “And what a science Natural History will
be, when we are in our graves, when all the laws of change are thought one
of the most important parts of Natural History” (July 30, 1856), “When-
ever naturalists can look at species changing as certain, what a magnificent
field will be open,—on all the laws of variation,—on the genealogy of liv-
ing beings,—on their lines of migration &c &c” (July 13, 1858), “When
we are dead and gone what a noble subject will be Geographical Distribu-
tion!” (December 31, 1858) (Burkhardt and Smith 1990, 194, Burkhardt
and Smith 1991, 130, 231). Later in a letter to John Innes (December 28,
1860) he made the comparison to the Copernican revolution: “By far the
greater part of the opposition is just the same as that made when the sun
was first said to stand still & the world to go round” (Burkhardt et al. 1993,
540).2 But it is in the Origin that we find Darwin’s most specific remarks
on the nature of the revolution that he envisioned would begin. There he
says species will no longer be considered to have an essence (484–485),
terms such as “community of type” will have a clear meaning (485), “A
grand and almost untrodden field of inquiry will be opened up, on the
causes and laws of variation, on correlation of growth, on the effects of use
and disuse, on the direct action of external conditions, and so forth. The
study of domestic productions will rise immensely in value” (486). He goes
on to add that “Our classifications will come to be, as far as they can be
made so, genealogies” (486). Psychology, he says, “will be based on a new
foundation,” and “Light will be thrown on the origin of man and his his-
tory” (488). Moreover, rather than evolution lowering the dignity of man,
non-human animals will instead become “ennobled” (489). How utterly
right was Darwin in every one of these, from antivivisection and animal
cruelty laws to the discovery of the chromosomal basis of heredity to the
212 DARWIN AND THE NATURE OF SPECIES

discovery of DNA, to phylogenetic taxonomy, to paleoanthropology, to

sociobiology and evolutionary psychology, through to animal psychology
and ape language studies. What needs to be remembered is that Darwin,
not in the Origin but in other works such as The Descent of Man (1871) and
The Expression of the Emotions in Man and Animals (1872), himself opened
many of these further fields of research, in particular, evolutionary ethics
and evolutionary psychology.
Indeed I would go further than Mayr and suggest that there are really
three Darwinian revolutions, the first and second having already occurred as
Mayr argued (though with the exception of the species concept, since it is
still in turmoil), the third being what we are currently in the midst of. Dar-
win himself recognized this third revolution, as the previous paragraph
shows, but of course he could not possibly have envisioned the extent of
that revolution. What I am referring to is the spilling over of Darwinian
explanations outside the confines of biology as normally circumscribed. This
third revolution has been ably discussed by Daniel Dennett in his book
Darwin’s Dangerous Idea (1995). It is arguably the most important (and still
ongoing) scientific revolution of all time, which Dennett likens (approv-
ingly) to a “universal acid” (63). Darwinism is a universal acid, for Dennett
(and many others, e.g., Dewey 1910, 2, 19), not just because it ate through
what were our traditional and most cherished beliefs in biology (e.g., cre-
ationism), but because it has leaked out into virtually all areas of life, in-
cluding religion, ethics, politics, and psychology. This is the debate raging
in virtually every university in the world today (cf. Stamos forthcoming).
But enough of this. What has any of the above to do with Darwin on
the nature of species? Returning to the theme of Mayr’s first Darwinian
revolution, Darwin’s Origin effected an enormous paradigm shift on the
concept of species, changing it in roughly ten years or so from an essentially
static, creationist concept to an evolutionary (or in some cases nominalis-
tic) one. That was an enormous and truly revolutionary achievement.
Granted, Darwin’s own species concept was not understood (and who can
blame his contemporaries for that?), moreover the new species concepts
following the first revolution were teleological, as Bowler (1988, 5) rightly
points out. But that should not be used to obscure the fact that Darwin
effected a revolutionary change. For a start, he abolished essentialism for
species. As long as species were conceived to be fixed types, there could be
no evolution of species. Moreover, he radically raised the importance of
variations (as well as varieties), in contradistinction to their previously low
status. This change was accepted, in one way or another, throughout
Bowler’s non-Darwinian revolution. But just as important with all of this,
 Darwin and the New Historiography 213

Darwin abolished the belief in what were thought to be two laws of species,
belief (as Darwin well knew) that posed a major obstacle to the change
from a creationist to an evolutionary view, namely, the belief in the law of
limited variation and the law of reversion (cf. chapters 2 and 7).3
With regard to species and Mayr’s second Darwinian revolution, Dar-
win, of course, did not effect a concept change, since (contrary to what Mayr
and some others would like to believe) the concept of species in modern
biology, if we look from the time of the Synthesis up to the present, was never
settled. In fact it may even be thought to be more unsettled today than thirty
years ago (Stamos 2003). That means, of course, that it is not over yet, and
it still may be that two key features of Darwin’s species concept—namely, his
emphasis on the priority of the horizontal dimension for species reality as
well as his advocacy of pattern over process, discussed in chapters 3 and 6
respectively—may eventually win the day. Only time will tell.
There is a second aspect of Bowler’s book (1988) that I said I wanted
to get to, but before I get to it we’re still not quite finished with the first
point. As I pointed out above, according to Bowler, Darwin eliminated
teleology from his species concept. This is because variation, for Darwin,
the very stuff that feeds natural selection, is essentially random with respect
to the environment, and the evidence for this Darwin got from the breed-
ers (29). But the Origin did not eliminate teleology from biology, not in the
post-Origin period up to the Modern Synthesis. Instead, teleology re-
emerged in various forms of non-Darwinian evolution, using the develop-
mental model of embryogenesis, even in self-proclaimed Darwinists such
as Haeckel (13, 83–90). As Bowler puts it, “the developmental model, in
one form or another, continued to dominate late nineteenth-century
thought. The materialist aspects of Darwin’s theory which appeal to mod-
ern biologists were not typical of his own time; indeed, they were so radi-
cal that hardly anyone could accept them. . . . The antiteleological aspects
of Darwin’s thinking prized by modern biologists were evaded or subverted
by the majority of his contemporaries” (5).
Interestingly, the claim that Darwin’s species concept, at the core of his
theory of evolution, was non-teleological has been denied by Robert
Richards. According to Richards (1992), Darwin “conceived embryological
evolution and species evolution as really two aspects of the same process:
both kinds of evolution involved a gradual morphological change through
a sequence of the same type patterns” (xiv). Again he says “I will indicate
how, as I believe, embryological development became for Darwin a model
of descent, infusing his conception precisely with those attributes, especially
notions of progress, usually thought to characterize only non-Darwinian,
214 DARWIN AND THE NATURE OF SPECIES

romantic theories of evolution in the nineteenth century” (3). Noting that

biologists today agree that three older proposals, common in the ninteenth
century, should be dismissed—“that species evolution should be modeled
on individual evolution, that embyogenesis recapitulates phylogenesis, and
that evolution is progressive” (179–180)—Richards points out that Darwin
in his early notebooks had all three. Dividing the “long gestation” of Dar-
win’s evolutionism into three stages (which begins roughly when he returned
from the Beagle voyage and continues through to 1859), Richards finds that
in the first stage Darwin “considered species to be comparable to individu-
als,” with a predetermined life span and the adapting mechanism of herita-
ble effects of environmental agents. In the second stage, Darwin retained
the adapting mechanism of heritable effects of environmental agents, gave
up the predetermined life span, and took on branching evolution. There is
no problem so far. It is the third stage that is troublesome. In this stage, says
Richards, Darwin added the Lamarckian emphasis on habit and the in-
heritance of acquired characteristics (but without the Lamarckian compo-
nent of conscious will), and he of course added as well (after reading
Malthus) his own contribution of natural selection. According to Richards,
“Though he regarded these ‘Lamarckian’ instruments as no longer central to
the production of new species, he did retain them as various auxiliary mech-
anisms in the Origin of Species” (84). Again, still no problem. The problem
begins when Richards claims that with natural selection, even in the Origin,
Darwin maintained that “species evolution should be modeled on individ-
ual evolution” (179). As he puts it, Darwin considered evolution of the em-
bryo and evolution of the species “as virtually the same process” (168).
Again, he says Darwin used “embryological evolution as a model of species
evolution” (169) and that “indeed, for Darwin embryological evolution be-
came part of the causal matrix that produced species evolution” (169).
In all of this Richards (90–99, 104–108, 111–166) makes much of
Darwin’s apparent use of recapitulation in the Origin (that ontogeny reca-
pitulates phylogeny). He claims to have “discovered that Darwin’s theory,
from its conception to its maturation, pulsed to the rhythms of that ever-
fascinating principle” (xiv), so much so that Darwin’s evolutionism con-
tained “teleological factors, more than a whiff of which the history of
recapitulation exudes” (176). Granted, as Richards recognizes, natural
selection is a force operating on a species from without. Nevertheless, he
sees Darwin being heavily influenced by, and thus retaining even in the
Origin, the embryological model of evolution proffered by Continental em-
bryologists. Although Darwin “did reject the hypothesis of an intrinsic cause
of necessary progress buried in the interstices of organization,” Darwin, says
 Darwin and the New Historiography 215

Richards, simply externalized the cause so that natural selection, along

with the subsidiary Lamarckian mechanisms, “would exert, as it were, an
external pull, drawing most organisms to greater levels of complexity and
perfection” (86).
This is a matter of ongoing debate, in particular between Richards and
Michael Ruse (e.g., Ruse 1993a; Richards 2002; Ruse 2004; Richards
2004). One problem I find is with Richards’ claim that the Continental
embryologists were also species evolutionists. According to Richards, the
recapitulation theories of Kielmeyer (19), Tiedemann (43–45), Treviranus
(45–46), and Meckel (54)—but excluding Serres (167)—were wedded to
a view of species evolution. None of the passages he cites in support of this
interpretation, however, with the exclusion of Serres, are convincing. For
example, he quotes Kielmeyer as saying “the force by which the series of
species has been brought forth is one and the same in its nature and laws
as that by which the different developmental stages [in embryogenesis] are
produced.” It is quite a stretch to read this as a belief in species evolution,
especially given that the Great Chain of Being, believed in by almost every-
one from the time of Locke to Darwin, was conceived to be an essentially
static, nonhistorical series (Lovejoy 1936). Thus the “force” that Kielmeyer
is referring to, for all that Richards’ readers know, could simply be God, re-
sponsible equally for both the static series of species through time and also
for the developmental stages in embryogenesis. The quotations from Tiede-
mann and Treviranus are hardly any better, and can easily be read as refer-
ring to species in the static Chain, given that the authors use words like
“seems” (Tiedemann) and “appears” and “can be compared” (Treviranus).
Meckel uses “probably,” but we are not given the context so that we can
determine what he means by “larger and smaller collections of organisms.”
Interestingly, Lyell (1832, 62–64) discussed the recapitulation theories of
Tiedemann and Serres, but rejected them as providing any support for
species evolution. Darwin, of course, would have read this, as he consumed
Lyell (1832) while onboard the Beagle. He would also have noticed that
Lyell does not ascribe to the two Continental morphologists a belief in
species evolution. Instead he says their recapitulation findings “has appeared
to some persons to afford a distant analogy, at least” to the theory of “pro-
gressive development” (62). For Lyell, instead, the recapitulation evidence
only provides evidence of a “unity of plan” (64).
Another problem I find with Richards’ thesis is his appeal to Darwin’s
use of recapitulation in the Origin. First of all, Darwin’s acceptance of it is
actually quite limited. Rather than say the development of the embryo of
a higher mammal recapitulates the evolutionary stages of vertebrates from
216 DARWIN AND THE NATURE OF SPECIES

fish, to reptiles, to birds, to mammalian features, he simply tries to explain

why this might appear so and he uses it as further evidence for evolution. For
example, he says “the adult differs from its embryo, owing to variations
supervening at a not early age, and being inherited at a corresponding age.
This process, whilst it leaves the embryo almost unaltered, continually adds,
in the course of successive generations, more and more difference to the
adult” (338; cf. 439–450). This is simply an attempt to explain, from his
own evolutionary point of view, something that many others had claimed
to observe. Moreover, recapitulation for Darwin only “partially shows us”
earlier stages (449), and he explicitly refrains from going as far as Agassiz in
proclaiming recapitulation a law of nature4—which is not at all what we
should expect him to do if Richards is right! Recapitulation was neverthe-
less important for Darwin on two fronts, so that he hoped it would be
proven a law (449). First, it provided further evidence for evolution, since
“The point of structure, in which the embryos of widely different animals
of the same class resemble each other, often have no direct relation to their
conditions of existence” (439–440). In this sense it was like homologies, a
point Darwin himself stressed (440). In either case the resemblances made
no sense from a creationist point of view. Second, recapitulation, being a
largely accepted part of embryology, would give further importance to em-
bryology as an aid in classification, settling, for example, that a barnacle is
really a crustacean (441). Once again, for Darwin, the importance is that
“community in embryonic structure reveals community of descent” (449).
In all of this it is quite a stretch to see teleology in Darwin’s use of re-
capitulation. To the extent that he accepted it, whatever that was, recapit-
ulation was always for Darwin something entirely backward looking, not
forward looking. As such, there is nothing teleological there. Embryology
itself, on the other hand, was at the time of the Continental morphologists
(with its etymological use of the word “evolution” in its embryology), as
well as today (with its concept of the DNA program in development), en-
tirely forward looking. Whether old or new, embryology is teleological.
This difference, between Darwin’s entirely backward-looking use of reca-
pitulation and the forward-looking meaning of embryology, is enormous,
and it seriously undermines any claim that for Darwin embryological evo-
lution and species evolution “are virtually the same process.”
There is, of course, much more that should erode our confidence in
Richards’ thesis, and we have already seen much of it in this book. We have
seen (i) that Darwin gave up the organism analogy for species in favor of
the language analogy, (ii) that like languages he conceived of species as pri-
marily horizontal entities, (iii) that his species concept was a pattern species
 Darwin and the New Historiography 217

concept, not a process species concept, and as such could have no teleology
in it, (iv) that he avoided the word “evolution,” preferring “descent with
modification” instead, and refused to call his theory “developmental,” (v)
that in the Origin he called variation under domestication “our best and
safest clue” (4), (vi) that he repeatedly called species “plastic” (12, 31, 80,
132), (vii) that he maintained throughout that their plasticity followed from
his view that variation, without which natural selection can do nothing (82),
is random with respect to the environment (30, 46, 61, 81, etc.), and (viii)
that in spite of his language in the Origin he did not think of natural selec-
tion as a real force or power.5 What more could one possibly want for a non-
teleological species concept and a non-teleological process of evolution?
Richards, however, seems consumed with selectivity of evidence. He
recognizes that the received view gets support from Darwin himself, in his
repeated claim “against Lamarck’s idea of an ‘innate tendency toward pro-
gressive development’” (85). In his footnote, Richards (n. 45) cites as his
source an 1872 letter from Darwin to Alpheus Hyatt, and he says a simi-
lar remark is to be found in Darwin’s Sketch of 1842 (Darwin 1909, 47).
And of course there is much more in this vein (e.g., Darwin 1859, 412,
351, 1863a, 79, 1876a, 87; Burkhardt et al. 1993, 577–578). But the real
point is that, all things considered, Richards has provided a paltry effort at
examining contrary evidence. Instead, in his book he continually presses
on with his thesis, trying to shove his square peg into a round hole.
It seems indisputable that the force of ideology, not evidence, is here at
the core. Richards says the received view on Darwin’s non-progressionism
is itself laced with “the darker byway of ideology” (xv), that ideology being
the Modern Synthesis, which eschews recapitulation and progress
(174–180). Richards ever so briefly hints that his own reinterpretation of
Darwin also has behind it a motivating ideology, when he writes “My own
narrative undoubtedly depends on some considerations that bend history
in ways that would appear distorting to others” (174). But anything more
than this is met with silence. I suggest that the ideology that drives Richards
is a common one among modern historians, the one we find in Desmond
and Moore and in other authors examined in this chapter, namely, to keep
Darwin in his place, to not allow him to be a man ahead, let alone well ahead,
of his time. This is the rallying cry of the new historiography, with respect
to scientists in general and to Darwin in particular, and with which I am here
in dispute. And indeed at the very end of his book Richards seems to briefly
show his hand, when he writes “Darwin was indeed the architect of the the-
ory that has been reconstructed as neo-Darwinism. But the architect was
our ancestor, who dwelt happily enough in the nineteenth century” (180).
218 DARWIN AND THE NATURE OF SPECIES

As if to further appease adherents of the received view, Richards adds at the

beginning of his book that “The Darwin that emerges from this study will
appear decidedly more venerable than the rejuvenated evolutionist who has
been injected by some historians with the monkey glands of a modern
scientific ideology” (3–4). But it is hard to see how Darwin becomes more
venerable from Richards’ study. Rather than more, it would seem less. And
that indeed seems the whole point to the new historiography.
I said earlier that there is a second point to Bowler’s book that I would
get to later on, and here is a good place to deal with it. Noting that Dar-
win had not only read Malthus but also Adam Smith and other political
economists, Bowler (1988) writes

The concept of divergence through specialization reflects the economic

advantages supposed to accrue from the division of labor. Even the Dar-
winian concept of species seems to reflect the individualist model of
society. The orthodox typological view of species can be compared with
a totalitarian political philosophy; individual organisms must conform
to the specific type just as individual human beings are supposed to sub-
merge themselves in the higher reality of the state. To some extent, Dar-
win saw the species as a population of varying individuals with no fixed
type, just as laissez-faire economics sees society as composed of individu-
als with divergent interests. [36]

This is exactly the sort of thinking that one would expect from the new his-
toriography, and it is particularly interesting since it focuses on the choice of
species concept, as something culturally determined. It is not entirely clear
whether or to what extent Bowler himself subscribes to the view that he ex-
presses here. The passage appears in the context of Bowler’s discussion on the
debate between the Darwin industry (which Bowler says basically adheres
to the myth of the Darwinian revolution) and the sociologists of science.
Bowler deserves the benefit of the doubt, however, since immediately follow-
ing the above passage he states “The possibility that Darwinism reflects cer-
tain aspects of Victorian society is accepted by the majority of modern
scholars, but this does not commit them to the view that natural selection is
nothing more than a projection of the capitalist ethic onto nature” (36).
Contrary to what Bowler suggests at the time of his writing, it would
seem that epistemological relativism, in the form of sociocultural relativism,
has become much more common in professional history of science. Indeed
Bowler’s statement above was written just before the books by Desmond
(1989) and Desmond and Moore (1992), both of which received book
awards and have been hailed by an increasing number of professional
 Darwin and the New Historiography 219

historians as paradigms of history of science writing (cf. Mauskopf 1992,

279; Shortland and Yeo 1996, x)—and one can only imagine the flock of
graduate students following. Times have indeed changed since 1988. Hence
David Hull (2000) writes of the “sociocultural turn” (72) in history of sci-
ence, with its “epistemological relativism” (75) and its “postmodernists”
(80), and Mary Winsor (2001) makes a powerful appeal for the “emancipa-
tion” of professional history of science from its self-imposed “Taboo Prob-
lem,” a taboo that amounts to “blacking out the very center of the topic we
claim to study” (240), the center being the “progressive direction of scien-
tific change” (241)—by which she clearly means epistemological progress.
At any rate, in the case of Darwin the new historiography tends to
make much of the supplanting of the old aristocracy by the rising middle
class of the Industrial Revolution, of which Darwin’s family, on both sides,
was part of. It also tends to make much of the influence of the backward
Victorian attitude toward sex. I shall return to the latter thesis below. Be-
ginning with the former, one could easily relate Darwin’s view of diverg-
ing species, of species evolution driven not only by natural selection but
also by ecological divergence, with the emphasis on the division of labor by
the capitalist economists of Darwin’s time. In the Origin Darwin himself
hints at a relation between the two, stating that “The advantage of diversi-
fication in the inhabitants of the same region is, in fact, the same as that of
the physiological division of labour in the organs of the same individual
body—a subject so well elucidated by Milne Edwards” (115–116). In Nat-
ural Selection (Stauffer 1975, 233) the analogy is likewise to physiology,
but elsewhere and earlier in his notes Darwin explicitly drew the analogy to
economics (cf. Ospovat 1981, 181). Desmond and Moore (1992) make
much of this. The problem for Darwin was to explain the branching, tree-
like diversification of nature, reflected in the static hierarchical classifica-
tions of his fellow naturalists. The answer for Darwin, claim Desmond and
Moore, came not from nature itself but from the political economy that
dominated his society as well as his own personal life. Thus not only do they
describe Darwin’s mechanism of natural selection as “his Malthusian, cap-
italist, competitive mechanism” (413), but we’re told that “Just as his
Malthusian insight had come from population theory, so his mechanism for
creating diversity looked like a blueprint for industrial progress. Darwin
was a heavy investor in industry. His Wedgwood cousins were among the
pioneers of factory organization. They created the production-line mental-
ity with a marked division of labour among the work force” (420). Again,
for Darwin “The creation of wealth and the production of species obeyed
similar laws. Division of labour was nature’s way as well as man’s” (420–421).
220 DARWIN AND THE NATURE OF SPECIES

Again, Darwin used “the zoologist Milne-Edwards’s use of the term ‘division
of labour’ rather than an economist’s” because “A political taint would have
made natural selection too much of a target” (421). We are told further-
more that Darwin had alternative models to choose from, such as that of the
economist Jean Sismondi (his wife Emma’s uncle), according to whom the
division of labor was unjust and those who worked the land should own it.
But Darwin rejected this and other models, we’re told, because “he was
himself an absentee landlord in Lincolnshire” (421). We find that Darwin
“put his money where his mouth was,” that “He spent tens of thousands of
pounds on railway companies, and twenty years of his life revealing the
competitive, specialized, and labour-intensive aspect of Nature’s ‘work-
shops,’” that “He was placing Nature on industry’s side” (421).
In all of this there is nothing about evidence but only hidden motive. A
theory of motive, however, is an empirical thesis, and empirical theses don’t
become facts simply by being claimed. What is required is supporting evi-
dence. There are certainly interesting correlations between the socioeconomic
conditions in Darwin’s time and Darwin’s theory of evolution by natural se-
lection and divergence. But correlation is not causation, and any argument
from correlation to causation needs something more—something much
more, as students of fallacies and medical researchers know only too well.
But Desmond and Moore don’t seem much interested in weighing evi-
dence pro and con. Correlation seems sufficient for them. Nor do they seem
interested in the principle of charity, giving Darwin the benefit of the doubt
until there is strong enough evidence against him. Instead, they have an
agenda, and they spare no effort in pushing it. For example, barely (448) do
they mention Darwin’s enormous work on small and large genera, in which
he argued, from tables of classifications by naturalists and related information
supplied by Watson, Hooker, and Gray, that species in large and wide-rang-
ing genera have more varieties than species in small genera, important corrob-
oration for his principle of divergence (Stauffer 1975, 134–164; Darwin
1859, 53–59). Perhaps the numerical analysis he accumulated was not after
all good from the viewpoint of modern statistics (Parshall 1982). Of greater
significance is what Kohn (1985b) has argued, that Darwin’s principle of di-
vergence was of the nature of an internal dialogue motivated by explanatory
unification, that the theory came first and the attempt to test it came shortly
after. From his biogeographical studies Darwin came to realize that natural
selection, even when combined with geographic isolation, seemed insuffi-
cient to explain hierarchical classification (something recognized by all, and
that Darwin temporalized as treelike). Natural selection therefore needed
something more, which he found in the concept of “places in the economy
 Darwin and the New Historiography 221

of nature,” sympatric speciation meaning for him that selection favors diver-
sification within a species. What is surely significant is that Darwin, as Kohn
(1985b, 256) has shown, developed his principle of divergence first, in
November of 1854, and then only later, much later in September of 1856,
connected it to economics and the division of labor (cf. also Ospovat 1981,
181). As Kohn puts it, “To mistake the labeling for the conception would, I
believe, be a misinterpretation of Darwin’s developmental process” (256).6
It is still possible that in developing and emphasizing his principle of
divergence Darwin was influenced by the political economy of his time, and
perhaps the emphasis on laissez-faire economics influenced his species con-
cept with its varieties within a species, along with the importance of the in-
dividual variants. Perhaps also Darwin’s (and Wallace’s) “discovery” of
natural selection was not really a discovery but a product of the same polit-
ical economy, projected onto nature, and that Darwin’s continued allegiance
to natural selection is to be explained by this environment (cf. Radick 2003
and Bowler 1988, ch. 2 as a counterbalance). But to “pull focus”—to use
Moore’s (1997, 296) cinematic metaphor—like that, to shift focus from the
individual in the foreground to the environment in the background, is to
miss so much more that was going on, such as Darwin’s observation of
variation in barnacles, his consilient argument for evolution by natural
selection (which included especially the evidence from breeders and bio-
geography), as well as his argument against the so-called laws of limited vari-
ation and reversion. Kohn (1985b) offers the following reconstruction
concerning Darwin’s principle of divergence: “As he [Darwin] comes to see
the species of local genera as the primary locus of divergence, he comes to
see small locales with no chance of geographic isolation as the primary cites
of speciation. Appreciating full well the swamping effect of crossing, he is
forced to invoke vigorous selection as the only effective countervailing force.
But more important, this line of thinking leads Darwin to look for the local,
hence ecological, conditions that favor vigorous selection” (255). To ignore
the role of problem, evidence, internal dialogue, and testing in Darwin’s
theorizing is simply poor scholarship.
Rather than there simply being a social constructionist influence from
the political economy of Darwin’s day (to which he seemed to subscribe)
to Darwin’s theory or principle of divergence, it may well be that Darwin
keyed into an important process that actually exists in multiple domains.
We have seen in chapter 3 that Darwin argued that natural selection is not
domain specific, that it also functions in language evolution. This is an in-
sight embraced by not a few linguists today (e.g., Lass 1997, 376–381).
Many today would also argue that Darwin recognized real similarities across
222 DARWIN AND THE NATURE OF SPECIES

two other domains, namely, economics and ecology, including the adaptive
response of diversification and specialization. In fact a whole journal now
exists for the single purpose of exploring these and other connections be-
tween biology and economics, appropriately titled the Journal of Bioeco-
nomics, founded in 1999 by Michael Ghiselin and Janet Landa. To dismiss
all of this as mere social construction or will to power is the easiest thing to
do, as long as one is not willing to take evidence seriously. The real skill is
to make inferences to the best explanation based on all the evidence avail-
able, but many, influenced by certain intellectual fashions, do not seem
willing to do this.
Whether multiple domains or domain specific, it is surely interesting,
and a further point in Darwin’s favor, that in recent years, in a longstand-
ing economic climate that embraces competition but disdains laissez-faire
economics, biologists have more and more been reassessing their previous
rejection of sympatric speciation (e.g., Ghiselin 1997, 159), so much so
that some biologists now claim that of all speciation models “the mecha-
nism of sympatric speciation proposed by Darwin . . . is the most plausi-
ble one” (Kondrashov et al. 1998, 97).
Another area the new historiography likes to make much of is sex, in
the case of Darwin and his contemporaries the Victorian attitude toward
sex. Desmond and Moore (1992) relate Darwin’s theory of sexual selec-
tion to his family life, in that “The Darwins fit the picture perfectly. The
Descent was essentially their story. Natural and sexual selection had made
and maimed them. Charles had strutted like ‘a peacock admiring his tail’
courting Emma. Coy and impressionable, she had selected him, admiring
his ‘courage, perseverance, and determined energy’ after a voyage around
the world” (580). She had selected him, never mind that he had seriously
weighed the pros and cons of marriage, as shown in his notes on marriage
of April and July 1838 (Burkhardt and Smith 1986, 443–445).
Here, as elsewhere, one can easily get carried away with social construc-
tionism. Gordon McOuat (2000), in his review of volume 10 of Darwin’s
Correspondence (Burkhardt et al. 1997), hints that the emphasis by Huxley
and others on the sterility criterion for species had a Victorian ring to it. As
McOuat puts it, for Huxley “Sexual sterility marks a true separation be-
tween species, and if natural selection could produce sterility, it could pro-
vide the origin of new species. At the very ground of the division of natural
kinds in biology was a sexual demarcation. There was something naughty,
forbidden here. In one way it rubbed against Victorian sensibilities” (191).
My God should Huxley now be thought more of a good Victorian? His
species were, after all, good Victorian species. And what of Darwin? Was he
 Darwin and the New Historiography 223

a decadent? Unlike Huxley, his species were bad Victorian species, some-
times with sterility within them,7 but worse—much worse—sometimes with
fertility between them. Surely there is sexual deviance afoot here, evidence
of a desire for promiscuity.8 Perhaps even bestiality! And wasn’t all of this
really what was behind the veil covering the Victorian attitude toward sex?
Didn’t Freud teach us that? Even more, Darwin’s rejection of “higher” and
“lower” in nature (e.g., Notebook B 74, 1859, 441, Burkhardt and Smith
1991, 228) may well be read as a symptom, and along with it as his own pro-
jection onto nature, of an underlying decay in Victorian society with regard
to its concept of “place” (i.e., the Victorian caste system with its layered
aristocracy at the top and the homeless poor on the bottom).
On the other hand, as we have seen in chapter 3, Darwin himself did
prefer a horizontal to a vertical species concept. This cannot be ignored, and
it should make one rethink one’s history. Correlation is causation, after all.
Somehow, someway, Darwin must be seen as “a cork bobbing on the sur-
face of the society of his day,” as Ruse (1993b, 229) characterized the new
historiography. Thus it may well be that Darwin was, in a sense, more of
a good Victorian after all, and by that I mean the overtly Victorian sense,
not the Freudian. Darwin almost certainly was influenced, probably sub-
consciously, by the Victorian insistence on the missionary position when he
conceived of species as primarily horizontal. And as I have shown in chap-
ter 3, he maintained that view throughout. His species were indeed, there-
fore, in a very different sense than before, good Victorian species (this was
his real torment). All of this, interestingly, can be traced back to Darwin’s
Beagle voyage, during which he got to observe much of the work of the
Christian missionaries, and we know from Desmond and Moore (1992,
173–176) that contrary to the critics back home he approved of their moral
effect. It is therefore only reasonable to suppose that Darwin knew very
well of the attempts by the missionaries to impose the horizontal position
on the poor savage aboriginals for when they conceive their children (hence
what we now call “the missionary position”). And probably this influence
on Darwin’s species concept was further reinforced by personal experiences
in his life, some of them not even overtly sexual, again such as his Beagle
voyage in which during seasickness “only a ‘horizontal position’ brought
relief ” (Desmond and Moore 1992, 115).
McOuat (2000) goes on to say “Freudians could have a field day with
Darwin’s envious astonishment over the size of the barnacle’s penis,” and
that in the correspondence “The married Darwin flirts; his sisters tittle”
(191). Did Darwin have an inferiority complex over his penis? Was this
common in Victorian society? Did he try to make up for his feelings of
224 DARWIN AND THE NATURE OF SPECIES

inadequacy by desiring sex outside the marriage? Was promiscuity seething

beneath the Victorian exterior? I do not say that McOuat is guilty of such
nonsense (he is not), but his remarks nevertheless invite the kind of garbage
that social constructionists seem to thrive on. It is tabloid historiography,
with all the irresponsibility and sensationalism that goes with the tabloid
journalism found in the neighborhood grocery store. Book awards, more-
over, don’t change this fact one bit.
Leaving my parody of the new historiography aside, a further area of
cultural influence that the new historiography likes to focus on is the scien-
tific culture of the time. Sometimes issues of scientific culture cannot be
clearly separated from politics and culture at large. This is the theme of Har-
riet Ritvo’s (1997) article on zoological nomenclature, where, following
Foucoult’s work (1971) on the eighteenth century, she finds the Victorian
debates driven mainly by the desire for dominion and control, whether
geopolitical or personal, including also patriotism and vanity. There is much
in her analysis that deserves merit and serious consideration, and yet, in a
way that has become all too typical among professional historians of science,
she goes too far. Confining ourselves to the case of Darwin, granted, Darwin
was involved in and supported the rules drafted by Strickland’s 1842 Com-
mittee. But with regard to an earlier draft he demurred (seconded by Leonard
Jenyns) on the side of foreigners where it stated that the British Association
for the Advancement of Science should properly be considered “the Parlia-
ment of Science” (Burkhardt and Smith 1986, 311). Moreover, he pointed
out in a letter to Jenyns (May–September 1842) that “Stricklands laws will
I think be useful in checking the egoism of some authors” (317). Darwin
himself, of course, had become impressed with nomenclatural problems dur-
ing his stay in London following his Beagle voyage and the cataloging of the
thousands of specimens he had brought back with him. Years later while
working on barnacles, in which he had inherited a nomenclatural mess from
his fellow naturalists, the need for rules and consistency became even more
important. But in none of Darwin’s practices do we find political or egoistic
concerns. He respected the priority of foreign authors (cf. Darwin 1854b,
2), and even when he felt the need to go against his fellow naturalists and call
a form a “species” he did not use his own name (e.g., Darwin 1854b, 250).
Nor in naming, for example, primroses and cowslips two species instead of
one was he trying to do something like raise the dignity of the common man
(who named them two species instead of one, contrary to scientific practice),
or any other such nonsense that the new historiography might come up with.
He kept the names the same. Instead, whenever he tried to correct the species
ascriptions of his fellow naturalists he did so purely for what he believed to
 Darwin and the New Historiography 225

be the soundest of scientific principles, namely, common descent partitioned

by adaptations produced by the law of natural selection.
This raises a further issue. Sometimes the influence of scientific cul-
ture is mixed very little if at all with the wider influences of politics and
culture at large. We have seen some of this already earlier in this chapter in
Richards’ (1992) attempt to make out of Darwin a Naturphilosoph. A much
more plausible approach is to look at the scientific culture of Darwin’s day
with its emphasis on explanatory laws of nature, along with its emphasis on
natural theology by way of the many beautiful adaptations to be found in
the living world. There is surely an influence here on Darwin’s species con-
cept—arguably more than any other area of influence—with its emphasis
on adaptations and their explanation by natural law. Not only was the dis-
covery of laws of nature highly prized in Darwin’s day, to the point of call-
ing phenomena laws that were not really laws, but laws also played a central
role, as we have seen, both in scientific explanation and scientific classifi-
cation. So great was this influence that Darwin called community of descent
a vera causa, even though this conflicted with his intuition that the extinc-
tion of a species is not necessarily forever.
However great the influence of this culture was on Darwin, we never-
theless still have to be very careful. For a start, adaptations were then as
now an extraordinary fact of nature, and they cry out for explanation. Dar-
win of course knew this, but his importance lies in the fact that he was the
first person in history to recognize that their explanation must be by nat-
ural selection.9 Modern biologists have become absolutely certain that Dar-
win was right here, and for this (and much more) they rightly honor him
(e.g., Dawkins 1986, ix, 6, 287).
In spite of Darwin’s importance here, however, which as we have seen
in this book is the key to understanding his species concept, some histori-
ans have tried to minimize Darwin’s importance by reevaluating the devel-
opments in taxonomy prior to the birth of Darwin’s crowning achievement.
Thus Mary Winsor (2003), for example, who for reasons given earlier I
greatly hesitate to include among the ranks of the new historiography, has
attempted to rewrite history by arguing that pre-Darwinian taxonomists,
contrary to the received view, were not in fact essentialists. There is noth-
ing new in this claim. It has been made, for example, by McOuat (1996),
as we have seen in chapter 8. What is new is Winsor’s claim that pre-
Darwinian taxonomists were clusterists (my term, not hers). Never mind
what they stated in their definitions, she argues, it’s what they did that mat-
ters, and their practice clearly indicates that they were not essentialists. Strict
essentialism classifies according to one or more conditions individually
226 DARWIN AND THE NATURE OF SPECIES

necessary and jointly sufficient for membership in the class. Clusterism, on

the other hand, involves a set of membership conditions, but no one condi-
tion is either necessary or sufficient for membership in the class, so that mem-
bership is a matter of degree. Clusterism, according to Winsor, is how we
should characterize pre-Darwinian taxonomists. What is more, following the
modern philosopher Richard Boyd (1999), who argues that species should be
conceived as homeostatic cluster classes, Winsor argues that because cluster-
ism does not in itself pose a barrier for evolutionism we should therefore look
at pre-Darwinian taxonomy as paving the way for the Darwinian revolution,
indeed as “constituting the foundation of Darwinism” (398).
The problem with her argument, at bottom, is not that Boyd and
others might be wrong, that homeostasis is a problematic concept and clus-
terism is arguably not compatible with evolutionism (as I have argued else-
where; cf. Stamos 2003, 224–230, 123–143). Instead, the fundamental
problem with her argument is that when dealing with pre-Darwinian tax-
onomists she does not discriminate between species and higher taxa. In
fact, all of her evidence for clusterism in the practices of pre-Darwinian
taxonomists comes from higher taxa. In my public debate with Winsor and
in my subsequent reply paper (Stamos 2005, 81–83), I provided further
evidence to support her in this area. For example, in a letter to Darwin
(April 26, 1844) George Waterhouse, a well-respected entomologist and
member of Strickland’s 1842 Committee, wrote “The term ‘typical species’
is used by Zoologists in two senses—it either refers to that species which
possesses in the highest degree of development some of the characters which
distinguish the group [‘I mean any assemblage of species’] to which it be-
longs from other groups; or, it has reference to that species which is sup-
posed to exhibit, in the best balanced condition, the greatest number of
characters common to the species forming the group of which it is a mem-
ber” (Burkhardt and Smith 1987, 30). Either way we have clusterism here,
and the evidence is overwhelming for such clusterism at the level of genera
and higher taxa. It was indeed the dominant view.
What Winsor has missed, however—and it is why we cannot afford to
not pay attention to what these naturalists say, including their definitions,
if we are to put labels on them—is that, again as I argued in the debate and
in my paper, the great majority of pre-Darwinian taxonomists did not be-
lieve in the reality of higher taxa from genera up. Buffon, Jussieu, Prichard,
Bentham, Lindley, Watson, and Hooker—representative examples with
enormous influence—were all higher taxa nominalists. Typical and uncon-
troversial here is what Agassiz (1857) wrote, that in spite of numerous dif-
ferent systems of classification “There is only one point in these innumerable
 Darwin and the New Historiography 227

systems on which all seem to meet, namely, the existence in nature of dis-
tinct species persisting with all their peculiarities . . . . Beyond species, how-
ever, this confidence in the existence of the divisions . . . diminishes greatly.
With respect to genera, we find already the number of naturalists who
accept them as natural divisions much smaller; few of them have expressed
a belief that genera have as distinct an existence in nature as species. As to
families, orders, classes, or any kind of higher divisions, they seem to be uni-
versally considered as convenient devices” (4–5). For most of these natural-
ists, then, they grouped species into higher categories simply out of practical
necessity. To call them clusterists is highly misleading. Methodologically
they were clusterists, but ontologically they were nominalists. For the great
majority of them, only species were real.
It is therefore an enormous mistake to look at these pre-Darwinian nat-
uralists and taxonomists, including Darwin’s contemporaries, and paint
them with the same brush when it came to species. Not every received view
is mistaken, although professional historians of science often seem bent on
disproving one received view after another. In the case of pre-Darwinian
naturalists and taxonomists, not only is the received view correct in that
most of them were species realists, but the majority of them were also species
essentialists. Again the evidence for the received view here is overwhelming.
For a start, as I argue more fully in my reply paper (Stamos 2005), most of
these naturalists and taxonomists believed in primordial descent, with each
species beginning with a single individual or pair. Second, most of them
were splitters, not lumpers, not allowing for varieties and making the vari-
eties of lumpers into species. Third, most of them believed in reversion as a
law, which allowed for some variation in characters but those characters
would revert back to the primordial type following the removal of the per-
turbing conditions (typically they thought this law relevant only for domes-
ticated species). Fourth and finally, most of them believed in an underlying
causal essence, passed on through generation, which accounted for rever-
sion and limited variation. We have seen some of this in chapter 7, in Lin-
naeus’ “unity in generation” where variation is in the “outside shell,” in
Buffon’s moule intérieur, in Jussieu’s “any individual whatever is the true
image of the whole species,” in Agassiz’s essences “forever fixed” in the mind
of God, and in Dana’s “specific law of force, alike in all.” Moreover, as we
have seen in chapter 1, what went hand in hand with this essentialism was
what Wollaston called the “axiom” of independent creation.
With the concept of species at its core, then, what Darwin’s revolu-
tion signifies is an enormous departure from the paradigm that existed be-
fore. Pre-Darwinian taxonomists and naturalists, contrary to Winsor, did
228 DARWIN AND THE NATURE OF SPECIES

not lay the foundation for Darwinism. Instead, they posed an enormous
barrier. To think and argue otherwise is to play into the hands of the new
historiography. It is, yet once again, to try to minimize Darwin’s impor-
tance, to see him as a mere extension of his culture.10 It is to follow a facile,
myopic, and ultimately pernicious trend.11
If we really want to see science as a social construction, we can do no
better than look at the Lysenko affair that occurred in Communist Russia
beginning in the mid-1930s (cf. Harman 2003). This episode illustrates as
perfectly as can be what happens when science really is a social construc-
tion, when it is based primarily on culture and ideology rather than on
nature. Previously among the leaders in the world, Russian biology under
Lysenko—with its forced rejection of Mendelian genetics (including the im-
prisonment and death of many geneticists), and also of Darwinian selection
on random variation, both of which were viewed as contrary to Marxist
dialectics, unlike the naive Lamarckism that Lysenko made mandatory—
came to a grinding halt, with enormous negative consequences for its agri-
culture. Only in the mid-1960s, when Lysenkoism was officially dismissed
by the Soviet state, did Russian biology and agriculture begin to resume
where they had left off, trying to play catch-up ever since.
The Darwinian revolution, on the other hand, with its five theories at
its core, has proved the exact opposite. It has opened up one fruitful re-
search program after another, leading to spectacular successes in knowl-
edge and understanding, so much so that biologists today routinely agree
with Dobzhansky that “nothing in biology makes sense except in the light
of evolution.” Darwin was a Victorian, to be sure. His was a time very dif-
ferent than our own in many ways, of course. But in so much of his think-
ing he belongs more in our day than his own. Biologists have recognized
this time and again, and not without good reason. Natural selection, for ex-
ample, has been amply documented in the wild (Endler 1986), it has
proved essential in understanding phenomena such as the evolution of bac-
terial resistance to antibiotics (Sonea and Panisset 1983, 86–87) and insect
resistance to pesticides (Ridley 1993, 101–106), and it is the key to under-
standing immune systems and the evolution of viruses such as HIV (Free-
man and Herron 1998, 3–25). In fact biologists, and to a lesser extent
philosophers, continue to find helpful insights and inspiration in Darwin’s
writings. Sexual selection is a good example, since the process itself, as well
as Darwin’s insights into it, have come to be appreciated by mainstream
biology only in the past three decades (Andersson 1994). Sympatric speci-
ation is an even more recent example (Kondrashov et al. 1998). While out-
side of biology proper, Darwin’s principle of divergence has come to be
 Darwin and the New Historiography 229

seen as a useful explanation of divergence in personality types among

siblings (Sulloway 1996, 84–85). Indeed evolutionary psychology, begun
by Darwin, finds that it owes much to Darwin (Crawford and Krebs 1998),
as does evolutionary ethics (Thompson 1995).
Darwin still has much to teach us in many areas, in spite of his mistakes
and shortcomings here and there. With mass extinction #6 in full swing as
the twenty-first century moves forward, now more than ever we need to
learn and take into full consideration what he has to say about the nature
of species. In his own time it would have fallen on deaf ears. But to ignore
him today, or to diminish his importance as the mere product of his time,
is simply arrogant folly. I do not know whether Darwin’s insights into the
nature of species, as I have brought them to light in this book, will ever ex-
perience any of the good fortune that so many of his other ideas have.
(Whiggish history this book is not, although my hope is that it someday will
be read as if it were.) But whatever the future holds, or should hold, surely
the time has come for Darwin to get a fair hearing on this head.
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 Notes


CHAPTER 1
1. Throughout the present book all references to the Origin are to the 1859 edition,
unless otherwise indicated.
2. This was indeed a common view of genera in Darwin’s day and before, even though
it was not shared by Linnaeus. For example, Bentham, who Darwin explicitly refers to on this
topic on p. 419, had long maintained (Bentham 1836) that “A genus . . . has seldom any real
existence in nature as a positively determined group, and must rather be considered as a mere
contrivance for assisting us in comparing and studying the enormous multitude of species,
which, without arrangement, our minds could not embrace” (xlvii; cf. Bentham 1858; cf. also
Stevens 1994, 99–109, for references on higher taxa nominalism in Bentham and in other
naturalists). In like manner Watson, in a letter to Darwin (March 23, 1858), confided that
“I look upon the orders & genera of plants as purely conventional arrangements, not natural
groups,—that is, not groups in nature, but only as groups in books & herbaria” (Burkhardt
and Smith 1991, 54). Similarly Hooker, in a letter to Darwin (February 25, 1858), stated that
“Genera in short are almost purely artificial as established in Botany” (Burkhardt and Smith
1991, 35). That genera and higher tax nominalism was the general view, cf. also Watson
(1845a, 142) and Agassiz (1857, 4–5), as well as Stamos (2005).
3. It is sometimes claimed that this was in fact not a common feature of species con-
cepts in Darwin’s day (e.g., McOuat 1996, 511, 515). And yet it is easy to find contempo-
rary sources who agree with Darwin on this point. For example, Watson (1845a) states that
“the very definition of the term ‘species,’ as usually given, involves an assumption of non-
transition” (147). He would later repeat this in a letter to Darwin (May 10, 1860), in which
he wrote “Until a faith in certain impassable barrier between existent species becomes thor-
oughly shaken, naturalists will resist your views, & hail difficulties as if conclusive argu-
ments on the contra side. Differently as these unseen barriers are traced or placed, they are
believed in about as strongly by almost all” (Burkhardt et al. 1993, 203). More specifically
Wollaston (1860), in a critical review of Darwin’s Origin, states that “The opinion among
naturalists that species were independently created, and have not been transmitted one from
the other, has been hitherto so general that we might almost call it an axiom” (133).

231
232 Notes to Chapter 1

4. Interestingly, Darwin seems to have got more from Watson than just the above strat-
egy. For in his correspondence with Darwin, Watson, in spite of what he says about the vari-
ability in species concepts between his fellow naturalists, seems still to have thought that
species are in a sense real entities in nature. For example, in a letter to Darwin (October 11,
1855) he wrote “I look upon the words ‘Orders, genera, species (of books), & varieties,’ only
as terms to indicate passably well the grades of resemblance between objects” (Burkhardt and
Smith 1989, 479). That Watson includes only book species is highly significant. Elsewhere
he made explicit his distinction between book species, which he considered arbitrary, and
natural species, species found in nature (Watson 1843, 618). Granted, a little later (Watson
1845b), as we shall see below, after focusing on the messy situation presented by primroses
and cowslips, and how they lend themselves to evolutionism, he came close to drawing a
nominalistic conclusion about species (219). However, he never went all the way, and in
later writings, in spite of his evolutionism, he seems clearly to have thought that there are real
species in nature, moreover that their reality is only or primarily horizontal in nature, at a
given time slice. For example, in a letter to Darwin (November 8, 1855) he wrote “I must
confess a pretty strong bias towards the view, that species are not immutably distinct;—altho’
in our time-narrowed observation of the individuals they seem to be so” (Burkhardt and
Smith 1989, 499). Similarly, in a later letter to Darwin (December 20, 1857), he wrote “In
writing the final volume of my Cybele Britannica, I find myself unable to carry out the ideas
or inquiries originally intended. And why?—Mainly, because the limits of species are so un-
certain in nature,—so dissimilar in books. . . . This leads me to devote many pages to my own
notions about species & classifications,—rather irrelevant in a book on local botany;—and
perhaps somewhat limping over that same ground which will be better trod by yourself. I can-
not find the proof of species being definite & immutable, whatever they may seem to be at
any one time & spot” (Burkhardt and Smith 1990, 511). Cf. Watson (1859, 31–34), where
he distinguishes between true species in nature, which he defines from an evolutionary point
of view (implicitly horizontal), and what he calls “false species,” for which he provides four
tests. Watson’s repeated emphasis on the general horizontal reality of species will prove
especially significant when we look at Darwin’s own view in Chapter 3.
5. In spite of Lewes’ confusion between species as a category and species as a taxon (cf.
Chapter 2), I find Lewes’ position extremely interesting for the simple reason that after the
above passage he immediately proceeds to provide an ontology for species based on similar-
ity relations, and one that happens to be close to what I shall argue in later chapters was in
fact Darwin’s view. According to Lewes, “Nature produces individuals; these individuals
resemble each other in varying degrees; according to their resemblances we group them to-
gether as classes, orders, genera, and species; but these terms only express the relations of
resemblance, they do not indicate the existence of such things as classes, orders, genera, or
species” (443). Later I will argue that Darwin conceived of species as horizontal similarity
complexes demarcated objectively by the law of natural selection. Moreover in the final
chapter of my book on the modern species problem (Stamos 2003), I develop an ontology
of species as relations by modifying Darwin’s ontology to include demarcating causal
processes besides natural selection. In all of this I cannot help but find in Lewes a most
interesting precursor.
6. This last clause is important, for in spite of his nominalism Lamarck (1809) did
allow that what are typically called species “have really only a constancy relative to the du-
ration of the conditions in which are placed the individuals composing it” (36). Thus for
 Notes to Chapter 3 233

Lamarck species on the traditional view, which involves “an absolute constancy in nature” as
well as the belief that “the existence of these species is as old as nature herself” (35), have no
real referents in nature. Their reality is conceptual at best. Does that mean that for Lamarck
species have a reality in nature in a different sense, in particular so long as there are no causal
changes acting up them, where “them” shares the traditional definition of “Any collection of
like individuals which were produced by others similar to themselves” (35)? This indeed is a
possible reading of Lamarck, although it was not the way he was commonly understood.
7. Particularly disappointing on this topic is the recently published anthology The
Cambridge Companion to Darwin (Hodge and Radick 2003). Not only does it contain next
to nothing on Darwin on the ontology of species, but Radick (2003, 165 n. 26) lumps my ear-
lier writings with Beatty, Hodge, and McOuat, no less as sources backing his view that for
Darwin naturalists and taxonomists, not nature, “invented” (162) and “divided species from
one another” (151), while Gayon (2003, 264 n. 59) uses my earlier writings as the sole refer-
ence for his view that Darwin made taxonomic categories, including that of species, “a mat-
ter of mere convenience” (260), even though I have never subscribed to the received view.
8. It is interesting to track the effect this line had on Gray. In Gray’s review of Dar-
win’s reviewers (Gray 1860c), published for October 1860, although he still ascribes to
both Agassiz and Darwin the view that species and varieties exist “as categories of thought”
(420–421), to which he adds the word “only” (421), he nevertheless explicitly ascribes to
Darwin the view that species in nature are “temporary” (406, 420), and he even goes so far
as to claim that “Darwin clearly maintains—what the facts warrant—that the mass of a
species remains fixed so long as it exists at all, though it may set off a variety now and then”
(424)—a view remarkably similar (though Darwin in fact did not have it) to the modern
theory of punctuated equilibria developed and maintained by Eldredge and Gould (1972).
Apparently when Gray wrote this paper he had the benefit of Darwin’s letter, and thus
could no longer, as he did in an earlier review (1860a) discussed near the middle of the
present chapter, simply ascribe to Darwin the view that species are “subjective.”

CHAPTER 2
1. According to Ghiselin (1989), among good candidates are Mayr’s theory of
allopatric speciation, according to which most species evolve as the result of a founder
effect, which Ghiselin says should be called “Mayr’s Law” (58); another is Bergmann’s Rule,
“which states that in colder climates animals get larger” (61); a further example is Dollo’s
Law (64), which will be discussed below.

CHAPTER 3
1. Interestingly, this rejected theory of species aging made its way into the first edition
of Darwin’s Journal of Researches (Darwin 1839), but was excised for the second edition
(Darwin 1845). In the first edition (1839), at the very end of his first chapter on Patagonia,
Darwin wrote “All that at present can be said with certainty, is that, as with the individual,
234 Notes to Chapter 4

so with the species, the hour of life has run its course, and is spent” (212). Cf. Gruber (1981, 261,
268–271) for an explanation of why this dated view of Darwin’s made it into the first edition.
2. Darwin insisted on gradualism for a number of reasons. In the Origin he gives at
least six: variation itself is a slow process, niche changes are generally very slow (recall Lyell’s
uniformitarianism), intercrossing further retards the process (108), saltations (macromu-
tions) are rare (10) and almost always are either maladaptive or nonadaptive (44), saltations
do not explain the ubiquitous co-adaptations both within species and between species (3–4),
and there is almost no known organ which is not known to have transitional steps (194). It
must be added that with his gradualism Darwin was not dogmatic. In reply to the botanist
Harvey, who argued that in the Royal Botanic Gardens at Kew a new species of Begonia had
arisen via saltation, which he called B. frigida, Darwin in his correspondence in 1860
(Burkhardt et al. 1993) replied to Hooker (February 20) that “Harvey’s is a good hit” but
that “It would take a good deal more evidence to make me admit that forms have often
changed by saltum” (97). To Lyell (February 18–19) he replied that “Harvey does not see
that if only a few (as he supposes) of the seedlings inherited his monstrosity natural selec-
tion would be necessary to select & preserve them” (93) and that (February 23) “On the
whole I still feel excessively doubtful whether such abrupt changes have more than very
rarely taken place” (102). And to Harvey himself (September 20–24) he wrote “About sud-
den jumps; I have no objection to them; they would aid me in some cases: all I can say is,
that when I went into the subject, & found no evidence to make me believe in jumps, & a
good deal pointing in the other direction—” (373). It is to be noted that in none of his
replies does Darwin speak of B. frigida as a “species”; instead he speaks of it only as a “form.”
This is significant, as we shall see in Chapter 5.
3. For examples from Darwin’s barnacle monograph that pretty clearly correspond to
this passage, cf. Darwin (1854b, 197, 243, 308).
4. Cf. Ospovat (1981, 255–256 n. 3) as well as Darwin in his Historical Sketch
(Burkhardt et al. 1993), in which he ascribes to Lamarck a belief in “a law of progressive de-
velopment” (573) and to Chambers a belief in “vital forces” such that “organization pro-
gresses by sudden leaps” (574).

CHAPTER 4
1. As for the “competent judges” Darwin mentions in the Origin, I have been unable
to ascertain their identity. Darwin’s section in Variation on cattle (Darwin 1868 I, 82–97)
is of little help here. The only pre-Origin source he mentions is a Memoir by a Professor
Nilsson (84 n. 37).
2. It is interesting to note that, contrary to Huxley who told Darwin in a letter
(before October 3, 1857) that he thought classification should be irrelevant of pedigree
(Burkhardt and Smith 1990, 461), Darwin thought that genealogical classification will
someday be achieved more or less. In a letter to Huxley (September 26, 1857) he speculated
that “The time will come I believe, though I shall not live to see it, when we shall have fairly
true genealogical trees of each great kingdom of nature” (456). Moreover in a letter to Lyell
(September 22–23, 1860), Darwin provided two very similar hypothesized genealogical
trees for mammals (Burkhardt et al. 1993, 379–380).
 Notes to Chapter 5 235

3. Interestingly, reproductive isolation is in fact not the only consequence of poly-

ploidy. In a detailed review of a considerable amount of literature on polyploidy, Levin
(1983) concluded that “The biological, physiological, and developmental changes which
incidentally accompany chromosome doubling may immediately adapt polyploids to con-
ditions other than those to which their diploid progenitors are adapted. In a sense, chromo-
some doubling may propel a population into a different resource or habitat space. . . .
Polyploidy not only alters the adaptive gestalt of populations, it promotes a series of genetic
and chromosomal changes which compound the differences between the polyploid popu-
lation and its progenitors. . . . Polyploidy may evoke large, discontinuous effects, which in
turn may lead to abrupt, transgressive and manifest shifts in the adaptive gestalt of popula-
tions which selection might accomplish slowly or not at all” (15–16). Levin’s use of the
words “adapt” and “adaptive,” of course, are problematic, since in modern evolutionary
biology, following Darwin, these terms are normally restricted to products of natural selec-
tion (cf. West-Eberhard 1992). We shall return to this issue in the next chapter.

CHAPTER 5
1. Darwin’s reasons for concluding common descent are many, and are sprinkled
throughout his chapter on the races of man.
2. In Natural Selection Darwin provides the example of North American Asters dis-
cussed by Torrey and Gray (Stauffer 1975, 196).
3. The only problem with this theory, as I see it, is that Darwin fully recognized that
there may be different adaptations within the same species. The problem is not so much with
the products of sexual selection, as Darwin regarded these as relatively superficial (cf. Dar-
win 1859, 88–89, 1871 I, 258–259), much the same as the products of artificial selection
compared with the products of natural selection (as we shall see below). Instead, the prob-
lem is with the differences within a species that are adaptations produced by natural selec-
tion, namely, sexual dimorphism where the differences are to “wholly different habits of life
in the two sexes” (cf. Darwin 1859, 87–88, 1871 I, 254–256), caste polymorphism (cf. Dar-
win 1859, 236–242), and ontogenetic dimorphism (cf. Darwin 1859, 440, 1868 II, 51–52).
Darwin definitely did not divide these differences into different species. Nor did anyone else.
As Darwin put it in the Origin, “With species in a state of nature, every naturalist has in fact
brought descent into his classification; for he includes in the lowest grade, or that of a species,
the two sexes; and how enormously these sometimes differ in the most important characters,
is known to every naturalist . . . yet no one dreams of separating them. The naturalist includes
as one species the several larval stages of the same individual, however much they may differ
from each other and from the adult” (424). What all of this indicates, it would seem, is that
Darwin’s use of the term “form” (e.g., Darwin 1859, 480, 485) had a populational aspect to
it. He only used different adaptations to separate different populations into different species,
never different sexes, castes, or ontogenetic stages. This is something that we shall see
repeated consistently by Darwin in example after example.
4. One can only imagine the shock and sting on Darwin’s part, then, after sending
Herschel a complimentary copy of the Origin, upon hearing that Herschel dismissed it as
“the law of higgledy-piggledy,” which Darwin heard “by round about channel” and which
236 Notes to Chapter 5

he took to be “very contemptuous,” adding that “If true this is great blow & discourage-
ment” (Burkhardt and Smith 1991, 423).
5. It is interesting to note that Whewell’s response to the Origin was much kinder
than Herschel’s. In a letter to Darwin (January 3, 1860) he wrote “I have to thank you for
a copy of your book on the ‘Origin of Species.’ You will easily believe that it has interested
me very much, and probably you will not be surprised to be told that I cannot, yet at least,
become a convert to your doctrines. But there is so much of thought and fact in what you
have written that it is not to be contradicted without careful selection of the ground and
manner of the dissent, which I have not now time for” (Burkhardt et al. 1993, 6). In a let-
ter to Lyell written the next day (January 4) Darwin wrote “Possibly you might like to see
enclosed note from Whewell, merely as showing that he is not horrified with us” (15).
6. Or rather Darwin restricted the term to process laws. In Variation Darwin (1868 I)
defines laws as “only the ascertained sequence of events” (7). His influence here might be
from his reading (either primary or secondary) of Hume. Although there is no evidence
that Darwin ever read Hume’s Treatise or Enquiry, we do know that he had read Hume’s
Dialogues (Burkhardt and Smith 1988, 458), which contains a brief (Philo in part 2) of
Hume’s famous theory of causation as mere regularity, as well as Burton’s Life and Corre-
spondence of David Hume (472, 479). At any rate, Darwin’s expansion (or restriction) of laws
to include process laws such as natural selection might help to explain Herschel’s rejection
of natural selection as “the law of higgledy-piggledy” in note 4 above.
7. Darwin, interestingly, often inveighed against splitters. Since varieties on his view
are incipient species and the history of life is treelike (Darwin, again, was a pluralist with
regard to speciation), he required what today is called a “polytypic” conception of species
(species with varieties or subspecies). As he wrote to Jenyns (April 28, 1858), “One chief
reason why I have not accumulated more facts of variation in state of nature is, that natu-
ralists so invariably turn around & say oh they are not varieties, but species” (Burkhardt and
Smith 1991, 86). Little wonder, then, that Darwin corresponded so much with lumpers,
notably Hooker, Watson, and Gray. For an interesting discussion on the diverse reasons and
motivations for Hooker as a lumper, cf. Stevens (1997).
8. Application of the vera causa ideal to species classification was not without prece-
dent. For a detailed discussion on, for example, A.P. de Candolle’s view on botany, natural
classification, and the laws of crystallography, cf. Stevens (1984).
9. Interestingly, when Maw in a letter to Darwin (March 15, 1861) brought out the
analogy between mineral classification and organism classification and the problem of su-
perficial similarity between different forms “which look most singularly like genealogical
affinity” (Burkhardt et al. 1994, 56), Darwin replied (March 17) “I have been particularly
struck with your observations on the classification of mineral bodies. The idea has crossed
my mind in a very vague & feeble manner: it is so difficult to be honest that I fear that I
must have unconsciously banished the idea as disagreeable. I see now the full force of the
difficulty & I will make a note not to forget this subject. I must own that classification may
be closely like that due to descent & yet have no relation to it” (56–57). Beginning with the
fourth edition of the Origin Darwin included references to the analogy between mineral
and biological classification. In the sixth edition (1876c), for example, on the problem of
convergence, he wrote “The shape of a crystal is determined solely by the molecular forces,
and it is not surprising that dissimilar substances should sometimes assume the same form;
 Notes to Chapter 6 237

but with organic beings we should bear in mind that the form of each depends on an
infinitude of complex relations, . . . It is incredible that the descendants of two organisms,
which had originally differed in a marked manner, should ever afterwards converge so closely
as to lead to a near approach to identity throughout their whole organization. If this had oc-
curred, we should meet with the same form, independently of genetic connection, recurring
in widely separated geological formations; and the balance of evidence is opposed to any such
admission” (100–101). Again he wrote “We know, for instance, that minerals and the ele-
mental substances can be thus arranged [‘either artificially by single characters or more nat-
urally by a number of characters’]. In this case there is of course no relation to genealogical
succession, and no cause can at present be assigned for their falling into groups. But with
organic beings the case is different, and the view given above accords with their natural
arrangement in group under group; and no other explanation has ever been attempted”
(364). Cf. also Descent (1871 I, 231) for the same point about convergence and the dis-
analogy between mineral and organism classification.

CHAPTER 6
1. To the modern mind, with its emphasis on the genetic basis of sterility, it is perhaps
difficult to believe that Darwin would have believed that mere domestication could eventu-
ally eliminate such a physical character as sterility between species. Given Darwin’s claim in
Notebook C (161) that “My definition of species has nothing to do with hybridity, is simply,
an instinctive impulse to keep separate, which no doubt be overcome, but until it is the ani-
mals are distinct species” (Barrett et al. 1987, 289), combined with his view in the Origin that
“Natural instincts are lost under domestication” (215), it might be natural to suppose that
according to Darwin the domestic breeds of dogs are interfertile because their domestication
eliminated the instinct to keep separate between the interfertile wild species from which they
were derived. Although a plausible theory on the surface, it nevertheless does not express Dar-
win’s view. Darwin makes this clear enough in the Origin, at the end of his discussion on
domestic pigs and cattle, in which he wrote “we must look at sterility, not as an indelible char-
acteristic, but as one capable of being removed by domestication” (254).
2. This interpretation finds further corroboration in a letter from Darwin to John
Gould, the ornithologist who classified the bird specimens sent to him from Darwin’s Bea-
gle voyage. In 1861 Gould sent to Darwin a copy of his book An Introduction to the
Trochilidæ, or Family of Humming-Birds published earlier that year. In that book Gould re-
ferred to three species of hummingbirds found on the island of Juan Fernandez and two
species of hummingbirds found on the island of Jamaica, stating in each case that no cross-
ing or interbreeding had ever been observed between the species. In a letter to Gould (Oc-
tober 6, 1861), Darwin wrote “I see you allude to the crossing of Birds in a state of nature;
I, for one, repudiate this notion” (Burkhardt et al. 1994, 295). Gould evidently employed
in his species concept reproductive isolation, and Darwin here, quite consistently with every-
thing we have seen and shall further see in this chapter, evidently rejects that concept.
3. Even as early as his Sketch of 1842 (Darwin 1909), Darwin argued that “sterility,
though a usual, is not an invariable concomitant” (50), to which he adduced evidence from
Kölreuter, who we know from Darwin’s reading notebooks he had read in January of 1840
(Burkhardt and Smith 1988, 461).
238 Notes to Chapter 6

4. Interestingly, during 1862 Darwin toyed with, and possibly even held, the idea
that interspecific sterility can be naturally selected. His strongest statement to this effect is
found in a letter to Hooker (December 12), in which he wrote “By the way my notions on
hybridity are becoming considerably altered by my dimorphic work: I am now inclined to
believe that sterility is at first a selected quality to keep incipient species distinct. If you have
looked at Lythrum, you will see how pollen can be modified merely to favour crossing; with
equal readiness it could be modified to prevent crossing” (Burkhardt et al. 1997, 598). Dar-
win quickly gave up this idea, however, and never toyed with it again. Kottler (1985,
402–405) claims that Darwin reverted to his former view the more he studied Lythrum sali-
caria, publishing his results in 1865. Burkhardt et al. (1997, 700–711), however, argue that
Darwin’s Lythrum work was part of a wider rejection of selection for sterility, beginning
with a plan for breeding experiments on pigeons for the purpose of artificially selecting
sterility. In late December 1862 Darwin suggested to his friend, the pigeon fancier Teget-
meier, that he perform the necessary experiments, which Tegetmeier did with negative re-
sults (cf. Variation 1868 I, 200). Elsewhere in Variation (1868 II, 169–172) Darwin gives
three reasons for why interspecific sterility could not be selected for, the first being that in-
terspecific sterility could not possibly be selected for in geographically isolated varieties, the
second being that selection could not possibly explain the many cases of asymmetry in the
sterility/fertility of reciprocal crosses, the third being that it could be of no advantage to in-
dividuals of one variety to have reduced fertility with individuals of another variety and
thus leave fewer offspring. Burkhardt et al. (1997) argue that Darwin wrote the first draft
of this section of Variation between April and June of 1863 and that “there is no evidence
to suggest that the first draft differed from the published version in this regard” (704).
5. It is interesting to compare what Darwin says in the Origin on primroses and
cowslips with what he says in his 1869 paper. As we have seen in chapter 5, Darwin argued
that primroses and cowslips should be classified as two species instead of one and that it is
very doubtful that the intermediate links are hybrids. In his later paper on the subject, Dar-
win (1869) gives essentially the same reasons for classifying primroses and cowslips as dif-
ferent species, but he argues instead that the common oxlip is indeed a hybrid between the
two. Nevertheless Darwin nowhere in his paper calls the common oxlip a “species” (which
if he did it would create a serious problem for my thesis in chapter 5). Ghiselin, however,
is highly misleading in the passage that I quoted at length earlier in the present chapter. He
implies that the “third species” is the common oxlip. But it is not. The three species Dar-
win there refers to are not the primrose, cowslip, and common oxlip, but rather the prim-
rose (Primula vulgaris), cowslip (P. veris), and Bardfield oxlip (P. elatior), the latter which
Darwin argues (449–451), much like the former two, is not a hybrid but should be consid-
ered a good and true species in its own right.
6. Kottler does indeed seem to think that Darwin’s mature species concept is closer
to Mayr’s than anyone else’s (Kottler 1978, 280 n. 9, 292, 297).
7. Of further relevance and interest is what Darwin has to say only a few pages earlier
on the origin of “permanent varieties”: “The formation of a permanent variety, implies not
only that the modifications are inherited, but that they are not disadvantageous, generally
that they are in some degree advantageous to the variety, otherwise it could not compete
with its parent when inhabiting the same area. The formation of a permanent variety must
be effected by natural selection” (Stauffer 1975, 240). To this we should recall to mind
Darwin’s characterization of species in the Origin, as “only well-marked and permanent
 Notes to Chapter 7 239

varieties” (133), or in other words “only strongly marked and fixed varieties” (155), as well
as his claim that varieties are “incipient species” (111).
8. Similar though much abbreviated statements can be found in the Origin
(101–106).
9. Darwin’s passing references in his review paper to examples of preferential mat-
ing are apparently taken from his much more careful discussion in Natural Selection (Stauf-
fer 1975), in which he gives thirteen examples. Especially significant in his discussion is the
example of two forms of Caribou deer, distinguished primarily by different kinds of ranges
and migration patterns. Darwin quotes a source as saying that they constitute “two well-
marked & permanent varieties” (258).
10. The same point recurs in Darwin’s reply to Watson on the topic of convergence
at the species level and in his subsequent writings on the topic, as examined in chapter 4.
Of further interest is that one can read in Darwin a distinction between adaptations in terms
of function and adaptations in terms of structure. In the Origin Darwin gives the example
of the upland goose, which although it has webbed feet it rarely or never goes near the water,
so that “habits have changed without a corresponding change of structure. The webbed feet
of the upland goose may be said to have become rudimentary in function, though not in
structure” (185). Such structures are nevertheless still adaptations in terms of structure, and
are not to be confused with rudimentary organs or structures in the strict sense, which are
no longer adaptations in either function or structure because they are atrophied (cf. 416,
450–456, 480, 483).

CHAPTER 7
1. The same might be said of other responses to Darwin’s Origin. Masters (1860),
for example, while maintaining “the existence of ‘species,’ endowed with a very variable,
but a limited power of variation” (224), also maintained that varieties are “confessedly ar-
tificial distinctions” (226). Less explicitly, Hopkins (1860) claimed the same when he ac-
cused Darwin of confounding “artificial with natural species” (78)—the latter being “formed
by nature,” the former being manmade and “arbitrary” (747)—in that for Hopkins natural
species have a clear distinction from varieties but artificial species do not. Sedgwick (1860)
also implied that varieties in nature are not real. According to Sedgwick, “We all admit the
varieties, and the very wide limits of variation, among domestic animals,” but “the varieties,
built upon by Mr. Darwin, are varieties of domestication and human design. Such varieties
could have no existence in the old world” (285). What I shall argue in subsequent pages,
however, is that what we see here in the examples of Agassiz, Masters, Hopkins, and Sedg-
wick is probably not really a rearguard action in response to the Origin, but rather the mak-
ing explicit, caused by the Origin, of a view that had been generally accepted by naturalists
prior to the Origin but that had remained pretty much implicit until that time.
2. Winsor (1979, 104–111) takes issue with Mayr’s claim that Agassiz’s variety nom-
inalism “forced” him to split species when there was a lack of intergrading. Instead she claims
that prior to the Origin, in his Essay, Agassiz allowed for the existence of subcategories, in-
cluding varieties. In all of this, however, her argument, including the example of three- and
four-toed box turtles (99–102), as well as the single passage in the Essay for subcategories that
240 Notes to Chapter 8

she provides as direct evidence (101), does not seem at all to me convincing. (The subcategories
briefly discussed by Agassiz are subclasses, suborders, subfamilies, subgenera, and varieties.) The
passage she quotes as direct evidence is the following: “The individuals of a species, occupy-
ing distinct fields of its natural geographical area, may differ somewhat from one another, and
constitute varieties, etc.” But it is highly unlikely that Agassiz would have thought that sub-
categories, including varieties, also belong to God’s “mode of thinking.” Otherwise what was
there to stop the next step of God thinking in terms of evolution? Revealing on this matter is
one of Agassiz’s early footnotes in his Essay, in which he states “It must not be overlooked
here that a system may be natural, that is, may agree in every respect with the facts in nature,
and yet not be considered by its author as the manifestation of the thoughts of a Creator, but
merely as the expression of a fact existing in nature—no matter how—which the human mind
may trace and reproduce in a systematic form of its own invention” (8–9 n. 7). This footnote
needs to be related to the passage in the Essay, which Winsor quotes from, in which Agassiz
discusses subcategories. Agassiz specifically states that although they “must be acknowledged
in a natural zoological system,” they are a matter of “propriety” used “to express all the vari-
ous degrees of affinity of the different members of any higher natural group” (180), such that,
on Agassiz’s view, until an objective principle is discovered for the limits of these subdivi-
sions—and he doubts that this will ever happen because they are based on “degrees”—they
“must be left to arbitrary estimations,” and he hopes that “such arbitrary estimations are for-
ever removed from our science, as far as the categories themselves are concerned” (181). Given
all of this, it seems highly doubtful that Agassiz changed his expression on the objective exis-
tence of varieties following the publication of the Origin. He seems never to have believed in
their objective existence. Instead his only change was a change in emphasis.
3. According to Hodge (1987), Linnaeus is largely to be credited for laying the foun-
dation for “the historical and teleological conception of ‘permanent varieties’ that has been
developed by Prichard’s time” (232), but what we have seen already from Linnaeus above
undermines to my mind that assessment.
4. This letter may well be the source for an interesting passage in Natural Selection, in
which Darwin wrote “Dr. Hooker objects to my whole manner of treating the present sub-
ject because varieties are so ill defined; had he added that species were likewise ill defined,
I should have entirely agreed with him; for my belief is that both are liable to this imputa-
tion; varieties more than closely allied species, & these more than strongly marked species”
(Stauffer 1975, 159–160). We shall see the significance of this further below.

CHAPTER 8
1. For a discussion on Aristotle’s essentialism, with his distinction between genos and
eidos (which is not to be confused with the modern taxonomic distinction between genus
and species), cf. Stamos (2003, 102–111) and Grene and Depew (2004, 10–34). Ghiselin’s
explicit source for his understanding of Aristotelian essentialism and categories is Hull
(1965), who himself did not use primary but only secondary sources.
2. This is indicated by the fact that it has been reprinted in an anthology devoted to
the species problem, namely, Ereshefsky (1992a), in the direct following that its line of in-
terpretation has gained (e.g., Bowler 1988, 69; Stevens 1992, 305; Kitcher 1993, 32 n. 45;
 Notes to Chapter 8 241

de Queiroz 1999, 83 n. 24; Grene and Depew 2004, 213 n. 20; LaPorte 2004, 192 n. 13),
and in the slight twists that have been added to it by others (e.g., Hodge 1987, 248–249;
McOuat 1996, 475, 514–515).
3. This theory isn’t entirely original. Ellegård (1958), for example, claimed that
“Varietal distinctions were accidental and fluctuating, while specific ones were essential and
permanent. Thus the immutability doctrine was, as it were, guaranteed by the current de-
finition of species, and this was one of the first obstacles that Darwin had to remove in
order to prepare the way for his transmutation theory” (198). A few years later Simpson
(1961) claimed that “What he [Darwin] did was to take as given the classifications then
current and to show, first, that they were consistent with the theory that their taxa originated
by evolution, and second, that evolutionary phylogeny could explain the order that had
already been found among organisms” (53).
4. To all of this one must keep in mind that Darwin began his transmutation note-
books in July 1837 (Barrett et al. 1987, 167; cf. Darwin 1876a, 83).
5. “With species in a state of nature, every naturalist has in fact brought descent into
his classification; for he includes in his lowest grade, or that of a species, the two sexes; and
how enormously these sometimes differ . . . yet no one dreams of separating them. The nat-
uralist includes as one species the several larval stages of the same individual, however much
they may differ from each other and from the adult; . . . He includes monsters; he includes
varieties, not solely because they closely resemble the parent-form, but because they are
descended from it” (Darwin 1859, 424).
6. “Several eminent naturalists have of late published their belief that a multitude of
reputed species in each genus are not real species; but that other species are real, that is,
have been independently created” (Darwin 1859, 482).
7. “Hence I believe a well-marked variety may be justly called an incipient species;
but whether this belief be justifiable must be judged of by the general weight of the several
facts and views given throughout this work” (Darwin 1859, 52).
8. Hooker let Darwin see this letter to him. Interestingly, in response to Lyell’s fear
quoted above, Darwin wrote to Hooker (July 30, 1856) “I differ from him greatly in think-
ing that those who believe that species are not fixed will multiply specific names: I know in
my own case my most frequent source of doubt was whether others would not think this
or that was a God-created Barnacle & surely deserved a name. Otherwise I shd. only have
thought whether the amount of difference & permanence was sufficient to justify a name”
(Burkhardt and Smith 1990, 194).
9. Burkhardt et al. (1993) point out that, according to Francis Darwin, his father was
in the habit of periodically burning accumulated letters, such that “This process, carried on
for years, destroyed nearly all letters received before 1862” (110). This would explain why
in Darwin’s extant correspondence prior to 1862 more of the letters are from Darwin than
to Darwin.
10. The closest reference that one can find is Hooker’s letter to Darwin (March 14,
1858), in which Hooker wrote “The long & short of the matter is, that Botanists do not at-
tach that definite importance to varieties that you suppose” (Burkhardt and Smith 1991, 49).
11. According to Burkhardt and Smith (1989), as of mid-1855 “both Joseph
Dalton Hooker and Louis Agassiz regarded species to be fixed, but Hooker thought all
242 Notes to Chapter 8

species originated from a single parent or pair whereas Agassiz believed in multiple
creation” (364 n. 6).
12. Indeed it was typical of Darwin to mean by “given up species” no more than the
rejection of the belief that species are immutable, that is, that one species can evolve into an-
other. For example, in his reading notebooks Darwin wrote of Watson (1845a) that he
“gives up permanent species” (Burkhart and Smith 1988, 449). We have already seen in
chapter 1 that even though Watson was an early evolutionist he never believed that species
are not real. Moreover Darwin never believed him to have taken such a nominalist position.
13. This is in spite of having written in the Origin, after listing a number of eminent
paleontologists and geologists who do not doubt that species are immutable, that “I have rea-
son to believe that one great authority, Sir Charles Lyell, from further reflexion entertains
grave doubts on this subject” (310).
14. An interesting and contemporaneous use of the word “entity” is to be found in
Bentham (1858): “He [Linnaeus] accordingly, treating his genera as entities (to use a word
of Jeremy Bentham’s) as natural as species, distributed them for practical purposes into his
well-known artificial Classes and Orders” (30). The connotation of realism in this passage
is pretty clear, so that a nonentity is not real. Darwin’s use of the term “entity” in the pas-
sage above, then, might seem to imply species nominalism, but it doesn’t necessarily. Dar-
win’s use of the term “entity” only applies to the sterility criterion of species distinctness,
for Darwin immediately continues in the above passage to say “Is it not that the dog case
injures the argument from fertility; so that one main argument that the races of man are
varieties & not species, i.e. because they are fertile inter se is much weakened?” (397).
15. This is a prime example of the kind of reasoning that has become popular in pro-
fessional history of science, which I take on in chapter 10. The present case, however, de-
serves a few comments here. First, no reference is given for the “no jot of ‘demonstrative
evidence in its favour’” line, which is given to imply that Huxley converted for non-eviden-
tial reasons, and which is then contradicted by the reference to Darwin on pigeon breeds,
a matter of evidence that Darwin surely would have raised in his discussions with Huxley
and others at his home. Second, Desmond (675 n. 24) refers to Lyell’s journal on the species
question (Wilson 1970) as historical support. If we look at what Lyell actually wrote, how-
ever, all he wrote in his journal (dated April 29, 1856) is that “After conversation with Mill,
Huxley, Hooker, Carpenter & Busk at Philos. Club, conclude that the belief in species as
permanent, fixed & invariable, & as comprehending individuals descending from single
pairs or protoplasts is growing fainter—no very clear creed to substitute” (56–57). This is
hardly good evidence for an “about-face” on Huxley’s or anyone else’s part. Third and most
importantly, Huxley had been a longtime friend of Herbert Spencer, the evolutionary pro-
gressivist par excellence. Had Huxley’s main motive for converting to evolutionism been po-
litical, the main catalyst should have been Spencer, not Darwin, and it should have occurred
much sooner. A much more plausible explanation, then, would be that it was scientific ev-
idence and argument, raised mainly by Darwin and lacking in Spencer, which got Huxley
turned around, evidence and argument that took a couple of years for him to digest and that
he subsequently used in his campaign to professionalize science. To this it is interesting to
add Huxley’s own reflections on the matter recorded many years later. Noting that he first
met Spencer in 1852 and that shortly afterward they had become friends, he adds that
“Many and prolonged were the battles we fought on this topic [evolution]. But even my
friend’s rare dialectic skill and copiousness of apt illustration could not drive me from my
 Notes to Chapter 9 243

agnostic position. I took my stand upon two grounds: firstly, that up to that time, the
evidence in favour of transmutation was wholly insufficient; and, secondly, that no sugges-
tion respecting the causes of the transmutation assumed, which had been made, was in any
way adequate to explain the phenomena. Looking back at the state of knowledge at that
time, I really do not see that any other conclusion was justifiable” (Darwin 1892, 178). In
what I call “the new historiography,” all of this means nothing.
16. Cf. the correspondence between Darwin and Huxley (late September 1857) on the
importance of genealogy versus structure in classification (Burkhardt and Smith 1990, 456,
461–462).
17. It might seem odd that Darwin would read Wallace (1855) as a creationist, and
continued to do so for some time. For one thing, Lyell himself never did, or did so only hes-
itatingly. In his index book to his first journal on the species question, begun on November
28, 1855, Wallace’s article is the first topic of writing, and Lyell wrote that Wallace “goes far
towards Lamarck’s doctrine” (Wilson 1970, 66). During a visit to Darwin’s home from April
13–16, 1856, Lyell apparently recommended Wallace’s paper for Darwin to read and warned
him that he might be forestalled (Burkhardt and Smith 1991, 108 n. 4). What may have con-
tributed to Darwin’s reading of Wallace (1855) as a creationist, in spite of Lyell, is that Wal-
lace repeatedly used the word “creation” (188, 193), he repeatedly referred to the closely-allied
ancestral species in his “law” as the “anti-type” (186, 187, 188, 191, 192), and even though
he used the tree metaphor (187, 191) he thought that “We have no reason for believing that
the number of species on the earth at any former period was much less than at present” (194).
Thus even though Wallace used the tree metaphor, argued for a kind of gradualism in change
between species (191, 195–196), and argued for the importance and significance of rudi-
mentary organs (195), Darwin read Wallace as a successive creationist, which indeed at the
time was nothing new. Many, such as Lyell and Agassiz, had combined the natura non facit
saltum creationism of Linnaeus with the idea of many successive creations.

CHAPTER 9
1. For his maintenance of “generally,” cf. Kuhn (1970, 181) and Hoyningen-Huene
(1993, 232–233).
2. Cf., for example, Lakatos (1970, 93) and Chalmers (1999, 115, 123–124) for a
reading of Kuhn as meaning religious conversion, and Hoyningen-Huene (1993, 257–258,
239–252) for a disavowal that religious conversion was what Kuhn had meant.
3. Kuhn was quite apparently an instrumentalist, as, for example, when he (Kuhn
1970) compares scientific theories to tools and says “As in manufacture so in science—
retooling is an extravagance reserved for the occasion that demands it. The significance of
crisis is the indication they provide that an occasion for retooling has arrived” (76).
4. On the topic of age as a factor, Hull et al. (1978), restricting themselves to the Dar-
winian revolution, claim that “age explains less than ten percent of the variation in accep-
tance,” so that “the connection between age and acceptance is not as important as people
such as Max Planck have claimed” (58). Sulloway (1996) calls age “a reasonably good
predictor of attitudes toward scientific innovation” (36), but adds the qualification that
244 Notes to Chapter 10

with firstborns age does not seem to play much of a factor, whereas with laterborns it
“exerts a substantially greater influence on attitudes toward evolution” (35). Darwin, for
what it’s worth, thought that age would be a large factor. In a passage surprisingly quoted
with approval by Kuhn (1970, 151), Darwin wrote at the end of the Origin that he looked
mainly to “young and rising naturalists” to transform creationist biology into evolutionary
biology, since he thought they “would be able to view both sides with impartiality” (482).
This was a theme that Darwin would repeat throughout his correspondence (cf. Burkhardt
and Smith 1991, 279, 404, 431; Burkhardt et al. 1993, 115, 507, 514; Burkhardt et al.
1994, 45, 102, 135).
5. Originally Kripke and Putnam argued that what unites samples of a natural kind is
an underlying structural essence, whether chemical kinds such as gold (the essence being pro-
ton number 79) or biological kinds including species such as tiger (the essence being in the
chromosome structure). Whether the causal theory of reference is necessarily committed to
essentialism for natural kinds (which of course is utterly implausible for biological species,
since variation is the norm) is discussed and rejected, for example, by Hacking (1983, 82).
6. As Putnam (1973) put it, “concepts which are not strictly true of anything may yet
refer to something; and concepts in different theories may refer to the same thing . . . real-
ists have held that there are successive scientific theories about the same things: about heat,
about electricity, about electrons, and so forth; and this involves treating such terms as ‘elec-
tricity’ as trans-theoretical terms” (197).
7. Again as Putnam (1973) put it, “Scientists . . . are trying to maximize truth (or im-
prove their approximation to truth, or increase the amount of approximate-truth they know
without decreasing the goodness of the approximation, and so forth)” (212).
8. Beatty in his writings about Darwin on species makes no explicit reference to the
causal theory of meaning; indeed he coyly hides his debt to it, claiming that his discussion is
in terms of nineteenth century thought. Nevertheless one clue to his debt is that he approv-
ingly refers to Kitcher’s (1978) discussion on “recent developments in semantics” and claims
that Darwin did indeed recognize and take advantage of the distinction between meaning
as reference and meaning as definition (cf. Beatty 1982, 221–222 n. 6, 1985, 280–281 n. 3).
9. Bowler (2000) claims that Darwin’s conversion was “like a gestalt-shift” (95), that Dar-
win “jumped suddenly and wholeheartedly” (97), but this seems to me not only to ignore the
evidence to the contrary but to be the result of a confusion. Granted, in his autobiography
Darwin (1876a) states that the theory of natural selection “at once struck” him (120) in Octo-
ber 1838 after reading Malthus, and also that he can remember “the very spot in the road, whilst
in my carriage” (120) at which the principle of divergence “occurred” to him many years later,
but it does not follow that his conversion to these theories was saltational rather than gradual. For
one, he also states that it was not until “about 1839” that “the theory [of evolution by natural
selection] was clearly conceived” (124). For another, his principle of divergence was a theory
that he subsequently went to great lengths to test (as we shall see in the next chapter).

CHAPTER 10
1. This is, ironically and paradoxically, an epistemological premise, a statement about
reality, specifically people including scientists, which is claimed to be true, and yet it is
 Notes to Chapter 10 245

thoroughly immune to testing—since by its own principles no amount of evidence could

logically refute it!
2. The Copernican revolution, it should be noted, did not follow immediately after
the publication of Copernicus’ De Revolutionibus, but instead took roughly 150 years to be
completed (Kuhn 1957).
3. It is interesting that Darwin, a few years after the publication of the Origin, claimed
that the most important change was to the belief in species evolution, not to natural selec-
tion, although he did think that the latter would eventually be accepted. In a letter to
Hooker (January 13, 1863), for example, he wrote “as I look at it, the great gain is for any
good man to give up immutability of species: the road is then open for progress; it is com-
paratively immaterial whether he believes in N. Selection; but how any man can persuade
himself that species change unless he sees how they become adapted to their conditions is
to me incomprehensible” (Burkhardt et al. 1999, 36). Similarly in a letter to A. de Candolle
(January 14, 1863) he wrote “But the great point, as it seems to me, is to give up the im-
mutability of specific forms; as long as they are thought immutable, there can be no real
progress in ‘epiontology’ [the attempt to unify biogeography and phylogeny]” (40). One can
also find the same basic claim in one of his articles (Darwin 1863b), in which he wrote
“Whether the naturalist believes in the views given by Lamarck, or Geoffroy St.-Hilaire, by
the author of the ‘Vestiges,’ by Mr. Wallace and myself, or in any other such view, signifies
extremely little in comparison with the admission that species have descended from other
species and have not been created immutable; for he who admits this as a great truth has a
wide field opened to him for further inquiry. I believe, however, from what I see of the
progress of opinion on the Continent, and in this country, that the theory of Natural
Selection will ultimately be adopted, with, no doubt, many subordinate modifications and
improvements” (Barrett 1977 II, 81). It should be added that Darwin never gave up his be-
lief in the importance of adaptation and natural selection, as indicated by what he wrote in
his autobiography (1876a). On the topic of adaptation, he wrote “I had always been much
struck by such adaptations, and until these could be explained it seemed to me almost use-
less to endeavour to prove by indirect evidence that species have been modified” (199). This
needs to be compared with what he wrote on the topic of natural selection: “The old argu-
ment of design in nature, as given by Paley, which formerly seemed to me so conclusive, fails,
now that the law of Natural Selection has been discovered” (87).
4. To be specific, Agassiz confined recapitulation to each of Cuvier’s embranchements
(cf. Ospovat 1981, 135).
5. Darwin’s correspondence is often useful for finding clarifications and emphases
perhaps missing in his published work. For example, in a letter to Lyell (March 12, 1860)
he wrote that his view of progress depends “on the conditions” (Burkhardt et al. 1993, 128),
while in a letter to Harvey (September 20–24, 1860) he wrote “I consider Natural Selection
of such high importance, because it accumulates successive variations in any profitable
direction; & thus adapts each new being to its complex conditions of life” (371, italics
mine). Equally important, in a letter to Falconer (October 1, 1862) he wrote “This [adap-
tation] seemed to me and does still seem the problem to solve, and I think natural selection
solves it, as artificial selection solves the adaptation of domestic races for man’s use. But I
suspect that you mean something further,—that there is some unknown law of evolution
by which species necessarily change; and if this be so, I cannot agree” (Burkhardt et al. 1997,
441). Similarly he wrote in a letter to Scott (March 6, 1863) “I cannot help doubting from
246 Notes to Chapter 10

your expression of an ‘innate . . . . . selective principle’ whether you fully comprehend what
is meant by Natural Selection” (Burkhardt et al. 1999, 213). Cf. also Natural Selection
(Stauffer 1975, 238, 271, 273), Variation (1868 II, 425–428), and Darwin’s autobiography
(1876a, 87).
6. Sulloway (1996) calls the historiography of Desmond and Moore “Marxist” (240).
Moreover, as a result of his enormous statistical analysis that he conducted over many years,
Sulloway argues, against Desmond (1989), that “the correlation between social class and sup-
port for evolution is almost zero” (237), so that “The idea that social class explains something
about the Darwinian revolution is an illusion” (240). He argues further that Desmond’s analy-
sis of social class and acceptance of evolution “is the product of biased sampling that con-
firmed his working hypothesis” (239). I would argue that bias is precisely what characterizes
Desmond and Moore (1992). Indeed a few years later, looking back at the co-authoring,
Moore (1996) confides that “our historiographic aim was shared: to embed Darwin the man,
his practices and theories, in a shifting social order” (275), a historiography that sees “progress”
in “the degree to which the pure waters of Darwin’s science are muddied by the rich surround-
ing soil of political economy, natural theology, urban radicalism and provincial Dissent” (271).
Mud is exactly what we find (cf. Moore 1997 who does the same for Wallace).
7. Referring to Darwin’s seed experiments on Lythrum salicaria (1865), in which, as we
have seen in chapter 6, Darwin found sterility in homomorphic though not heteromorphic
unions, in what he called “illegitimate” as opposed to “legitimate unions” (120), Desmond
and Moore (1992) remark that “Sterile seed counts somehow fitted an unromantic, data-
crunching age” (520). Although they obviously prefer to substitute Darwin’s word “unions”
above with “marriages,” they nevertheless miss the point about Darwin’s species, which is that
they undermined the Victorian value of family, with lots of children gathered round the
hearth. Indeed, as Secord (1989) points out, it was partly because Chambers in his Vestiges
(1844) played into the hands of this Victorian domestic virtue—with his generative model
of parental and offspring species, such that “the birth of a new species was no more or less to
be feared than the birth of a child” (184)—that his book became a Victorian best-seller.
8. It is to be noted that this was indeed part of Darwin’s family heritage. Desmond
and Moore (1992) refer to the “bastardizing experiments” (520) of his grandfather, Eras-
mus Darwin (an early evolutionist, it should be added, famous author of Zoönomia), while
Browne (1995) points out that “Erasmus Darwin had a passionate and earthy nature, at
one with the liberal sexual views of eighteenth-century Paris” (41). Charles Darwin’s species,
then, rather than being good Victorian species, were good Parisian species! All the more odd
that the Origin never took off in France.
9. Or rather he was one of the first, since the nod for the basic idea should probably
go to Patrick Matthew, who published his theory in the appendix to his On Naval Timber
and Arboriculture (cf. Burkhardt et al. 1993, 584–589), published in 1831. And yet credit
must still go to Darwin for being the first in history to give a scientific explanation of adap-
tations by natural selection, which included not only a wealth of evidence but also recogni-
tion and explanation of co-adaptations, such as moths and orchids. Striking is his prediction
concerning the Madagascar orchid Angraecum sesquipedale, with its nectar receptacle “eleven
and a half inches long, with only the lower inch and a half filled with nectar” (Darwin
1862a, 162). Darwin predicted that, in order to harvest the nectar, “there must be moths
with proboscides capable of extension to a length of between ten and eleven inches!” (163).
The discovery of the moth was made 40 years later in Madagascar, long after Darwin was
 Notes to Chapter 10 247

dead (cf. Browne 2002, 178; interestingly the prediction and discovery are omitted from
Desmond and Moore 1992).
10. The immediate ancestor of Winsor’s article is Camardi (2001), between which
there is much overlap, notably the rejection of the interpretations of essentialism by Hull
and Mayr, and the acceptance of the clusterism of Whewell and Boyd. But Camardi pre-
sents a different angle than Winsor. Based mainly on his study of the work of von Baer and
Owen, he states that “if we read in sequence von Baer, Owen and Darwin, we understand
that the Darwinian concept of evolution was not born full-fledged merely from Darwin’s
own speculation. Rather, it is a result of the long historical accumulation on such concepts
as development, generation, reproduction, a work that involved both Owen and Darwin,
among others” (500). Here again we see an example of the new historiography, which sees
Darwin’s theories as the result of a “long historical accumulation,” and in which we find
phrases such as “must have” (495), “should be” (496), and “sounds like” (509) used to con-
vince us that von Baer and Owen were already testing the evolutionary waters prior to Dar-
win’s coming out, an interpretation that is stretched at best. On the other hand, there is
something in Grene and Depew’s (2004) claim that “thanks to Darwin, we think of homol-
ogy as a historical concept. It is difficult for us to think of it in pre-Darwinian, non-histor-
ical terms” (215). Of course it is easy for us (and professional historians of biology are
apparently no exception) to see in von Baer’s law of branching development in embryos, or
in Owen’s work on archetypes and homologies, theories leading to the Darwinian view.
We can just as easily see the common belief in the Great Chain of Being, whether the older
linear version, or the later branching version established by Cuvier’s theory of embranche-
ments, or the doctrine of successive creations, paving the way for Darwin. But we seriously
need to remember that, unlike for us today, it was quite easy, natural, and common for bi-
ologists at the time of Darwin and before to accept the above theories and still view nature
in static terms. This was not only because their biology was mixed with religion, with its cre-
ationism, but also because they had a belief in species essentialism and in related laws such
as the laws of reversion and limited variation. These were enormous obstacles that Darwin
needed to overcome, and there were many more. Not surprisingly, von Baer rejected evo-
lution, and pathetically little evidence of evolutionary thinking on the part of Owen exists
before 1859. Indeed most of Camardi’s evidence for Owen as a moderate evolutionist and
“forerunner of Darwin” is post-1859, which is entirely irrelevant. But such is the new his-
toriography. It goes out of its way, seemingly obsessed, to find and overemphasize anything
that takes away from Darwin’s originality and achievement, while it ignores or underempha-
sizes anything that supports it. So far, Darwin’s species concept has suffered the same fate.
11. It is pernicious not only for its treatment of scientific epistemology but also (re-
latedly) for its treatment of individual scientists. For a good example of how the new histo-
riography can be pernicious in both ways, cf. Moore (1997), where the rejection of, even
the intense dislike for, the concepts of scientific genius and scientific discovery is quite ap-
parent (cf. esp. 295, 307), and where the historiography that views the most basic ideas of
scientists as projections from society and culture is forwarded as progress (290–292).
This page intentionally left blank.
 References


All references are to reprints where indicated

Agassiz, Louis (1857). Essay on Classification. Boston: Little, Brown & Co. (1859). 2nd ed.
London: Longman, Brown, Green, Longmans and Roberts. Reprinted in Edward Lurie,
ed. (1962). Cambridge: Harvard University Press.
——— (1860a). “Minutes of Meeting of March 13.” Proceedings of the American Academy
of Arts and Sciences 4, 410.
——— (1860b). “On the Origin of Species.” American Journal of Science and Arts 30 (2nd
ser.), 142–154.
Alter, Stephen G. (1999). Darwinism and the Linguistic Image: Language, Race, and Natural
Theology in the Nineteenth Century. Baltimore: Johns Hopkins.
Andersson, Malte (1994). Sexual Selection. Princeton: Princeton University Press.
Anon. (1959). “Darwin’s Origin of Species.” The Saturday Review Dec. 24, 775–776.
Anon. (1860a). “Professor Owen on the Origin of Species.” The Saturday Review May 5,
573–574.
Anon. (1860b). “Species.” All the Year Round June 2, 174–187; July 7, 293–299.
Armstrong, D.M. (1983). What is a Law of Nature? Cambridge: Cambridge University
Press.
Arthur, J.C. (1908). “The Physiological Aspect of the Species Question.” American Naturalist
42, 243–248.
Ashton, Paul A., and Abbott, Richard J. (1992). “Multiple Origins and Genetic Diversity
in the Newly Arisen Allopolyploid Species, Senecio Cambrensis Rosser (Compositae).”
Heredity 68, 25–32.
Babinski, Edward T. (1995). Leaving the Fold: Testimonies of Former Fundamentalists.
Amherst, NY: Prometheus Books.
Barrett, Paul, ed. (1977). The Collected Papers of Charles Darwin. Two volumes. Chicago:
University of Chicago Press.

249
250 References

Barrett, Paul H., et al., eds. (1987). Charles Darwin’s Notebooks, 1836–1844. Ithaca:
Cornell University Press.
Bates, Henry Walter (1862). “Contributions to an Insect Fauna of the Amazon Valley.
Lepidoptera: Heliconidæ.” Transactions of the Linnean Society 23, 495–566.
Baum, David A., and Shaw, Kerry L. (1995). “Genealogical Perspectives on the Species
Problem.” In Peter C. Hoch and A.G. Stephenson, eds. (1995, 289–303). Experimental
and Molecular Approaches to Plant Biosystematics. St. Louis: Missouri Botanical Gardens.
Beatty, John (1982). “What’s In a Word?: Coming to Terms in the Darwinian Revolu-
tion.” Journal of the History of Biology 15, 215–239.
——— (1985). “Speaking of Species: Darwin’s Strategy.” In Kohn (1985a, 265–281).
Beer, Gillian (1989). “Darwin and the Growth of Language Theory.” In John Christie and
Sally Shuttlesworth, eds. (1989, 152–170). Nature Transfigured: Science and Literature,
1700–1900. Manchester, UK: Manchester University Press.
Bentham, George (1836). Labiatarum Genera et Species. London: James Ridgway and Sons.
——— (1858). “Memorandum on the Principles of Generic Nomenclature in Botany.”
Journal of the Proceedings of the Linnean Society (Botany) 2, 30–33.
——— (1861) “On the Species and Genera of Plants, Considered With Reference to Their
Practical Application to Systematic Botany.” Natural History Review 1, 133–151.
Bessey, Charles E. (1908). “The Taxonomic Aspect of the Species Question.” American
Naturalist 42, 218–224.
Blyth, Edward (1835). “An Attempt to Classify the ‘Varieties’ of Animals.” Magazine of
Natural History 8, 40–53.
Bowen, Francis (1860). “Darwin on the Origin of Species.” North American Review 90,
474–506.
Bowler, Peter J. (1988). The Non-Darwinian Revolution: Reinterpreting a Historical Myth.
Baltimore: Johns Hopkins.
——— (2000). “Philosophy, Instinct, Intuition: What Motivates the Scientist in Search of
a Theory?” Biology & Philosophy 15, 93–101.
Boyd, Richard (1999). “Homeostasis, Species, and Higher Taxa.” In Wilson (1999,
141–185).
Browne, Janet (1995). Charles Darwin: Voyaging. New York: Knopf.
——— (2002). Charles Darwin: The Power of Place. New York: Knopf.
Burkhardt, Frederick, and Smith, Sydney, eds. (1985). The Correspondence of Charles
Darwin, Volume 1, 1821–1836. Cambridge: Cambridge University Press.
——— (1986). The Correspondence of Charles Darwin, Volume 2, 1837–1843. Cambridge:
Cambridge University Press.
——— (1987). The Correspondence of Charles Darwin, Volume 3, 1844–1846. Cambridge:
Cambridge University Press.
——— (1988). The Correspondence of Charles Darwin, Volume 4, 1847–1850. Cambridge:
Cambridge University Press.
 References 251

——— (1989). The Correspondence of Charles Darwin, Volume 5, 1851–1855. Cambridge:

Cambridge University Press.
——— (1990). The Correspondence of Charles Darwin, Volume 6, 1856–1857. Cambridge:
Cambridge University Press.
——— (1991). The Correspondence of Charles Darwin, Volume 7, 1858–1859. Cambridge:
Cambridge University Press.
Burkhardt, Frederick, et al., eds. (1993). The Correspondence of Charles Darwin, Volume 8,
1860. Cambridge: Cambridge University Press.
——— (1994). The Correspondence of Charles Darwin, Volume 9, 1861. Cambridge:
Cambridge University Press.
——— (1997). The Correspondence of Charles Darwin, Volume 10, 1862. Cambridge:
Cambridge University Press.
——— (1999). The Correspondence of Charles Darwin, Volume 11, 1863. Cambridge:
Cambridge University Press.
——— (2001). The Correspondence of Charles Darwin, Volume 12, 1864. Cambridge:
Cambridge University Press.
——— (2002). The Correspondence of Charles Darwin, Volume 13, 1865. Cambridge:
Cambridge University Press.
Burkhardt, R.W., Jr. (1987). “Lamarck and Species.” In Roger and Fischer (1987, 161–180).
Camardi, Giovanni (2001). “Richard Owen, Morphology and Evolution.” Journal of the
History of Biology 34, 481–515.
Carpenter, William Benjamin (1860). “Darwin on the Origin of Species.” National Review
10, 188–214.
Chalmers, A.F. (1999). What Is This Thing Called Science? 3rd ed. Indianapolis: Hackett.
Chambers, Robert (1844). Vestiges of the Natural History of Creation. London: John
Churchill.
——— (1859). “Charles Darwin on the Origin of Species.” Chambers’s Journal 12,
388–391.
Claridge, M.F., Dawah, H.A., and Wilson, M.R., eds. (1997). Species: The Units of Biodi-
versity. London: Chapman & Hall.
Coleman, William (1962). “Lyell and the ‘Reality’ of Species: 1830–1833.” Isis 53,
325–338.
Coulter, J.M. (1908). “Discussion of the Species Question.” American Naturalist 42,
272–281.
Cowan, S.T. (1962). “The Microbial Species—A Macromyth?” Symposium—Society for
General Microbiology 12, 433–455.
Cowles, H.C. (1908). “An Ecological Aspect of the Conception of Species.” American
Naturalist 42, 265–271.
Cracraft, Joel (1983). “Species Concepts and Speciation Analysis.” Current Ornithology 1,
159–187. Reprinted in Ereshefsky (1992a, 93–120).
252 References

——— (1987). “Species Concepts and the Ontology of Evolution.” Biology & Philosophy
2, 329–346.
——— (1989). “Species as Entities of Biological Theory.” In Ruse (1989, 31–52).
——— (1997). “Species Concepts in Systematics and Conservation Biology—An Ornitho-
logical Viewpoint.” In Claridge et al. (1997, 325–339).
Crawford, Charles, and Krebs, Dennis L., eds. (1998). Handbook of Evolutionary Psychology:
Ideas, Issues, and Applications. Mahwah, NJ: Lawrence Erlbaum Associates.
Crawfurd, John (1859). “On the Origin of Species.” The Examiner Dec. 3, 772–773.
Creath, Richard, and Maienschein, Jane, eds. (2000). Biology and Epistemology. Cambridge:
Cambridge University Press.
Cronquist, Arthur (1978). “Again, What Is a Species?” Biosystematics in Agriculture: Beltsville
Symposia in Agricultural Research 2, 3–20.
Dana, James D. (1857). “Thoughts on Species.” Annals and Magazine of Natural History 20
(2nd ser.), 485–497.
Darwin, Charles (1839). Journal of Researches into the Geology and Natural History of the
Various Countries Visited by H.M.S. Beagle, Under the Command of Captain Fitzroy, R.N.
from 1832 to 1836. London: Henry Colburn.
——— (1845). Journal of Researches into the Natural History and Geology of the Various
Countries Visited During the Voyage of H.M.S. Beagle Round the World. 2nd ed. London:
John Murray.
——— (1851a). A Monograph on the Fossil Lepadidæ. London: The Paleontological Society.
——— (1851b). A Monograph on the Sub-Class Cirripedia, the Lepadidæ. London: The Ray
Society.
——— (1854a). A Monograph on the Fossil Balanidæ. London. The Paleontological Society.
——— (1854b). A Monograph on the Sub-Class Cirripedia, the Balanadæ. London: The Ray
Society.
——— (1858). “On the Agency of Bees in the Fertilization of Papilionaceous Flowers, and
on the Crossing of Kidney Beans.” Annals and Magazine of Natural History 2 (3rd ser.),
459– 465. Reprinted in Barrett (1977 II, 19–25).
——— (1859). On the Origin of Species by Means of Natural Selection. London: John Murray.
——— (1862a). The Various Contrivances by which Orchids are Fertilised by Insects. London:
John Murray. 2nd ed. (1877).
——— (1862b). “On the Two Forms, or Dimorphic Condition, in the Species of Prim-
ula, and on Their Remarkable Sexual Relations.” Journal of the Proceedings of the Lin-
nean Society (Botany) 6, 77–96. Reprinted in Barrett (1977 II, 45–63).
——— (1863a). “The Doctrine of Heterogeny and Modification of Species.” Athenaeum
9, 554–555. Reprinted in Barrett (1977 II, 78–80).
——— (1863b). “Origin of Species.” Athenaeum 9, 617. Reprinted in Barrett (1977 II, 81).
——— (1863c). “A Review of H.W. Bates’ Paper on ‘Mimetic Butterflies.’” Natural History
Review 3, 219–224. Reprinted in Barrett (1977 II, 87–92).
 References 253

——— (1865). “On the Sexual Relations of the Three Forms of Lythrum salicaria.” Journal
of the Proceedings of the Linnean Society (Botany) 8, 169–196. Reprinted in Barrett (1977
II, 106–131.
——— (1868). The Variation of Animals and Plants Under Domestication. Two volumes.
London: John Murray. 2nd ed. (1875).
——— (1869). “On the Specific Difference Between Primula veris, Brit. Fl. (var. acaulis,
Linn.), and P. elatior, Jacq.; and on the Hybrid Nature of the Common Oxlip. With
Supplementary Remarks on Naturally-Produced Hybrids in the Genus Verbascum.” Journal
of the Linnean Society (Botany) 10, 437–454.
——— (1871). The Descent of Man, and Selection in Relation to Sex. Two volumes. London:
John Murray.
——— (1872). The Expression of the Emotions in Man and Animals. London: John Murray.
——— (1876a). Autobiography. In Nora Barlow, ed. (1958). The Autobiography of Charles
Darwin, 1809–1882. London: Collins.
——— (1876b). The Effects of Cross and Self Fertilisation in the Vegetable Kingdom. London:
John Murray. 2nd ed. (1878).
——— (1876c). The Origin of Species. 6th ed. London: John Murray.
——— (1880). “Fertility of Hybrids from the Common and Chinese Goose.” Nature 21,
207. Reprinted in Barrett (1977 II, 219–220).
Darwin, Francis, ed. (1892). Charles Darwin: His Life Told in an Autobiographical Chapter
and in a Selected Series of his Published Letters. New York: Appleton.
———, ed. (1909). The Foundations of the Origin of Species: Two Essays Written in 1842 and
1844. Cambridge: Cambridge University Press.
Davies, Anna Morpurgo (1987). “‘Organic’ and ‘Organism’ in Franz Bopp.” In Hoenigswald
and Wiener (1987, 81–107).
Dawkins, Richard (1983). “Universal Darwinism.” In D.S. Bendall, ed. (1983, 403–425).
Evolution from Molecules to Men. Cambridge: Cambridge University Press.
——— (1986). The Blind Watchmaker. Harlow, UK: Longman Scientific & Technical.
Dawson, John William (1860). “Review of ‘Darwin on the Origin of Species by means of
Natural Selection.’” Canadian Naturalist 5, 100–120.
de Queiroz, Kevin (1999). “The General Lineage Concept of Species and the Defining
Properties of the Species Category.” In Wilson (1999, 49–89).
Dennett, Daniel (1995). Darwin’s Dangerous Idea: Evolution and the Meanings of Life. New
York: Simon & Schuster.
Desmond, Adrian (1989). The Politics of Evolution: Morphology, Medicine, and Reform in
Radical London. Chicago: University of Chicago Press.
——— (1997). Huxley: From Devil’s Disciple to Evolution’s High Priest. Reading, MA:
Addison-Wesley.
Desmond, Adrian, and Moore, James (1992). Darwin: The Life of a Tormented Evolutionist.
New York: Warner Books.
254 References

Dewey, John (1910). The Influence of Darwin on Philosophy and Other Essays. New York:
H. Holt & Co.
Di Gregario, Mario A. (1982). “The Dinosaur Connection: A Reinterpretation of T.H.
Huxley’s Evolutionary View.” Journal of the History of Biology 15, 397–418.
Di Gregario, Mario A., and Gill, Nick W., eds. (1990). Charles Darwin’s Marginalia. Volume
1. New York: Garland Publishing.
Dobzhansky, Theodosius (1937). Genetics and the Origin of Species. New York: Columbia
University Press.
——— (1973). “Nothing in Biology Makes Sense Except in the Light of Evolution.”
American Biology Teacher 35, 125–129.
Dupré, John (1981). “Natural Kinds and Biological Taxa.” The Philosophical Review 90,
66–90.
Dupree, A. Hunter (1959). Asa Gray: American Botanist, Friend of Darwin. Cambridge:
Harvard University Press.
Ehrlich, Paul R. (2002). Human Natures: Genes, Cultures, and the Human Prospect. London:
Penguin Books.
Ehrlich, Paul R., and Raven, Peter H. (1969). “Differentiation of Populations.” Science
165, 1228–1232. Reprinted in Ereshefsky (1992a, 57–67).
Eiseley, Loren (1958). Darwin’s Century: Evolution and the Men Who Discovered It. Garden
City, NY: Doubleday.
Eldredge, Niles (1985). Time Frames. Princeton: Princeton University Press.
Eldredge, Niles, and Cracraft, Joel (1980). Phylogenetic Patterns and the Evolutionary Process.
New York: Columbia University Press.
Eldredge, Niles, and Gould, Stephen Jay (1972). “Punctuated Equilibria: An Alternative to
Phyletic Gradualism.” In T.J.M. Schopf, ed. (1972, 82–115). Models in Paleobiology. San
Francisco: Cooper and Company.
Ellegård, Alvar (1958). Darwin and the General Reader: The Reception of Darwin’s Theory of
Evolution in the British Periodical Press, 1859–1872. Stockholm: Almqvist and Wiksell.
Endersby, Jim (2003). “Escaping Darwin’s Shadow.” Journal of the History of Biology 36,
385–403.
Endler, John A. (1986). Natural Selection in the Wild. Princeton: Princeton University Press.
——— (1989). “Conceptual and Other Problems in Speciation.” In Otte and Endler (1989,
625–648).
Ereshefsky, Marc, ed. (1992a). The Units of Evolution: Essays on the Nature of Species.
Cambridge: MIT Press.
——— (1992b). “Eliminative Pluralism.” Philosophy of Science 59, 671–690.
Erwin, Douglas H., and Anstey, Robert L., eds. (1995). New Approaches to Speciation in
the Fossil Record. New York. Columbia University Press.
Farber, Paul L. (1972). “Buffon and the Concept of Species.” Journal of the History of Biology
5, 259–284.
 References 255

Fawcett, Henry (1860). “A Popular Exposition of Mr. Darwin on the Origin of Species.”
Macmillan’s Magazine 3, 81–92.
Fodor, Jerry A. (1975). The Language of Thought. New York: Crowell. Reprinted (1980).
Cambridge: Harvard University Press.
Foucault, Michel (1971). The Order of Things: An Archaeology of the Human Sciences. New
York: Random House.
Fox, Anthony (1995). Linguistic Reconstruction: An Introduction to Theory and Method.
Oxford: Oxford University Press.
Freeman, Scott, and Herron, Jon C. (1998). Evolutionary Analysis. Upper Saddle River, NJ:
Prentice Hall.
Frost, Darrel L., and Wright, John W. (1988). “The Taxonomy of Uniparental Species,
with Special Reference to Parthenogenetic Cnemidophorus (Squamata: Teiidae).” Systematic
Zoology 37, 200–209.
Futuyma, Douglas J. (1986). Evolutionary Biology. 2nd ed. Sunderland, MA: Sinauer Associates.
——— (1998). Evolutionary Biology. 3rd ed. Sunderland, MA: Sinauer Associates.
Gayon, Jean (2003). “From Darwin to Today in Evolutionary Biology.” In Hodge and
Radick (2003, 240–264).
Ghiselin, Michael T. (1966). “On Psychologism in the Logic of Taxonomic Controversies.”
Systematic Zoology 15, 207–215.
——— (1969). The Triumph of the Darwinian Method. Berkeley: University of California
Press.
——— (1974). “A Radical Solution to the Species Problem.” Systematic Zoology 23,
536–544.
——— (1987). “Species Concepts, Individuality, and Objectivity.” Biology & Philosophy 2,
127–143.
——— (1989). “Individuality, History and Laws of Nature in Biology.” In Ruse (1989,
53–66).
——— (1997). Metaphysics and the Origin of Species. Albany: State University of New York
Press.
Ghiselin, Michael, and Jaffe, Linda (1973). “Phylogenetic Classification in Darwin’s Mono-
graph on the Sub-Class Cirripedia.” Systematic Zoology 22, 132–140.
Gorovitz, Samuel, et al. (1979). Philosophical Analysis: An Introduction to Its Language and
Techniques. 3rd ed. New York: Random House.
Gould, Stephen Jay (1970). “Dollo on Dollo’s Law: Irreversibility and the Status of Evolu-
tionary Laws.” Journal of the History of Biology 3, 189–212.
——— (1980). The Panda’s Thumb. New York: W.W. Norton.
——— (1981). The Mismeasure of Man. New York: W.W. Norton.
——— (1982). “Darwinism and the Expansion of Evolutionary Theory.” Science 216,
380–387. Reprinted in Michael Ruse, ed. (1989, 100–117). Philosophy of Biology. New
York: Macmillan Publishing Company.
256 References

——— (1989). Wonderful Life: The Burgess Shale and the Nature of History. London:
Hutchinson Radius.
Grant, Verne (1957). “The Plant Species in Theory and Practice.” In Mayr (1957b, 39–80).
Gray, Asa (1860a). “Review of Darwin’s Theory on the Origin of Species by means of Natural
Selection.” American Journal of Science and Arts 29 (2nd ser.), 153–184.
——— (1860b). “Darwin on the Origin of Species.” Atlantic Monthly 6, 109–116, 229–239.
——— (1860c). “Darwin and His Reviewers.” Atlantic Monthly 6, 406–425.
Grene, Marjorie, and Depew, David. (2004). The Philosophy of Biology: An Episodic History.
Cambridge: Cambridge University Press.
Gruber, Howard E. (1981). Darwin on Man: A Psychological Study of Scientific Creativity.
2nd ed. Chicago: University of Chicago Press.
Hacking, Ian (1983). Representing and Intervening: Introductory Topics in the Philosophy of
Natural Science. Cambridge: Cambridge University Press.
Hamilton, Alexander (1820). “Review of Bopp, Conjugations System.” Edinburgh Review 33, 431.
Harman, Oren Solomon (2003). “C.D. Darlington and the British and American Reaction to
Lysenko and the Soviet Conception of Science.” Journal of the History of Biology 36, 309–352.
Harris, Randy Allen (1993). The Linguistics Wars. Oxford: Oxford University Press.
Hattiangadi, J.N. (1987). How Is Language Possible?: Philosophical Reflections on the Evolu-
tion of Language and Knowledge. La Salle, IL: Open Court.
Haughton, Samuel (1860). “Biogenesis.” Natural History Review 7, 23–32.
Haynes, Robert H. (1987). “The ‘Purpose’ of Chance in Light of the Physical Basis of Evo-
lution.” In John M. Robson, ed. (1987, 1–31). Origin and Evolution of the Universe.
Kingston and Montreal, CA: McGill-Queen’s University Press.
Hennig, Willi (1966). Phylogenetic Systematics. D. Dwight Davis and Rainer Zangerl, trans.
Urbana: University of Illinois Press.
Herschel, John F.W. (1830). A Preliminary Discourse on the Study of Natural Philosophy.
London: Longman, Rees, Orme, Brown, Green, and J. Taylor.
Hodge, M.J.S. (Jon) (1977). “The Structure and Strategy of Darwin’s ‘Long Argument.’”
British Journal for the History of Science 10, 237–246.
——— (1987). “Darwin, Species and the Theory of Natural Selection.” In Roger and Fischer
(1987, 227–252).
——— (1989). “Darwin’s Theory and Darwin’s Argument.” In Ruse (1989, 163–182).
——— (2000). “Knowing about Evolution: Darwin and His Theory of Natural Selection.”
In Creath and Maienschein (2000, 27–47).
Hodge, Jonathan, and Radick, Gregory, eds. (2003). The Cambridge Companion to Darwin.
Cambridge: Cambridge University Press.
Hoenigswald, Henry M., and Wiener, Linda F., eds. (1987). Biological Metaphor and
Cladistic Classification: An Interdisciplinary Perspective. Philadelphia: University of
Pennsylvania Press.
 References 257

Hooker, Joseph Dalton (1859). “Review of the Origin of Species.” The Gardeners’ Chronicle
and Agricultural Gazette Dec. 31, 1051–1052.
Hopkins, William (1860). “Physical Theories of the Phenomena of Life.” Fraser’s Magazine
61, 739–752; 62, 74–90.
Howard, Daniel J., and Berlocher, Stewart H., eds. (1998). Endless Forms: Species and
Speciation. New York: Oxford University Press.
Howard, Jonathan (1989). Darwin. Oxford: Oxford University Press.
Hoyningen-Huene, Paul (1993). Reconstructing Scientific Revolutions: Thomas S. Kuhn’s
Philosophy of Science. Alexander T. Levine, trans. Chicago: Chicago University Press.
Hull, David L. (1965). “The Effect of Essentialism on Taxonomy: Two Thousand Years of
Stasis.” British Journal for the Philosophy of Science 15, 314–326; 16, 1–18.
——— (1978). “A Matter of Individuality.” Philosophy of Science 45, 335–360.
——— (1988). Science as a Process. Chicago: University of Chicago Press.
——— (2000). “The Professionalization of Science Studies: Cutting Some Slack.” Biology
& Philosophy 15, 61–91.
Hull, David L., Tessner, Peter D., and Diamond, Arthur M. (1978). “Planck’s Principle:
Do Younger Scientists Accept New Scientific Ideas with Greater Alacrity than Older
Scientists?” Science 202, 717–723. Reprinted in Hull (1989, 43–61). The Metaphysics of
Evolution. Albany: State University of New York Press.
Huxley, Julian S. (1912). The Individual in the Animal Kingdom. Cambridge: Cambridge
University Press.
Huxley, Thomas H. (1859a). “On the Persistent Types of Animal Life.” Proceedings of the
Royal Institution of Great Britain 1858–1862 3, 151–153.
——— (1859b). “Time and Life: Mr. Darwin’s ‘Origin of Species.’” Macmillan’s Magazine
1, 142–148.
——— (1859c). “Darwin on the Origin of Species.” The Times Dec. 26, 8.
——— (1860a). “On Species and Races, and their Origin.” Proceedings of the Royal Institu-
tion of Great Britain 1858–1862 3, 195–200.
——— (1860b). “On the Origin of Species by means of Natural Selection.” Westminster
Review 17, 541–570.
Keller, Evelyn Fox, and Lloyd, Elisabeth A., eds. (1992). Keywords in Evolutionary Biology.
Cambridge: Harvard University Press.
Kitcher, Philip (1978). “Theories, Theorists and Theoretical Change.” The Philosophical
Review 87, 519–547.
——— (1993). The Advancement of Science: Science without Legend, Objectivity without
Illusions. New York: Oxford University Press.
Kohn, David, ed. (1985a). The Darwinian Heritage. Princeton: Princeton University Press.
——— (1985b). “Darwin’s Principle of Divergence as Internal Dialogue.” In Kohn (1985a,
245–257).
258 References

Kondrashov, Alexey S., Yampolsky, Lev Yu, and Shabalina, Svetlana A. (1998). “On the
Sympatric Origin of Species by Means of Natural Selection.” In Howard and Berlocher
(1998, 90–98).
Kottler, Malcolm Jay (1978). “Charles Darwin’s Biological Species Concept and Theory of
Geographic Speciation: the Transmutation Notebooks.” Annals of Science 35, 275–297.
——— (1985). “Charles Darwin and Alfred Russel Wallace: Two Decades of Debate Over
Natural Selection.” In Kohn (1985a, 367–432).
Kuhn, Thomas S. (1957). The Copernican Revolution. Cambridge: Harvard University Press.
——— (1970). The Structure of Scientific Revolutions. 2nd ed. Chicago: University of
Chicago Press.
——— (1977). The Essential Tension: Selected Studies in Scientific Tradition and Change.
Chicago: University of Chicago Press.
——— (1993). “Afterwords.” In Paul Horwich, ed. (1993, 311–341). World Changes:
Thomas Kuhn and the Nature of Science. Cambridge: MIT Press.
Lakatos, Imre (1970). “Falsification and the Methodology of Scientific Research Pro-
grammes.” In Lakatos and Alan Musgrave, eds. (1970, 91–196). Criticism and the Growth
of Knowledge. Cambridge: Cambridge University Press.
Lamarck, Jean Baptiste de (1809). Philosophie Zoologique. Paris: Dentu. Hugh Elliot, trans.
(1914). Zoological Philosophy. New York: Macmillan.
Lambert, David M., and Spencer, Hamish G., eds. (1995). Speciation and the Recognition
Concept: Theory and Applications. Baltimore: Johns Hopkins.
LaPorte, Joseph (2004). Natural Kinds and Conceptual Change. Cambridge: Cambridge
University Press.
Laporte, Léo F. (1994). “Simpson on Species.” Journal of the History of Biology 27, 141–159.
Larson, James L. (1968). “The Species Concept of Linnaeus.” Isis 59, 291–299.
Lass, Roger (1997). Historical Linguistics and Language Change. Cambridge: Cambridge
University Press.
Leifchild, John R. (1859). “On the Origin of Species by Means of Natural Selection.” The
Athenæum Nov. 19, 659–660.
Leikola, Anto (1987). “The Development of the Species Concept in the Thinking of
Linnaeus.” In Roger and Fischer (1987, 45–59).
Levin, Donald A. (1979). “The Nature of Plant Species.” Science 204, 381–204.
——— (1983). “Polyploidy and Novelty in Flowering Plants.” American Naturalist 122, 1–25.
Lewes, George Henry (1860). “Studies in Animal Life.” Cornhill Magazine 4, 438–447.
Lightman, Bernard, ed. (1997). Victorian Science in Context. Chicago: University of Chicago Press.
Lipton, Peter. (1990). “Contrastive Explanation.” In Dudley Knowles, ed. (1990, 247–266).
Explanation and its Limits. Cambridge: Cambridge University Press.
Lovejoy, Arthur O. (1936). The Great Chain of Being: A Study of the History of an Idea.
Cambridge: Harvard University Press.
 References 259

——— (1959). “Buffon and the Problem of Species.” In Bentley Glass, et al., eds. (1959,
84–113). Forerunners of Darwin: 1745–1859. Baltimore: Johns Hopkins.
Luckow, Melissa (1995). “Species Concepts: Assumptions, Methods, and Applications.”
Systematic Botany 20, 589–605.
Lyell, Charles (1830). Principles of Geology. Volume 1. London: John Murray.
——— (1832). Principles of Geology. Volume 2. London: John Murray.
——— (1833). Principles of Geology. Volume 3. London: John Murray.
——— (1838). Elements of Geology. London: John Murray.
——— (1863). The Geological Evidences of the Antiquity of Man, with Remarks on Theories
of the Origin of Species by Variation. London: John Murray.
Lyons, Sherrie (1999). “In Search of Huxley the Scientist.” Biology & Philosophy 14,
585–591.
Mallet, James (1995). “A Species Definition for the Modern Synthesis.” Trends in Ecology
and Evolution 10, 294–299.
Masters, Maxwell T. (1860). “On the Relation between the Abnormal and Normal Forma-
tions in Plants.” Notices of the Proceedings at the Meetings of the Members of the Royal
Institution of Great Britain (1858–1862) 3, 223–227.
Mauskopf, Seymour H. (1992). “Prize Announcements.” Isis 83, 278–279.
Maynard Smith, John (1975). The Theory of Evolution. Harmondsworth, UK: Penguin
Books. Reprinted (1993). Cambridge: Cambridge University Press.
Mayr, Ernst (1942). Systematics and the Origin of Species. New York: Columbia University
Press.
——— (1957a). “Species Concepts and Definitions.” In Mayr (1957b, 1–22).
———, ed. (1957b). The Species Problem. Washington, DC: American Association for the
Advancement of Science.
——— (1970). Populations, Species, and Evolution. Cambridge: Harvard University Press.
——— (1976). Evolution and the Diversity of Life. Cambridge: Harvard University Press.
——— (1982). The Growth of Biological Thought. Cambridge: Harvard University Press.
——— (1985). “Darwin’s Five Theories of Evolution.” In Kohn (1985a, 755–772).
——— (1987). “The Ontological Status of Species: Scientific Progress and Philosophical
Terminology.” Biology & Philosophy 2, 145–166.
——— (1988). Toward a New Philosophy of Biology. Cambridge: Harvard University Press.
——— (1991). One Long Argument: Charles Darwin and the Genesis of Modern Evolutionary
Thought. Cambridge: Harvard University Press.
——— (1992). “A Local Flora and the Biological Species Concept.” American Journal of
Botany 79, 222–238.
Mayr, Ernst, and Short, Lester L. (1970). Species Taxa of North American Birds. Cambridge:
The Nuttall Ornithological Club.
260 References

McDade, Lucinda A. (1995). “Species Concepts and Problems in Practice: Insight from
Botanical Monographs.” Systematic Botany 20, 606–622.
McOuat, Gordon R. (1996). “Species, Rules and Meaning: The Politics of Language and
the Ends of Definitions in 19th Century Natural History.” Studies in History and Philosophy
of Science 27, 473–519.
——— (2000). “Networks, Hybrids and Forms of Life.” Annals of Science 57, 189–195.
——— (2001). “Cataloguing Power: Delineating ‘Competent Naturalists’ and the Meaning
of Species in the British Museum.” British Journal for the History of Science 34, 1–28.
Mishler, Brent D., and Donoghue, Michael J. (1982). “Species Concepts: A Case for
Pluralism.” Systematic Zoology 31, 491–503.
Moore, James (1996). “Metabiographical Reflections on Charles Darwin.” In Shortland
and Yeo (1996, 267–281).
——— (1997). “Wallace’s Malthusian Moment: The Common Context Revisited.” In
Lightman (1997, 290–311).
Morgan, T.H. (1903). Evolution and Adaptation. New York: Macmillan.
Murray, Andrew (1859). “On Mr. Darwin’s Theory of the Origin of Species.” Proceedings
of the Royal Society of Edinburgh (1857–1862) 4, 274–291.
Nagel, Ernest (1961). The Structure of Science. New York: Harcourt, Brace & World.
O’Hara, Robert J. (1993). “Systematic Generalization, Historical Fate, and the Species
Problem.” Systematic Biology 42, 231–246.
Ospovat, Dov (1981). The Development of Darwin’s Theory: Natural History, Natural
Theology, and Natural Selection, 1838–1859. Cambridge: Cambridge University Press.
Otte, Daniel, and Endler, John A., eds. (1989). Speciation and Its Consequences. Sunder-
land, MA: Sinauer Associates.
Owen, Richard (1860). “On the Origin of Species by means of Natural Selection.” Edinburgh
Review 3, 487–532.
Padian, Kevin (1999). “Charles Darwin’s Views of Classification in Theory and Practice.”
Systematic Biology 48, 352–364.
Parshall, Karen Hunger (1982). “Varieties As Incipient Species: Darwin’s Numerical Analysis.”
Journal of the History of Biology 15, 191–214.
Paterson, H.E.H. (1985). “The Recognition Concept of Species.” In E.S. Vrba, ed. (1985,
21–29). Species and Speciation. Pretoria, South Africa: Transvaal Museum.
Percival, W. Keith (1987). “Biological Analogy in the Study of Languages Before the Advent
of Comparative Grammar.” In Hoenigswald and Wiener (1987, 3–38).
Poulton, E.B. (1903). “What is a Species?” Royal Entomological Society of London, Transactions
(Appendix) 51, 77–116.
Prichard, James Cowles (1813). Researches into the Physical History of Man. London: John
and Arthur Arch.
Putnam, Hilary (1973). “Explanation and Reference.” In Putnam, ed. (1975, 196–214).
Mind, Language and Reality. Cambridge: Cambridge University Press.
 References 261

Quine, W.V.O. (1986). Philosophy of Logic. 2nd ed. Cambridge: Harvard University Press.
Radick, Gregory (2003). “Is the Theory of Natural Selection Independent of Its History?”
In Hodge and Radick (2003, 143–167).
Ramsbottom, J. (1938). “Linnaeus and the Species Concept.” Proceedings of the Linnean
Society of London May 24, 192–219.
Raup, David M. (1991). Extinction: Bad Genes or Bad Luck? New York: W.W. Norton &
Company.
Richards, Robert J. (1992). The Meaning of Evolution: The Morphological Construction and
Ideological Reconstruction of Darwin’s Theory. Chicago: University of Chicago Press.
——— (2002). The Romantic Conception of Life: Science and Philosophy in the Age of Goethe.
Chicago: University of Chicago Press.
——— (2004). “Michael Ruse’s Design for Living.” Journal of the History of Biology 37,
25–38.
Ridley, Mark (1989). “The Cladistic Solution to the Species Problem.” Biology & Philoso-
phy 4, 1–16.
——— (1993). Evolution. Boston: Blackwell Scientific Publications.
Rieppel, Olivier (1986). “Species Are Individuals: A Review and Critique of the Argument.”
In Max K. Hecht, et al., eds. (1986, 283–387). Evolutionary Biology. Volume 20. New
York: Plenum Press.
Ritvo, Harriet (1997). “Zoological Nomenclature and the Empire of Victorian Science.” In
Lightman (1997, 334–353).
Roger, Jacques, and Fischer, Jean-Louis, eds. (1987). Histoire du Concept d’Espèce dans les
Sciences de la Vie. Paris: Fondation Singer-Polignac.
Rosen, David (1978). “The Importance of Cryptic Species and Specific Identifications as
Related to Biological Control.” Biosystematics in Agriculture: Beltsville Symposia in
Agricultural Research 2, 23–35.
Rosen, Donn E. (1979). “Fishes from the Uplands and Intermontane Basins of Guatemala:
Revisionary Studies and Comparative Geography.” Bulletin of the American Museum of
Natural History 162, 267–376.
Ruse, Michael (1975). “Darwin’s Debt to Philosophy: An Examination of the Influence of
the Philosophical Ideas of John F.W. Herschel and William Whewell on the Develop-
ment of Charles Darwin’s Theory of Evolution.” Studies in the History and Philosophy of
Science 6, 159–181.
——— (1979). The Darwinian Revolution. Chicago: University of Chicago Press.
——— (1987). “Biological Species: Natural Kinds, Individuals, or What?” British Journal
for the Philosophy of Science 38, 225–242. Reprinted in Ereshefsky (1992a, 343–363).
———, ed. (1989). What the Philosophy of Biology Is. Dordrecht, Netherlands: Kluwer.
——— (1993a). “Were Owen and Darwin Naturphilosophen?” Annals of Science 50, 383–388.
——— (1993b). “Will the Real Charles Darwin Please Stand Up?” Quarterly Review of
Biology 68, 225–231.
262 References

——— (2000). “Darwin and the Philosophers: Epistemological Factors in the Develop-
ment and Reception of the Theory of the Origin of Species.” In Creath and Maienschein
(2000, 3–26).
——— (2004). “The Romantic Conception of Robert J. Richards.” Journal of the History
of Biology 37, 3–23.
Salthe, Stanley N. (1985). Evolving Hierarchical Systems. New York: Columbia University Press.
Schwartz, Stephen P., ed. (1977). Naming, Necessity, and Natural Kinds. Ithaca, NY:
Cornell University Press.
Schweber, Silvan S. (1985). “The Wider British Context in Darwin’s Theorizing.” In Kohn
(1985a, 35–69).
Secord, James A. (1989). “Behind the Veil: Robert Chambers and Vestiges.” In James R.
Moore, ed. (1989). History, Humanity and Evolution: Essays for John C. Greene. Cambridge:
Cambridge University Press.
Sedgwick, Adam (1860). “Objections to Mr. Darwin’s Theory of the Origin of Species.” The
Spectator Mar. 24, 285–286; Apr. 7, 334–335.
Shortland, Michael, and Yeo, Richard, eds. (1996). Telling Lives in Science: Essays on Scientific
Biography. Cambridge: Cambridge University Press.
Simpson, George Gaylord (1961). Principles of Animal Taxonomy. New York: Columbia
University Press.
Simpson, Richard (1860). “Darwin on the Origin of Species.” Rambler 2, 361–376.
Smith, Neil (1999). Chomsky: Ideas and Ideals. Cambridge: Cambridge University Press.
Sober, Elliott (1993). Philosophy of Biology. Boulder: Westview Press.
Sonea, Sorin, and Panisset, Maurice (1983). A New Bacteriology. Boston: Jones and Bartlett.
Stamos, David N. (1996). “Was Darwin Really a Species Nominalist?” Journal of the
History of Biology 29, 127–144.
——— (1998). “Buffon, Darwin, and the Non-Individuality of Species—A Reply to Jean
Gayon” Biology & Philosophy 13, 443–470.
——— (1999). “Darwin’s Species Category Realism.” History and Philosophy of the Life
Sciences 21, 21–70.
——— (2000). “Book Review of Stephen G. Alter’s Darwinism and the Linguistic Image:
Language, Race, and Natural Theology in the Nineteenth Century.” Annals of Science 57,
319–321.
——— (2002). “Species, Languages, and the Horizontal/Vertical Distinction.” Biology &
Philosophy 17, 171–198.
——— (2003). The Species Problem: Biological Species, Ontology, and the Metaphysics of
Biology. Lanham, MD: Lexington Books.
——— (2005). “Pre-Darwinian Taxonomy and Essentialism—A Reply to Mary Winsor.”
Biology & Philosophy 20, 79–96.
——— (2006). “Popper, Laws, and the Exclusion of Biology from Genuine Science.”
manuscript in review.
 References 263

——— (forthcoming). Evolution and the Big Questions: Sex, Race, Religion, and Other
Matters. Blackwell Publishing.
Stauffer, R.C., ed. (1975). Charles Darwin’s Natural Selection, Being the Second Part of His
Big Species Book Written From 1856 to 1858. Cambridge: Cambridge University Press.
Stevens, Peter F. (1980). “Evolutionary Polarity of Character States.” Annual Review of
Ecology and Systematics 11, 333–358.
——— (1984). “Haüy and A.-P. Candolle: Crystallography, Botanical Systematics, and
Comparative Morphology, 1780–1840.” Journal of the History of Biology 17, 49–82.
——— (1992). “Species: Historical Perspectives.” In Keller and Lloyd (1992, 302–311).
——— (1994). The Development of Biological Systematics: Antoine-Laurent de Jussieu,
Nature, and the Natural System. New York: Columbia University Press.
——— (1997). “J.D. Hooker, George Bentham, Asa Gray and Ferdinand Mueller on
Species Limits in Theory and Practice: A Mid-Nineteenth-Century Debate and Its Reper-
cussions.” Historical Records of Australian Science 11, 345–370.
Strickland, Hugh E. (1837). “Rules for Zoological Nomenclature.” Magazine of Natural
History 1, 173–176.
——— (1842). “Report of a Committee Appointed ‘to Consider of the Rules by which the
Nomenclature of Zoology may be Established on a Uniform and Permanent Basis.’”
British Association for the Advancement of Science (London), 105–121.
Sulloway, Frank J. (1979). “Geographic Isolation in Darwin’s Thinking: The Vicissitudes
of a Crucial Idea.” In William Coleman and Camille Limoges, eds. (1979, 23–65). Studies
in History of Biology. Volume 3. Baltimore: Johns Hopkins.
——— (1982). “Darwin’s Conversion: The Beagle Voyage and Its Aftermath.” Journal of
the History of Biology 15, 325–396.
——— (1996). Born to Rebel: Birth Order, Family Dynamics, and Creative Lives. New York:
Pantheon Books.
Taub, Liba (1993). “Evolutionary Ideas and ‘Empirical’ Methods: The Analogy Between
Language and Species in Works by Lyell and Schleicher.” British Journal for the History
of Science 26, 171–193.
Templeton, Alan R. (1989). “The Meaning of Species and Speciation: A Genetic Perspec-
tive.” In Otte and Endler (1989, 1–27).
Thompson, Paul (1989). The Structure of Biological Theories. Albany: State University of
New York Press.
———, ed. (1995). Issues in Evolutionary Ethics. Albany: State University of New York Press.
Trask, R.L. (1996). Historical Linguistics. London: Arnold.
Van Lalen, Leigh (1973). “A New Evolutionary Law.” Evolutionary Theory 1, 1–30.
——— (1976). “Ecological Species, Multispecies, and Oaks.” Taxon 25, 233–239.
Reprinted in Ereshefsky (1992a, 69–78).
Wallace, Alfred Russel (1855). “On the Law Which Has Regulated the Introduction of
New Species.” Annals and Magazine of Natural History 16 (2nd ser.), 184–196.
264 References

——— (1859). “On the Tendency of Varieties to depart indefinitely from the Original
Type.” Journal of the Proceedings of the Linnean Society (Zoology) 3, 53–62. Reprinted in
Barrett (1977 II, 10–19).
Watson, Hewett Cottrell (1843). “Remarks on the Distinction of Species in Nature, and in
Books; preliminary to the notice of some variations and transitions of character, observed
in the native plants of Britain.” London Journal of Botany 2, 613–622.
——— (1845a). “On the Theory of ‘Progressive Development,’ applied in explanation of
the Origin and Transmutation of Species.” The Phytologist 2, 108–113, 140–147,
161–168, 225–228.
——— (1845b). “Report of an experiment which bears upon the specific identity of the
Cowslip and Primrose.” The Phytologist 2, 217–219.
——— (1859). Cybele Britannica. Volume 4. London: Longman.
Wayne, Robert K., and Gittleman, John L. (1995). “The Problematic Red Wolf.” Scientific
American 273, 36–39.
Wedgwood, Hensleigh (1833). “Grimm’s Deutche Grammatik.” Quarterly Review 50,
169–189.
Weinberg, Steven (1992). Dreams of a Final Theory: The Scientist’s Search for the Ultimate
Laws of Nature. New York: Pantheon Books.
Weinert, Friedel, ed. (1995). Laws of Nature: Essays on the Philosophical, Scientific and
Historical Dimensions. Berlin: Walter de Gruyter.
Wells, Rulon S. (1987). “The Life and Growth of Language: Metaphors in Biology and
Linguistics.” In Hoenigswald and Wiener (1987, 39–80).
West-Eberhard, Mary Jane (1992). “Adaptation: Current Usages.” In Keller and Lloyd
(1992, 13–18).
Wheeler, Quentin D., and Meier, Rudolf, eds. (2000). Species Concepts and Phylogenetic
Theory. New York: Columbia University Press.
Whewell, William (1837). History of the Inductive Sciences. Three volumes. London: Parker.
——— (1840). Philosophy of the Inductive Sciences. Two volumes. London: Parker.
Wilberforce, Samuel (1860). “Darwin’s Origin of Species.” Quarterly Review 108, 225–264.
Wiley, E.O. (1981). Phylogenetics: The Theory and Practice of Phylogenetic Systematics. New
York: John Wiley & Sons.
Wilson, Edward O. (1992). The Diversity of Life. Cambridge: Harvard University Press.
Wilson, Leonard G., ed. (1970). Sir Charles Lyell’s Scientific Journals on the Species Question.
New Haven: Yale University Press.
Wilson, Robert A., ed. (1999). Species: New Interdisciplinary Essays. Cambridge: MIT Press.
Winsor, Mary Pickard (1979). “Louis Agassiz and the Species Question.” In William
Coleman and Camille Limoges, eds. (1979, 89–117). Studies in History of Biology.
Volume 3. Baltimore: Johns Hopkins.
——— (2001). “The Practitioner of Science: Everyone Her Own Historian.” Journal of the
History of Biology 34, 229–245.
 References 265

——— (2003). “Non-Essentialist Methods in Pre-Darwinian Taxonomy.” Biology &

Philosophy 18, 387–400.
Wollaston, Thomas Vernon (1856). On the Variation of Species, With Especial Reference to
the Insecta. London: John Van Voorst.
——— (1860). “On the Origin of Species by means of Natural Selection.” Annals and
Magazine of Natural History 5 (3rd ser.), 132–143.
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 Index


adaptations, xvi, 9, 33, 75, 88–89, 90, 155, 161, 173, 189, 216, 221, 224,
91–94, 103–106, 111, 118, 120, 122, 234n3, 241n8
126–129, 132, 148–150, 183, 200, Bates, Henry Walter, 116, 119–121, 122
225, 234n2, 235n3 (ch. 4), 235n3 Baum, David A., 63
(ch. 5), 239n10, 245n3, 245n5, Beatty, John, xi, xvii, 19, 26, 28, 81–82,
246n9 110, 133, 158–164, 166, 174, 95–196,
Agassiz, Louis, 12, 19, 42, 135–136, 145, 197–199, 205, 240n2, 244n8
172, 173, 181, 216, 226–227, 231n2, Beck, Henrick Henricksen, 146
239n1, 239n2, 241n11, 243n17, Beer, Gillian, 56
245n4 Bentham, George, 13, 26, 92, 183–184,
Alter, Stephen G., 52, 56 226, 231n2, 242n14
Alvarez, Luis, xi–xii Bergmann’s Rule, 233n1
“amount of difference,” 67, 85–88, 148, Big Bang, 22–23
241n8 biodiversity, xii–xiii, 189, 229
angiosperms, 74 biological species concept, xiii, 4, 18, 37,
apostasy, 203 39, 42, 76, 108, 111, 116, 117, 119,
archetypes, 181, 247n10 125, 127–128, 130, 156, 158, 237n2,
Archimedes, 151 238n6
argument from design, 34, 160, 245n3 Blyth, Edward, 68–69, 132, 137,
Aristotle, 24, 29–30, 43, 133, 134, 137, 140–141, 144, 147, 174, 185
145, 153–155, 240n1 Bopp, Franz, 55, 56
artificial selection, 34, 89–90, 91–92, 201, Bowler, Peter J., 209–210, 212, 213, 218,
235n3 (ch. 5), 245n5 244n9
asexual organisms, 125 Boyd, Richard, 226, 247n10
Ashton, Paul A., 74, 77 Brocchi, Giovanni, 41
Bronn, Heinrich, 55
Babbage, Charles, 54 Browne, Janet, 207, 246n8
Babington, Charles Cardale, 13 Bruno, Giordano, 189
Babinski, Edward T., 203 Buffon, Georges Louis-Leclerc Comte de,
baboons, 18 4, 138, 145, 226, 227
Baer, Karl von, 73, 247n10 Busk, George, 242n15
barnacles, 7–8, 46–49, 84, 86, 88, 148, Butler, Joseph, 146

267
268 INDEX

butterflies, 116, 119–120, 123 157, 166, 167, 168, 170, 179–180,
185–186, 192, 193, 200, 212, 216,
Camardi, Giovanni, 247n10 231n3, 241n6, 243n17, 247n10
Candolle, Alphonse de, 173 creationist species concept, 82, 83,
Candolle, Augustin-Pyramus de, 122, 165–166
170, 236n8 Crinum, 126
Carpenter, William Benjamin, 178, crystallography, 35, 101–102, 236n8, 236n9
242n15 Cuvier, Georges, 73, 181, 245n4, 247n10
caste polymorphism, 235n3 (ch. 3)
categories, 22–25, 27–28, 30, 153–156, Dana, James Dwight, 142–143, 144, 145,
190. See also species, category v. taxon 166, 227
cats, 141 Darwin, Emma, 222
cattle, 68–69, 113, 132, 190, 237n1 Darwin, Erasmus, 105, 246n8
Chalmers, A.F., 243n2 Darwin, Francis, 41, 241n9
Chambers, Robert, 9, 11, 57, 105, 161, Darwin-Knight law, 99, 115
163, 180, 184, 185, 192, 234n4, Davies, Anna Morpurgo, 55
245n3, 246n7 Dawkins, Richard, 23, 27, 61
Chomsky, Noam, 208 Dawson, John, 13
chronospecies, 62–63 deer, 120, 239n9
cladism, 38, 58, 66–67, 73, 74–79, definition, 29–30, 43–44, 46, 48, 81, 84,
127–128 88, 99, 133, 154–156, 159, 162, 163,
cladistic species concept, 51, 61, 63 165–166, 171, 174–175, 195, 240n4
classes, 21–22, 27, 29 Dennett, Daniel C., 212
classification. See natural system/ de Queroz, Kevin, 49
classification Desmond, Adrian, xvii, 173, 175,
clusterism, 225–226, 247n10 180–181, 208–209, 217, 218–220,
common descent, xvi, ch. 4, 85, 87, 90, 222, 223, 242n15, 246n6, 246n7,
91, 102, 127, 132, 168, 184, 200, 246n8, 246n9
210, 216, 225, 243n16 “developmental hypothesis,” 57
Comte, Auguste, 103 “Devil’s chaplain,” 160
“confessing a murder,” 172 Dewey, John, xvii, 187–189, 212
“consilience of inductions,” 96–97, 221 divergence, principle of, 9, 34, 46, 69, 73,
“constant and distinct” characters, 69, 85, 147, 219–222, 228–229, 244n9
87–88, 89–90, 114, 116–117, 118, division of labor, 219–222
120, 127, 132, 148–149, 155, Dobzhansky, Theodosius, xv, 4, 127, 209,
241n8 228
constellations, 1, 19, 26 dogs, 67–68, 70, 109–110, 113, 141, 179,
convergence, 69–70, 87, 236n9, 239n10 190, 237n1, 242n14
conversion, 105, 170–184, 191, 192–194, Dollo, Louis, 32
200, 201–, 242n15, 243n2, 243n4, Dollo’s Law, 32–33, 233n1
244n9 domestication, 237n1
Copernicus, Nicolaus, 211, 245n2 Donnellan, Keith, 195
correlation, 220 ducks, 72
Cowan, S.T., 18 Dupré, John, 59
cowslips. See primroses and cowslips Dupree, A. Hunter, 170, 171
Cracraft, Joel, xiii, 63, 129, 189
creationism, 3, 17, 82, 94, 123, 135, 153, Eaton, John Matthews, 90–91, 182–183
 Index 269

ecological niches, xvi, 9, 32, 45, 70, 72, Geoffroy Saint-Hilaire, M. Isidore, 245n3
90, 109, 118, 121–124, 220–221, Gérard, Frédéric, 7
234n2 Ghiselin, Michael T., x–xi, xviii, 19, 26,
ecological species concept, xiv 28–30, 39, 58, 65, 67, 81, 110–111,
Ehrlich, Paul R., 121 116, 133, 153–154, 158, 163, 164,
eidos, 187–188 222, 233n1, 238n5, 240n1
Einstein, Albert, 200 “giving up species,” 6, 176, 177, 202,
Eiseley, Loren, 200 242n12
Eldredge, Niles, 39, 51, 233n8 gold, 22, 23
Ellegård, Alvar, 19, 166, 241n3 Gorovitz, Samuel, 168
embryological evolution, 213–217 Gould, John, 237n2
Endersby, Jim, 186 Gould, Stephen Jay, xi, 233n8
Endler, John A., 38 gourds, 112
“entity,” 133, 179, 200, 242n14 gradualism, 46, 54, 94, 134, 155–156,
essentialism, 23–25, 30, 43–44, 72–73, 201, 202–204, 210, 234n2, 244n9
100, 102, 133, 134, 135–136, Grant, Verne, 16
137–139, 142, 145, 150, 153–156, gravity, 98, 99
165–166, 172, 187–188, 211, 212, Gray, Asa, 13, 86, 102, 144, 170–172,
218, 225–227, 240n1, 241n3, 180, 186, 202, 233n8
244n5, 247n10 Great Chain of Being, 215, 247n10
evil, problem of, 161 Grene, Marjorie, 247n10
“evolution,” 58, 216, 217 Grimm, Jacob, 55, 56
evolutionary ethics, 229 groups, 26–27, 32–33
evolutionary psychology, 229
extinction, xii–xiii, 32–33, 34, 40–41, 43, habit, 103–106, 214
45, 53, 54, 57, 61, 70–71, 74, 75, 78, Hacking, Ian, 244n5
85, 86, 122, 127–128, 149 Haeckel, Ernst, 213
Haldane, J.B.S., 15–16
Fawcett, Henry, 13, 198, 202 Hamilton, Alexander, 56
fertility, xvi, 4–5, 42, 108–109, 111, Harvey, William Henry, 58, 93, 234n2
113–116, 223, 234n2, 237n1 Hattiangadi, Jagdish, 190
fish, 135 Haüy, Abbé, 101
Flew, Antony, 18 Hennig, Willi, 127–128
Fodor, Jerry, 58 Henslow, John Stevens, 102
“forms,” 122, 124, 171, 235n3 (ch. 5) Herbert, William, 16–17, 126
fossils, 43 hermaphrodites, 114
Foucault, Michel, 208, 224 Herschel, John F.W., 54, 95–98,
Frost, Darrel L., 66 100–101, 102–103, 235n4, 236n6
“higgledy-piggledy,” 235n4, 236n6
Galapagos Archipelago, 123, 178 “higher” and “lower,” 223
Gärtner, Carl Friedrich, 83, 89, 112, 115 higher taxa, 5–6, 17, 31, 67, 73, 163,
Gayon, Jean, 233n7 226–227
geese, 116, 239n10 Hippeastrum, 126
genealogical species concept, 63 “historical entities,” 78–79
genealogy. See common descent Hodge, Jon, 19, 42, 97, 163–164, 240n3
genera, 2, 176, 184, 220, 231n2 hollyhocks, 112
genius, 204, 207, 247n10 homeostasis, 226
270 INDEX

homologies, 196, 216, 247n10 Kielmeyer, Karl Friedrich, 215

Hooker, Joseph Dalton, 7, 10, 17, 136, Kitcher, Philip, xvii, 164, 196–198,
144, 145–146, 171, 172–176, 177, 244n8
178, 180, 183, 186, 202, 226, 231n2, Kohn, David, 207, 220–221
240n4, 241n8, 241n10, 241n11, Kölreuter, Johann Gottlieb, 174, 237n3
242n15 Kottler, Malcolm Jay, 39–40, 82, 83,
Hopkins, William, 13, 198, 239n1 116–117, 119, 146–147, 238n4, 238n6
horizontal v. vertical dimension, xv, 4–5, Kripke, Saul, 195, 244n5
ch. 3, 77–79, 84, 87, 88, 90, 127–128, Kuhn, Thomas S., xvii, 160, 189,
129, 130, 156–157, 200, 213, 216, 190–192, 194–195, 198, 200, 201,
223, 232n4 205, 209, 243n1, 243n4, 245n2
horses, 110, 116, 138
Hoyningen-Huene, Paul, 243n1, 243n2 Lakatos, Imre, 243n2
Hull, David L., 65, 219, 240n1, 243n4, laissez-faire economics, 218, 221, 222
247n10 Lamarck, Jean Baptiste de, 7, 9, 10, 14,
human races, 84, 89, 128–129, 140, 155, 57, 104, 105, 136, 210, 214–215, 217,
162, 235n1, 242n14 228, 232n6, 234n4, 243n17, 245n3
Humboldt, Alexander von, 57, 95–96 languages, xv, 44, 51–62, 71, 74, 76,
Humboldt, Wilhelm von, 57 77–79, 102, 127, 130, 201, 216, 221
Hume, David, 236n6 LaPorte, Joseph, 195, 198–200
hummingbirds, 237n2 Larson, James L., 138
Huxley, Julian S., 41–42 Lavoisier, Antoine Laurent, 196–197
Huxley, Thomas Henry, 124–125, 145, laws of nature, xvi, ch. 2, 54, 55–57, 58,
172, 177, 178, 180–183, 186, 202, 95–96, 98–103, 128, 138, 167, 185,
208–209, 222–223, 234n2, 242n15, 192, 193, 200, 211, 216, 219, 225,
242n16 243n17, 245n5
hybridity, 17, 45, 69, 74, 75, 76, 113, Leikola, Anto, 24
126, 156, 238n4 Levin, Donald, 18, 235n3 (ch. 4)
Lewes, George Henry, 13, 232n5
“incipient species,” xvi, 3, 32, 70, 82, 133, Lindley, John, 226
134, 146, 147, 148, 149, 150, 168, Linnaeus, Carolus, 24, 31, 72, 93, 136,
176, 197, 236n7, 238n4, 238n7, 137–138, 144, 145, 154, 166, 192,
241n7 227, 231n2, 240n3, 242n14, 243n17
incommensurability, 160, 190–191, Lipton, Peter, 193
192–194, 195, 196–200, 201 “living fossils,” 32, 48–49, 51, 90
Industrial Revolution, 219 Lobelia, 126
inference to the best explanation, 193, 222 Locke, John, 18, 215
“infinite evolution,” 61, 62, 76 Lubbock, John, 172
instincts, 4–5, 127 Luckow, Melissa, xv, 129, 130
instrumentalism, 243n3 lumpers v. splitters, 11, 72, 101, 135–136,
145, 146, 162, 180, 227, 236n7
Jenyns, Leonard, 224 Lyell, Charles, 14, 17, 30, 41, 53–54, 70,
Jordan, Alexis, 11 71, 94, 95, 100, 108, 121–122,
Jussieu, Antoine-Laurent de, 31, 139, 141–142, 144, 145, 167, 168, 175,
144, 145, 226, 227 176–180, 185, 186, 202, 204, 215,
241n8, 242n13, 242n15, 243n17
Kant, Immanuel, 23–24 Lysenko, Trofim Denisovich, 228
 Index 271

Lythrum salicaria, 112–113, 125, 238n4, natural law. See laws of nature
246n7 natural selection, xvi, 9, 33–35, 43, 45,
53, 58, 60, 75, 88–89, 90, 91–92, 94,
Macculloch, John, 5, 98 97, 98–99, 103–106, 111, 118, 127,
maize, 112 128–129, 147, 149–150, 163, 185,
Mallet, James, xiv 188, 201, 210, 214, 217, 221, 225,
Malthus, Thomas Robert, 95, 96, 218, 228, 235n3 (ch. 4), 238n7, 244n9,
219, 244n9 245n3, 245n5, 246n9
mammals, 234n2 natural system/classification, 6, 53, 65–66,
man. See human races 85, 98, 100–102, 139, 183, 192, 211,
Masters, Maxwell T., 239n1 236n9, 243n16
Matthew, Patrick, 246n9 Naudin, Charles V., 85
Mayr, Ernst, xiii, 2, 4, 16–18, 27, 38, 39, new historiography, the, xvii, ch. 10,
45–46, 72–73, 86, 117, 127, 135, 242n15
146–147, 150, 156, 157–158, 210, Newton, Isaac, 98, 99, 210, 211
213, 239n2, 247n10 niches. See ecological niches
McDade, Lucinda A., 77, 156
McOuat, Gordon, 19, 164–166, 222–223, O’Hara, Robert J., 188
225, 231n3 ontogenetic polymorphism, 72, 73, 87,
meaning, theory of, 194–195, 196–198, 123, 168, 235n3 (ch. 5), 241n5
199, 205, 244n5, 244n6, 244n8 orchids, 92–93, 108, 114, 126, 246n9
Meckel, Johann Friedrich, 215 orthogenesis, 57, 210
“messy situations,” 2–3, 7, 8, 45, 88, Ospovat, Dov, 9, 234n4
156–157, 185, 232n4 Owen, Richard, 12, 247n10
Mill, John Stuart, 242n15
Milne-Edwards, Henri, 219–220 Padian, Kevin, 85
mimicry, 119–120 Paley, William, 34, 99, 245n3
mineralogy, 100–102, 236n9 Pallas, Pyotr, 68, 109
missionary position, 223 Pascal, Blaise, 204
Modern Synthesis, xii, 26, 129, 188, 210, Paterson, Hugh E.H., 108–109, 125,
213, 217 126, 188
monadic theory of evolution, 40–42, 51, pattern v. process, xvi, 130, 213, 216
177 Paul, St., 204
monophyly, xvi, 66–67, 71, 74 peacocks, 93
monstrosities, 72, 94, 141, 168, 241n5 Phillips, John, 171
Moore, James R., xvii, 173, 175, 208, 209, phlogiston, 196–197
217, 218–220, 221, 222, 223, 246n6, phylogenetic species concept, xiii, 63,
246n7, 246n8, 246n9, 247n10 76–77
morphological species concept, xiv, 37, pigeons, 90–91, 141, 182–183, 238n4,
76, 86, 124, 130, 158 242n15
moths, 246n9 pigs, 237n1
Müller, Max, 15 Planck, Max, 243n4
multiple origins, 33, 53, 60–61, 68, 69, polyploidy, 60–61, 74–75, 129, 235n3 (ch. 4)
73–76, 77, 78, 179, 241n11 postmodernism, 208 , 219
“mystery of mysteries,” 54 Poulton, E.B., 15, 88
Prichard, James Cowles, 5, 108, 139–140,
natural kinds, 27, 28, 195, 222, 244n5 141, 226, 240n3
272 INDEX

primroses and cowslips, 14, 72, 82–83, Secord, James A., 246n7
84, 86, 87, 88–89, 91, 92, 110, 112, Sedgwick, Adam, 102, 200, 239n1
120, 122, 124–125, 132, 148, 173, Serres, Etienne R.A., 215
190, 224, 232n4, 238n5 sets, 21–22, 26–27
process. See pattern v. process sex, 222–224, 246n7, 246n8
process laws, 99–100, 236n6 sexual dimorphism, 72, 73, 123, 168,
progress, 213–217, 219, 245n5 235n3 (ch. 5), 241n5
punctuated equilibria, 38, 39, 51, 66, sexual selection, 222, 228, 235n3 (ch. 5)
190–191, 201, 233n8 Shaw, Kerry L., 63
Putnam, Hilary, 195, 244n5, 244n6, sheep, 141
244n7 similarity, 66–67, 69–70, 74, 75–76, 77,
85, 86–87, 113–114, 127, 129, 130,
Quine, W.V.O., 58 132, 184, 189, 232n5
Simpson, George Gaylord, 26, 38, 51,
rabbits, 93, 116 121, 146–147, 150, 158, 241n3
Radick, Gregory, 233n7 Sismondi, Jean, 220
Raup, David M., 61 Smith, Adam, 218
recapitulation, 214–217, 245n4 Sober, Elliott, 18
recognition species concept, 108–109, 126 social constructionism, 208, 221, 222,
Red Queen hypothesis, 32 224, 228, 247n10
red wolf, xiii speciation, 33, 37, 44, 45–46, 51, 57,
reductio ad absurdum, 168 60, 62, 70–71, 74, 127–129, 147,
“reference potential,” 196–198 221, 222, 228, 233n1
reflecting goniometer, 101, 102 species:
relativism, epistemological, 218–219 aging, 40–41, 61, 233n1
reversion, law of, 30–32, 90, 138, 139, as individuals, xv, 28–30, 38, 40–42,
140–142, 144–145, 148, 192, 200, 57, 59–62, 63, 65–66, 75–76, 8–79,
213, 221, 227, 247n10 214, 216
Richards, Robert J., 103–104, 213–218, as lineages, 49–51
225 category v. taxon, xi, 2, 19, ch.2, 81, 133,
Ridley, Mark, 38, 76 153–157, 158–160, 163, 165,
Ritvo, Harriet, 224 166–167, 199, 232n5
Rosen, David, 188 Darwin’s concepts of, 2–3, 4–5, 117,
Rosen, Donn E., 76–77 127, 163, 164, 237n1, 238n7
rudiments, 52, 53, 239n10, 243n17 definitions of Darwin’s contemporaries,
Ruse, Michael, 86, 96–97, 209, 215, 223 3, 11–12, 131–132, 135,
137–144, 162, 164–166, 170,
saltations, 90, 93–94, 105, 124, 173, 173–174, 179–180, 184, 241n11
201, 203, 204, 210, 234n2, 234n4, endemic, 160
243n17, 244n9 good, 44, 45, 47, 48–49, 52, 87, 88,
Saussure, Ferdinand de, 58 125, 129, 155
Schlegel, Friedrich von, 55 group v. rank, 26, 166–167, 182, 184
Schleicher, August, 55–56 nominalism, xiii, ch. 1, 26, 29, 39,
Schweber, Silvan, 97–98 81–82, 157, 159, 160, 163, 168,
scientific revolutions, 160, ch. 9, 209–212 170–186, 188, 199, 241n6, 242n14
Sclater, Philip, 93 plasticity, 60, 122, 179, 217
seals, 72 pluralism, xiii, 25, 129, 130, 188
 Index 273

problem, xii, xiv, xv, 21, 25 use and disuse, 34, 103–106
range, 45, 60
representative, 45 Van Valen, Leigh, 32, 121
sibling, 40, 116–117 variation, 45, 46, 47, 53, 57–58, 89, 90,
typical, 226 04–105, 118, ch 7., 156, 180, 212,
Spencer, Herbert, 99, 242n15 234n2
splitters. See lumpers v. splitters law of limited, 30, 54, 142, 144–145,
spontaneous generation, 40, 41 148, 192, 200, 213, 221, 247n10
Stamos, David N., x, xi, 18, 59, 76, 77, varieties, xvi, 2–3, 30–32, 83, 87, 90,
110, 129, 166, 226, 232n5 113, 114, 118, ch. 7, 168, 169, 197,
sterility, xvi, 4, 17, 40, 54, 83, 89, 108, 212, 238n7, 241n3, 241n5, 241n10.
109–116, 124–126, 127–128, 132, See also “incipient species”
140, 157–158, 164, 174, 200, intermediate, 43, 45, 82–84, 86,
222–223, 235n3 (ch. 4), 237n1, 93, 148
237n3, 238n4, 246n7 vera causa, 34–35, 74–75, 95–103, 114,
Stevens, Peter F., 26, 128, 173, 175, 129, 158, 182, 189, 201, 225
236n7, 236n8 Verbascum, 112, 115, 124, 125, 126
Strickland, Hugh E., 44, 155, 165–166,
224, 226 Wallace, Alfred Russel, 9, 31, 144, 170,
subspecies, 134 177, 185–186, 192, 211, 243n17,
Sulloway, Frank J., 19, 86, 117, 158, 245n3, 246n6
192, 193–194, 202, 204, 243n4, “war of nature,” 160
246n6 Waterhouse, George Robert, 226
“systematic affinity,” 113–114 Watson, Hewett Cottrell, 10–11, 14,
17, 69, 136, 144, 150–151, 162–163,
Tegetmeier, William Bernhard, 238n4 167, 184–185, 186, 200, 204,
teleology, 213–217 210–211, 226, 231n2, 231n3,
Templeton, Alan R., 121 232n4, 242n12
theistic evolution, 210 Wedgwood, Hensleigh, 56
Tiedemann, Friedrich, 215 Whewell, William, 14, 55, 96–97, 101,
tobacco, 112, 124 167, 236n5, 247n10
Tooke, John Horne, 52 Wiley, Edward O., 27, 78
Tree of Life, 37, 45, 49, 185, 243n17 willow wrens, 40, 116–117, 119, 120
Treviranus, Gottfried Reinhold, 215 Wilson, Edward O., xiii
turtles, 239n2 Winsor, Mary P., 219, 225–228, 239n2,
247n10
unity of science, 102, 103, 106 Wollaston, Thomas Vernon, 12, 14,
unity of type, 6, 211, 215 143–144, 166, 167, 177, 181, 192,
“universal acid,” 188, 212 198, 200, 227, 231n3
universal species concept, xiv, 28, 29–130 woodpeckers, 123
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