Darwinism

First published Fri Aug 13, 2004; substantive revision Tue May 26, 2015
Darwinism designates a distinctive form of evolutionary explanation for the history and diversity of life
on earth. Its original formulation is provided in the first edition of On the Origin of Species in 1859.
This entry first formulates Darwin's Darwinism in terms of five philosophically distinctive themes: (i)
probability and chance, (ii) the nature, power and scope of selection, (iii) adaptation and teleology, (iv)
nominalism vs. essentialism about species and (v) the tempo and mode of evolutionary change. Both
Darwin and his critics recognized that his approach to evolution was distinctive on each of these topics,
and it remains true that, though Darwinism has developed in many ways unforeseen by Darwin, its
proponents and critics continue to differentiate it from other approaches in evolutionary biology by
focusing on these themes. This point is illustrated in the second half of the entry by looking at current
debates in the philosophy of evolutionary biology on these five themes.
1. Introduction
2. Darwin and Darwinism
2.1 Darwin's Life
2.2 Darwin's Darwinism
2.3 Philosophical Problems with Darwin's Darwinism
3. The Five Core Philosophical Problems Today
3.1 The Roles of Chance in Evolutionary Theory
3.2 The Nature, Power and Scope of Selection
3.3 Selection, Adaptation and Teleology
3.4 Species and the Concept of Species
Bibliography
References
Further Reading
Academic Tools
Other Internet Resources
Related Entries

1. Introduction
Scientific theories are historical entities. Often you can identify key individuals and documents that are
the sources of new theoriesEinstein's 1905 papers, Copernicus 1539 De Revolutionibus,
Darwin's On the Origin of Species. Sometimes, but not always, the theory tends in popular parlance to
be named after the author of these seminal documents, as is the case with Darwinism.
But like every historical entity, theories undergo change through time. Indeed a scientific theory might
undergo such significant changes that the only point of continuing to name it after its source is to
identify its lineage and ancestry. This is decidedly not the case with Darwinism. As Jean Gayon has
recently put it:
 The Darwin-Darwinism relation is in certain respects a causal relation, in the sense that
Darwin influenced the debates that followed him. But there is also something more: a kind
of isomorphism between Darwin's Darwinism and historical Darwinism. It is as though
Darwin's own contribution has constrained the conceptual and empirical development of
evolutionary biology ever after. (Gayon 2003, 241)

Darwinism identifies a core set of concepts, principles and methodological maxims that were first
articulated and defended by Charles Darwin and which continue to be identified with a certain
approach to evolutionary questions.[1] We will thus need to begin with Darwin's Darwinism as
articulated in On the Origin of Species in 1859. We will then examine these same themes as they have
been discussed by evolutionary biologists and philosophers of biology from the beginnings of the Neo-
Darwinian Synthesis to the present.
Charles Darwin was not, as we use the term today, a philosopher, though he was often so described
during his lifetime.[2] Nevertheless, for an encyclopedia of philosophy what is needed is a discussion of
the impact of philosophy on Darwin's Darwinism, and the impact of Darwin's Darwinism on topics that
both he, and we, would consider philosophical. We focus here on the impact of philosophical
discussions about the nature of science during Darwin's lifetime on Darwin's scientific research,
thinking and writing; and on the impact of that research, thinking and writing on philosophy. Taking the
time to do such philosophical archaeology stems from a conviction that if the concept of Darwinism
has legitimate application today, it is due to a set of principles, both scientific and philosophical, that
were articulated by Darwin and that are still widely shared by those who call themselves Darwinians
or neo-Darwinians.

2. Darwin and Darwinism

2.1 Darwin's Life
Charles Darwin was born February 12, 1809 and died April 18, 1882. It was a time of radical changes
in British culture, and his family background put him in the midst of those changes. His grandfather,
Erasmus Darwin, was a prosperous and highly respected physician living in Western England, south of
Birmingham. He was also a philosophical radical, pushing ideas born in the French Enlightenment,
ideas about human equality and liberty, including the liberty to think freely about the existence of God
and about natural origins for the earth's creatures. He wrote a number of very popular works of natural
history, some in verse, in which he defended views about progress that included evolutionary
speculations about the upward progress of living things from primordial beginnings.
Erasmus Darwin was an early member of an informal group of free thinkers self-styled the Lunar
Society,[3] that met regularly in Birmingham to discuss everything from the latest philosophical and
scientific ideas to the latest advances in technology and industry. The Society included James Watt,
Joseph Priestly and Charles Darwin's other grandfather, Josiah Wedgwood. Wedgwood, like Erasmus
Darwin, lived in Staffordshire and was in the process of developing a family pottery works into a major
industrial concern by applying new scientific and technological ideas to the production of china. The
religious inclinations of the group were non-conforming and included a number of Unitarians, a sect
Erasmus Darwin referred to as a featherbed to catch a falling Christian. Looked upon with suspicion
by High Church conservatives, they actively promoted in Great Britain the revolutionary philosophical,
scientific and political ideas sweeping across Europe and the Americas. Most had spent considerable
time absorbing Enlightenment ideas in Edinburgh, Scotland.
Under the circumstances, it is not surprising that Robert Darwin, Charles father, should follow in his
father's footsteps and become a doctor, nor that he should end up marrying Susannah Wedgwood, by all
reports Josiah's favorite offspring. Politically and philosophically engaged, Susannah worked to
organize her children's education in the town of Shrewsbury, where she and Robert took up residence.
She sent her children to a day school operated by Unitarian minister Rev. George Case and this is
where Charles began his education. Unfortunately, Susannah died in 1817 when Charles was only 8,
and his father then transferred him to the Shrewsbury School, operated by Dr. Samuel Butler,
grandfather of the novelist (and sometime satirist of Darwin's work) of the same name. Nothing could
have been worse for the development of my mind than Dr. Butler's school Charles proclaimed in the
autobiography he wrote for his family, and he escaped down the street to his home whenever he could.
His older siblings took good care of him, under the Doctor's watchful eye. Early letters indicate that he
and his brother Erasmus were enthusiastic amateur chemists, and after his brother went up to
Cambridge their letters were often full of possible experiments, orders to purchase chemicals and
equipment for their laboratory, and discussions of the latest discoveries. This was an obvious enough
passion that his classmates nicknamed him Gas. During summers he helped his father on his rounds
to his patients, and when only 16 his father sent him and his brother to Edinburgh for the best medical
education Great Britain had to offer. Erasmus needed to move from Cambridge to a proper medical
school to complete his medical education, and young Charles was taken out of Shrewsbury School
early to accompany his brother to Edinburgh, apparently being prepared to follow in his father's and
grandfather's footsteps in medicine. The two brothers arrived in Edinburgh in October of 1825.
Erasmus left after the first year, leaving his brother on his own during his second year at Edinburgh.
Privately, Darwin early on decided he could not practice medicine. But his already serious inclination
toward science was considerably strengthened at Edinburgh both by some fine scientific lectures in
chemistry, geology and anatomy and by the mentoring of Dr. Robert Grant. Grant certainly knew that
young Charles was Erasmus Darwin's grandson; Grant expounded evolutionary ideas derived from
Jean-Baptiste Lamarck and Charles grandfather. But his primary gift to Charles was introducing him to
marine invertebrate anatomy and the use of the microscope as a scientific tool and as an aid to
dissecting extremely small creatures dredged out of the Firth of Forth. Darwin joined an Edinburgh
scientific society, the Plinean society, of which Grant was a prominent member, and presented two
lectures that reported discoveries he had made while working with Grant. This interest in marine
invertebrates was to be a life long obsession, climaxing in his massive four-volume contribution to the
comparative anatomy and systematics of fossil and living Cirripedia or barnacles. (Barrett & Freeman
1988, vols. 1113)
When he finally broke the news of his distaste for medicine to his father, he enrolled to take a degree in
Divinity at Christ College, Cambridge University, from which he graduated in January of 1831. As with
the Shrewsbury School and Edinburgh, his official course of study had very little impact on him, but
while in Cambridge he befriended two young men attempting to institute serious reforms in the natural
science curriculum at Cambridge, Rev. John Henslow, trained in botany and mineralogy, and Rev.
Adam Sedgwick, a leading member of the rapidly expanding community of geologists. Henslow and
his wife treated Darwin almost as a son, and through Henslow Darwin was introduced to the men
whose ideas were currently being debated in geology and natural history, as well as to men whom we
look back on as among the very first to take up the historical and philosophical foundations of science
as a distinct discipline, Sir John Herschel and Rev. William Whewell. As he wrote in his autobiography:
During my last year at Cambridge, I read with care and profound interest
Humboldt'sPersonal Narrative. This work, and Sir J. Herschel's Introduction to the Study
of Natural Philosophy,[4] stirred up in me a burning zeal to add even the most humble
contribution to the noble structure of Natural Science. No one or a dozen other books
influenced me nearly so much as these two.

In the next section we will discuss the influence of the philosophical ideals of Herschel and Lyell on
Darwin.
Furthering his scientific training, Adam Sedgwick on two occasions took Darwin on extended
geological tours of England and Wales. In addition Darwin and a cousin, William Darwin Fox, a year
ahead of him at Cambridge, developed what began as an amateur passion for bug collecting into
serious entomology.

2.2 Darwin's Darwinism

His Edinburgh and Cambridge mentors were to shape Darwin's philosophical attitudes and scientific
career decisively. It was Henslow who was the final link to Darwin in a chain connected to Captain
Robert Fitzroy of H. M. S. Beagle. Fitzroy sought a gentleman companion who could also collect
information on geology and natural history during a proposed circumnavigation of the globe.
Henslow's note to Darwin, asking if he would be interested in being recommended for this post, arrived
at the Darwin home, the Mount, while Charles Darwin was on a geological survey of Northern Wales
with Adam Sedgwick. After resistance from his father had been overcome, Darwin was offered the post
and accepted it.
The combination of meticulous field observation, collection and experimentation, note taking, reading
and thinking during what turned into the Beagle's five year journey through a very wide cross-section
of the earth's environments was to set the course for the rest of his life. During the voyage he read and
reread Charles Lyell's newly published Principles of Geology, a three-volume work that articulated a
philosophical vision of rigorously empirical historical science, oriented around five key ideas:
1. The geologist investigates both the animate and inanimate changes that have taken place during
the earth's history.
2. His principal tasks are to develop an accurate and comprehensive record of those changes, to
encapsulate that knowledge in general laws, and to search for their causes.
3. This search must be limited to causes that can be studied empiricallythose now in operation,
as Lyell puts it in the sub-title of his Principles.
4. The records or monuments of the earth's past indicate a constant process of the introduction
and extinction of species, and it is the geologist's task to search for the causes of these
 introductions and extinctions, according to the strictures note in 3., above.
5. The only serious attempt to do so according to the idea that species are capable of indefinite
modification, that of Jean Baptiste Lamarck, is a failure on methodological grounds. All the
evidence supports the view that species variability is limited, and that one species cannot be
transformed into another.
This vision influenced Darwin profoundly, as he freely admitted. While he became convinced by his
observations and reading that the fossil record and current distribution of species could only be due to
the gradual transformation of one species into another, he was determined to articulate a theory that
measured up to Lyell's principles. The crucial event in convincing him that this was to be his life's work
was likely a visit to Cape Town, South Africa during the Beagle's return trip to England. John F. W.
Herschel was in Cape Town on a mission to do for the Southern Hemisphere what his father William
had done for the Northern, namely to develop a comprehensive star map with the new powerful
telescopes developed by his father and aunt. As noted earlier, Darwin had been deeply impressed by
Herschel's Preliminary Discourse on the Study of Natural Philosophy when it first appeared a year
before the Beagle set sail, and in his private journal he referred to his meetings with Herschel during a
week long stop in Cape Town in June of 1836 as among the most profound events of the entire voyage.
Just five months before meeting Darwin, Herschel had finished reading the 2ndedition of
Lyell's Principles. He sent Lyell a long letter filled with detailed constructive commentary. The letter
opens by praising Lyell for facing the issue of the introduction of new species which Herschel calls
that mystery of mysteries scientifically, and for advocating that we search for intermediate causes
to explain these introductionscode for natural, as opposed to miraculous, causes.[5] This part of
the letter was quoted in Charles Babbage's Bridgewater Treatise, published in 1837 while Darwin was
struggling to develop just such a theory. Upon reading the Herschel quotation in Babbage, Darwin
wrote in his private species notebooks:
Babbage 2d Edit, p. 226.Herschel calls the appearance of new species. the mystery of
mysteries. & has grand passage upon problem.! Hurrah.intermediate causes. (Barrett et
al., 1987, 413; original punctuation)

He clearly recognizes that Herschel is here providing a philosophical justification for the project upon
which Darwin was secretly working. And, in the very first paragraph of On the Origin of Species,
Darwin looks back to this Hurrah, attributing the idea that the origin of species is that mystery of
mysteries to one of our greatest philosophers, without mentioning Herschel by name. The first
mention of the possibility of an evolutionary solution to this problem is in his Ornithological
Notebooks, in a note written shortly after departing Cape Town.[6]
Darwin's theoretical task was, by the time he opened his species notebooks, tolerably clear: the only
process that could produce the systematic patterns in the fossil record and the otherwise strange
biogeographic distribution of species he now understood so widely and deeply was a process of slow,
gradual transformation of species. He needed to come up with a natural, causal theory that would
account for such transformations, and every element of that theory had to identify causes now in
operation, causes that could be investigated empirically. The problem, and the methodological
constraints, had been advocated by his geological hero, and now close friend, Charles Lyell; and they
had been defended philosophically by his philosophical hero, Sir John Herschel.
Darwin, of course, expected, and got, outraged reactions from religiously conservative colleagues, such
as his old geology teacher Sedgwick, who in a review expressed his deep aversion to the theory;
because of its unflinching materialism;--because it has deserted the inductive track,--the only track that
leads to physical truth;--because it utterly repudiates final causes, and therby [sic] indicates a
demoralized understanding on the part of its advocates. What he had not expected was Lyell's refusal
to openly endorse his theory and Herschel's decisive (if polite) rejection of its key elements. After we
set out the theory in its Darwinian form, we can consider these reactions from those who apparently
shared Darwin's philosophical norms about scientific theory, explanation and confirmation.
The theory can be set out as a series of causal elements that, working together, will produce the needed
transformations.
1. Species are comprised of individuals that vary ever so slightly from each other with respect to
their many traits.
2. Species have a tendency to increase in numbers over generations at a geometric rate.
3. This tendency is checked, to use the language of Thomas Malthus' On the Principle of
Population, by limited resources, disease, predation, and so on, creating a struggle for survival
among the members of a species.
4. Some individuals will have variations that give them a slight advantage in this struggle,
variations that allow more efficient or better access to resources, greater resistance to disease,
greater success at avoiding predation, and so on.
5. These individuals will tend to survive better and leave more offspring.
6. Offspring tend to inherit the variations of their parents.
7. Therefore favorable variations will tend to be passed on more frequently than others and thus be
preserved, a tendency Darwin labeled Natural Selection.
8. Over time, especially in a slowly changing environment, this process will cause the character of
species to change.
9. Given a long enough period of time, the descendant populations of an ancestor species will
differ enough both from it and each other to be classified as different species, a process capable
of indefinite iteration. There are, in addition, forces that encouragedivergence among
descendant populations, and the elimination of intermediate varieties.
It will be noticed that there is no element of this theory that is incapable of empirical investigation
indeed by now the published confirmatory studies of this process would fill a small library.[7] One can
understand why devout and orthodox Christians would have problems; but why Darwin's philosophical
and scientific mentors? It would seem to be the model of Herschelian/Lyellian orthodoxy.

2.3 Philosophical Problems with Darwin's Darwinism

The answer lies in five philosophically problematic elements of the theory.

2.3.1 Probability and Chance

First, notice the use of the language of tendencies and frequencies in the above principles. Privately,
Darwin learned, Herschel had referred to his theory as the Law of higgledy-piggledy, presumably a
reference to the large element played in its key principles by chance and probability. Darwin's theory is,
as we would say today, a statistical theory. One cannot say that every individual with favorable
variation v will survive or will leave more offspring than individuals without it; one cannot say that no
environment will ever support all of the offspring produced in a given generation, and thus that
there must always be a competitive struggle. These are things that tend to happen due to clearly
articulated causes, and this allows us to make accurate predictions about trends, at the level of
populations, but not to make absolute claims about what must happen in each and every case. Only
well after Herschel's time did philosophers of science become comfortable with the idea of a theory of
this sort, and the proper philosophical understanding of such explanations is still debated.

2.3.2 The Nature, Power and Scope of Selection

For many people natural selection is the core of Darwin's theory. Perhaps because of his use of the
concept of selection, the core element of Darwin's theory seems to have baffled nearly everyone. Could
it be, as Lyell, Herschel and Darwin's great American defender Asa Gray would ask, an intermediate
cause, i.e. a causal principle instituted and sustained by God? Or is it in its very nature the antithesis of
such a principle, as his old geology teacher Sedgwick believed? Could it possibly create species, or is it
by its nature a negative force, eliminating what has already been created by other means? In one of his
copies of On the Origin of Species, Alfred Russell Wallace crosses out natural selection and writes
survival of the fittest next to it. Wallace always felt that selection inappropriately imported
anthropomorphic notions of Nature choosing purposefully between variants into natural history. And, in
a devastating review Fleeming Jenkin happily accepted the principle of natural selection but challenged
its power to modify an ancestral species into descendent species, and thus limited its scope to the
production of varieties. A number of reviewers, even some sympathetic ones, questioned the possibility
of extending the theory to account for the evolution of those characteristics that differentiate humans
from their nearest relatives.

2.3.3 Selection, Adaptation and Teleology

Moreover, because Darwin was very fond of describing natural selection as a process that worked for
the good of each species, Darwin's followers seemed to have diametrically opposed views as to
whether his theory eliminated final causes from natural science or breathed new life into them. In either
case, there was also serious disagreement on whether this was a good thing or a bad thing.[8]

2.3.4 Nominalism and Essentialism

There is a fundamental philosophical problem with the idea that a species can undergo a series of
changes that will cause it to become one or more other species. To illustrate it, look carefully at the first
question that Charles Lyell wishes to address in the second volume of thePrinciples of Geology:
first, whether species have a real and permanent existence in nature; or whether they are
capable, as some naturalists pretend, of being indefinitely modified in the course of a long
series of generations. (Lyell 1831, II. 1)

Lyell pretty clearly assumes that to allow for evolution is to deny the reality of species. For a species to
be real, it must have permanent existence in nature, or as he puts it elsewhere , fixed limits
beyond which the descendants from common parents can never deviate from a certain type. (Lyell
1831, II. 23) To accept evolutionary change, on this view, you must become comfortable with a variety
of nominalism about species. And Darwin seems to have become so.[9]
Hence I look at individual differences, though of small interest to the systematist, as of high
importance for us, as being the first step towards such slight varieties as are barely thought
worth recording in works on natural history. And I look at varieties which are in any degree
more distinct and permanent, as steps leading to more strongly marked and more permanent
varieties; and at these latter, as leading to sub-species, and to species. (Darwin 1859, 52)

Permanence for Darwin is a relative thing, and there are no fixed limits to variability within a species.
Given enough time the individual differences found in all populations can give rise to stable varieties,
these to sub-species, and these to populations that systematists will want to class as distinct species.
Moreover, he concludes the Origin with very strong words on this topic, words bound to alarm his
philosophical readers:
Systematists will be able to pursue their labours as at present; but they will not be
incessantly haunted by the shadowy doubt whether this or that form be in essence a species.
In short, we will have to treat species in the same manner as those naturalists treat
genera, who admit that genera 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. (Darwin 1859, 485)

Lyell, Herschel, Whewell, Sedgwick and many of Darwin's contemporaries certainly would not find
this a cheering prospect, since they were unrepentant essentialists about species.[10]Members of a
species possess a type established in the original parents, and this type provides fixed limits to
variability. Lyell clearly feels this is an empirically verifiable resultmost of chapters 24
of Principles Vol. II is devoted to collecting the evidence that such fixed limits exist; and after
the Origin's publication this evidence was canvassed again in Fleeming Jenkin's review. If this is so,
then species extinction is easy to account forthere are fixed limits to a species ability to track
environmental change. But a naturalistic account of species origination is more difficult, since there
will need to be, in sexually reproducing species, a natural production of a new pair of parents with a
new type. On the other hand, to adopt the sort of nominalism advocated above by Darwin seems to
have undesirable consequences as well. How are we to formulate objective principles of classification?
What sort of a science of animals and plants will be possible if there are no fixed laws relating their
natures to their characteristics and behaviors? A good deal of chapter 2 of Darwin's Origin is devoted to
convincing the reader that current best practice among botanists and zoologists accepts a natural world
organized as he is insisting rather than as his opponents claim:
It must be admitted that many forms, considered by highly competent judges as varieties,
have so perfectly the character of species that they are ranked by other highly competent
judges as good and true species. (Darwin 1859, 49)

From a Darwinian perspective, this is a predictable consequence of the fact that the organisms we today
wish to classify are merely the most recent stage of a slow, gradual evolutionary process. Organisms
within a genus have common ancestors, perhaps relatively recent common ancestors; some naturalists
may see ten species with a few varieties in each; others may rank some of the varieties as species and
give the same genus twenty species. Both classifications may be done with the utmost objectivity and
care by skilled observers. As systematists like to say, some of us are lumpers, some of us are
splitters. Reality is neither.

2.3.5 Tempo and Mode of Evolutionary Change

The question of nominalism regarding species points toward a final aspect of Darwin's theory with
which many of those otherwise sympathetic to him disagreed, his gradualism. For apart from the
question of whether his views entailed nominalism about natural kinds, they do seem to reflect a
belief that the evolutionary process must be a slow and gradual one. It is perhaps here that we see the
most lasting impact of Darwin's careful study of Charles Lyell'sPrinciples of Geology while on H.M.S.
Beagle. I stress slow and gradual, for it is clear that one could have a slow but non-
gradual evolutionary process (perhaps the long periods of evolutionary stasis punctuated by
geologically rapid periods of speciation postulated by Eldridge and Gould's punctuated equilibrium
model are such); and one could have a rapid but gradual one (for example the process George Gaylord
Simpson labeled adaptive radiation, where a population migrates to a location with a variety of
unexploited niches, and rapidly evolves to exploit them). Darwin stresses over and over again that he
conceives of natural selection adding up infinitely small variations, and that he imagines the process
of speciation to take place over a very long period of time.
One of the strongest arguments for insisting that Darwinism as it is used today is isomorphic to
Darwin's Darwinism, as Gayon puts it, is that each of these questions is still hotly debated, and has
been throughout the theory's history. With all of the amazing changes that have been wrought by the
genetic, biochemical, and molecular revolutions, with the development of mathematical models of
population genetics and ecology, of sophisticated techniques for both field and laboratory investigation
of evolutionary processes, and of cladistic analysis in systematics, it nevertheless remains true that one
can find evolutionary biologists who adhere to Darwin's Darwinism, and are recognized as doing so by
both themselves and their critics. In the next section of this article, I will develop a portrait of
contemporary Darwinism around each of these contested features.
By the same token, however, Darwinism has evolved. As one example of this truth, think for a moment
of contemporary debates about the nature of selection. The problems people had with natural selection
in the 19th century continue to be problematic, but there are a variety of problems that were either not
discussed, or discussed very differently, in the 19th century. Can, and does, natural selection work at
levels other than the level of Darwin's focus, individual organisms; is there a non-vacuous way to
formulate the theory abstractly; how are we to understand the relationships between the concepts of
fitness, selection and adaptation? How strong are the constraints on the selection process, and what
sorts of constraints are there? Are there other motors of evolutionary change besides selection, and if
so, how important are they? In particular, how important is drift, and how are we to differentiate it
from selection?
3. The Five Core Philosophical Problems Today
"Theories need both essences and histories."
Stephen Jay Gould (2002, 1)
So reads the heading of the very first section of the first chapter of Gould's monumental The Structure
of Evolutionary Theory. Opening with a subtle reading of an exchange of letters in 1863 between
paleontologist Hugh Falconer and Charles Darwin, Gould eventually explains what he has in mind by
this section heading:
In short, The structure of evolutionary theory combines enough stability for coherence
with enough change to keep any keen mind in a perpetual mode of search and challenge.
(Gould 2002, 6)

Gould, of course, was both an unabashed admirer of Charles Darwin and one of the most outspoken
critics of the neo-Darwinian synthesis. I will be using both his account of the Essence of Darwinism
in Part I of this magnum opus and his arguments for a Revised and Expanded Evolutionary Theory in
its Part II as touchstones and targets.
In the preceding section of this essay, I organized my discussion of the problems that Darwin's allies
had with Darwin's Darwinism around five issues: [i] the role of chance as a factor in evolutionary
theory and the theory's apparently probabilistic nature; [ii] the nature of selection; [iii] the question of
whether selection/adaptation explanations are teleological; [iv] the ontological status of species and the
epistemological status of species concepts; and [v] the implications of Darwin's insistence on the slow
and gradual nature of evolutionary change. I claimed that one very good reason for continuing to
characterize one dominant approach to evolutionary biology, that represented by the so-called Neo-
Darwinian Synthesis, as Darwinism is that its proponents side with Darwin on these issues (and on
many less fundamental ones besides). That in itself is remarkable, but it is the more so because the
Darwinian position on each of these issues is under as much pressure from non-Darwinian evolutionary
biologists today as it was in the wake of the Origin. It is not surprising, given the situation as I have
just characterized it, that philosophers of biology have made significant contributions to the discussion,
especially in pointing out underlying philosophical issues that are at stake and conceptual confusions
and ambiguities that stand in the way of resolving the issues at hand.
It is my conviction that a full understanding of the underlying philosophical disagreements on these
questions will only come from a patient historical study of how the Synthesis positions on these
various issues, and those of their critics, arose. That I cannot do here. Rather, in what follows I will
simply be presupposing certain answers to these questions of historical origins. The list of references at
the end of this essay includes a number of excellent pieces of work on this subject for those who share
my convictions about its importance.

3.1 The Roles of Chance in Evolutionary Theory

The evolutionary process, as Darwin understood it, involves the generation of variation and a process
producing a differential perpetuation of variation. One simple way to think about Darwinism in relation
to a logical space of alternatives, then, is by means of the followingvariation grid:
 Variations
Generation Perpetuation
Lamarck Darwin
Fitness Biased
Asa Gray Asa Gray
Darwin Lamarck
Not Fitness Biased
Neutralism Neutralism
The above grid might lead you to conclude that both non-fitness biased generation of variation and
non-fitness biased perpetuation of variation would be properly labeled chance. By seeing why that
would be a misleading conclusion to draw, we get to the heart of the problem of the concept of chance
in contemporary Darwinism.
Let us begin with the language Darwin uses when he first sketches his theory at the beginning of the
fourth chapter of the Origin:
Can it, then, be thought improbable, seeing that variations useful to man have undoubtedly
occurred, that other variations useful in some way to each being in the great and complex
battle of life, should sometimes occur in the course of thousands of generations? If such do
occur, can we doubt (remembering that many more individuals are born than can possibly
survive) that individuals having any advantage, however slight, over others, would have the
best chance of surviving and of procreating their kind? (Darwin 1859, 8081)

Unlike Darwin's contemporaries, the founders of the synthesis of Mendelian genetics and Darwinian
selection theory, Sewall Wright, Ronald Fisher and J. B. S. Haldane, were entirely comfortable with a
selection theory formulated in such terms. On this issue contemporary Darwinism agrees whole-
heartedly with Charles Darwin. Note one clear statement of thePrinciple of Natural Selection from the
philosophical literature:
If a is better adapted than b to their mutual environment E, then (probably) a will have
greater reproductive success than b in E. (Brandon1990, 11).

The theory trades pervasively in probabilities. To take a simple case: if there are three possible
combinations of alleles at a given locus in a population, we can characterize the outcome of a
reproductive cycle as chance if each of the three possible combinations occurs at a frequency
determined strictly by the laws of probability. In any given case of reproduction, we would say, which
genotype emerged is a matter of chance. Given the fact that evolutionary biologists, especially in so far
as they take their cues from population genetics, deal with large populations conceived as gene pools,
and think of evolution as long run changes in the frequencies of different combinations of genes from
generation to generation, it is clear that, in this sense, chance permeates contemporary Darwinism. The
models of population biology provide a means of assigning probabilities to various outcomes, given
information about population size, rates of mutation and migration (themselves given as averages and
estimates). That is, as Darwin notes, being relatively better adapted increases an organism's chances,
i.e. increases its probability, of leaving viable offspring. It does not guarantee it. Since natural selection
is a stochastic process, Darwinians from Darwin to the present rightly characterize it in terms of
influencing the chances of a given outcome, given variables such as selection pressure, population
size or mutation rate.
Conceptual confusion arises, however, from the fact that chance and randomness are often
contrasted, not with deterministic outcomes but with selected outcomes. For example, when John
Beatty describes random drift as changes in frequencies of variations due to chance in the following
passage, he presumably has something like a contrast with changes in frequencies due to selection in
mind.
In Darwin's scheme of things, recall, chance events and natural selection were consecutive
rather than alternative stages of the evolutionary process. There was no question as to
which was more important at a particular stage. But now that we have the concept of
random drift taking over where random variation leaves off, we are faced with just such a
question. That is, given chance variations, are further changes in the frequencies of those
variations more a matter of chance or more a matter of natural selection? (Beatty 1984,
196)

Notice that in the above quote we first get a substitution of random for chance in the phrases
random variation and chance variation, and then at least the suggestion that the concept of random
drift can be characterized as changes in frequencies of variations due to chance, where the contrast
class consists of similar changes due to natural selection.
With respect to the generation of variation, chapter 5 of On the Origin of Species opens with the
following apology:
I have hitherto sometimes spoken as if the variationsso common and multiform in
organic beings under domestication, and in a lesser degree in those in a state of naturehad
been due to chance. This, of course, is a wholly incorrect expression, but it serves to
acknowledge plainly our ignorance of the cause of each particular variation. (Darwin 1859,
131)

Here Darwin is noting that, though to speak of chance variations may seem to be citing chance as the
cause of the variations, in fact it is simply acknowledging that they appear to have no assignable
cause. But it is important to keep historical context in mind here. Whether Darwin himself ever flirted
with the idea of directed variation or not, he was acutely aware of two views from which his needed
to be distinguished, very different from each other, but both holding to the view that variations arose
for a purpose.[11] The most widely shared alternative was that found in natural theology. To quote the
Reverend William Paley's Natural Theology, regarding a beautiful instance of adaptation: A
conformation so happy was not the gift of chance. Likewise, among Darwin's followers, the American
botanist Asa Gray, in an essay entitled Natural Selection and Natural Theology, uses the same contrast
to advise Darwin against the notion of chance variation: we should advise Mr. Darwin to assume,
in the philosophy of his hypothesis, that variation has been led along certain beneficial lines.
Gray is here insisting that, since Darwin admits that using the term chance merely signals ignorance
of the true cause, and since the pervasive adaptations in nature suggest design, Darwin should avoid the
suggestion that variations are due to chance in the sense of absence of design.[12]
Darwin, in fact never refers to chance variations in the Origin, though occasionally he will note that if
a beneficial variation chances [i.e. happens] to appear, it will be favored by selection (see pp. 37, 82)
What Darwin has in mind, however, is clear from his concluding remarks in his chapter on Laws of
Variation:
Whatever the cause may be of each slight difference in the offspring from their parents
and a cause of each must existit is the steady accumulation, through natural selection, of
such differences, when beneficial to the individual, that gives rise to all the more important
modifications of structure (Darwin 1859, 170)

Whatever the cause of the generation of a variation may be, the role of selection is toaccumulate those
already present variations that happen to be beneficial. As Beatty put it, the generation of variations and
their selection are consecutive processes. But to call the generation of variation a chance process is
to use chance in this second sense, meaning not by design or for some end.
Apart from those urging Darwin to give up chance in favor of design, he had pressure to abandon
chance from another direction, the evolutionary philosophy of Jean-Baptiste Lamarck. Lamarck's is
a materialistic argument against the variation in nature being a matter of chance. On the Lamarckian
view, variations arise in an organism as a direct response to environmental stress or demand, giving rise
to a stimulus, which in turn elicits a physiological response, which finally can be passed on via
reproduction to offspring. Variations are not chance or random, since they are an appropriate response
to an environmental stress. Here chance signals a lack of relation or connection to adaptive needs, an
idea akin to, but ontologically quite distinct from, the contrast between chance and design.
The concept of random variation is today often used as a synonym for chance variation in precisely
this latter sense. Here are two examples of this notion of chance or randomness as used by
contemporary Darwinians.
mutation is a random process with respect to the adaptive needs of the species.
Therefore, mutation alone, uncontrolled by natural selection, would result in the breakdown
and eventual extinction of life, not in adaptive or progressive evolution. (Dobzhansky 1970,
65)

Thus the production of variations may be a chance process in that there are a number of possible
outcomes with assignable probabilities, but it is also a chance process in the sense that the probability
assignments are not biased by adaptive needs or fitness.
My second example is intended to take us back to problems with our first sense of random and
chance. Here, a champion of the neutral theory of molecular evolution characterizes his position:
the great majority of evolutionary changes at the molecular (DNA) level do not result
from Darwinian natural selection acting on advantageous mutants but, rather, from random
fixation of selectively neutral or very nearly neutral mutants through random genetic drift,
which is caused by random sampling of gametes in finite populations. (Kimura 1992, 225)
Here, it will be noticed, the focus is not on the generation of variations but on theperpetuation of
variations. The contrast is between a random sampling of gametes that leads to the fixation of
selectively neutral alleles and natural selection favoring advantageous variations. That is, the contrast
between chance and fitness biased processes is now being used to distinguish means of perpetuating
certain variations. We are contrasting two sampling processes. Drift samples without concern for
adaptation; selection samples discriminately on the basis of differences in fitness. Both samplings are
probabilistic, of course, but that in no way obviates the above contrast.
However, as Beatty has pointed out, it was quite common until fairly recently to characterize natural
selection in such a way as to make it almost indistinguishable from random drift (cf. Lennox 1992,
Lennox and Wilson 1994). Numerous accounts of fitness characterized the fitness of a genotype as
defined by its relative contribution to the gene pool of future generationsthe genotype contributing
the larger percentage being the fitter. But of course that could easily be the result of a randomnon-
fitness biased-- sampling process; which organisms would be declared fitter by this method might
have nothing to do with natural selection. In order to provide a proper characterization of the role of
chance in evolutionary change, then, it is critical to provide a more robust and sophisticated account of
fitness. (For further information, see the entry on: fitness.) This, in turn, requires that we discuss the
conceptual network that includes the notions of adaptation and natural selection, to which we will turn
shortly.
For now, let us assume that there is a way of characterizing fitness such that there is a substantial
empirical question of what role indiscriminate sampling of genotypes (or phenotypes) plays in
evolutionary change. This issue was first placed squarely before evolutionary biologists by Sewall
Wright in the early 1930s. As Wright pointed out, genes that are neutral with respect to fitness can, due
to the stochastic nature of any process of sampling from a population, increase their representation
from one generation to the next. The likelihood of this happening goes up as effective population size
goes down. Since Wright imagined that a quite typical scenario in evolutionary change was for species
to be broken up into relatively small, relatively isolated, populations (or demes), with significantly
more breeding within than between demes, the likelihood that such neutral genotypes could become
fixed at relatively high levels was significant. Though he gradually toned down this aspect of his work,
a significant school of mathematical population geneticists in the 1960s and 70s took these ideas and
ran with them, developing a Neutralist approach to evolutionary change. This is the position
characterized by Kimura (one of its most eloquent defenders) in the passage quoted above. Whether or
not such a process plays a significant role in evolution is not a philosophical issue, but it is highly
relevant to whether evolutionary biology should be seen as predominantly Darwinian. For if any view
is central to Darwinism, it is that the evolutionary process is predominantly guided by the fitness-
biasing force of natural selection, acting on variations that arise by chance. It is to natural selection and
related concepts that we now turn.

3.2 The Nature, Power and Scope of Selection

The greatest number of females will, of course, fall to the share of the most vigorous males;
and the strongest individuals of both sexes, by driving away the weakest, will enjoy the best
food, and the most favourable situations, for themselves and for their offspring. A severe
 winter, or a scarcity of food, by destroying the weak and the unhealthy, has had all the good
effects of the most skilful selection.

The words of Charles Darwin? No; these are the words of John Sebright, penned in The Art of
Improving the Breeds of Domestic Animals in 1809, the year of Charles Darwin's birth and fifty years
before On the Origin of Species was published. Darwin refers to this passage in Notebook C of his
Species Notebooks.[13] It will be noticed that Sebright is not discussing domestic selection, but is quite
clearly saying that processes leading to differential survival and reproduction in nature will have all
the good effects of the most skilful selection. Darwin, then, did not need to read Malthus to see what is
here so plainly and clearly statednamely, that the struggle for survival in nature will have the same
selective effects as the actions of the domestic breeder of plants and animals.
As this passage, and the argument of the Origin, shows, natural selection began life as the product of
analogical reasoning. Sebright sees clearly that the natural processes he is describing will have the
same effects as the breeder's selection, but he is not about to describe those processes as selection
processes. Darwin took that step, and Darwinism has followed.
Darwin himself consistently refers to natural selection as a power of preserving advantageous, and
eliminating harmful, variations. As noted in the last section, whether a particular variation is
advantageous or harmful is, in once sense of that term, a matter of chance; and whether an
advantageous variation is actually preserved by selection is, in another sense of the term, also a matter
of chance. For Darwinism, selection is the force or power that biases survival and reproduction in favor
of advantageous variations, or to look ahead to the next section, of adaptations. It is this that
distinguishes selection from drift.
In a recent monograph entitled Natural Selection: Domains, Levels and Challenges in theOxford Series
in Ecology and Evolution, George C. Williams has vigorously defendedDarwinian selection
theory against a variety of challenges that have emerged over the last few decades. Those challenges
can be placed into two broad categories: [i] proposed limitationson natural selection as an evolutionary
force; and [ii] expansions of the scope of natural selection to include new targets and levels. It will
be noted that in neither case is it obvious that the theory itself requires modification in the face of such
challengesin principle these might be nothing more than challenges to the theory's range of
application. However, if it turned out that most evolutionary change could be explained without
recourse to natural selection, this would be grounds for arguing that evolutionary biology was no longer
Darwinian. And if it turned out that the theory of natural selection could only be integrated with our
new understanding of the processes of inheritance and development by a wholesale modification of its
foundations, it might be best to see the new theory as a modified descendent of Darwinism, rather than
Darwinism itself. Theories may need essences, as Gould claims; but if what is fundamental to the
theory has changed, then so has its essence. To borrow a phrase from Paul Griffiths, perhaps it is not
that theories need histories andessencesperhaps what they need are historical essences.
Alfred Russell Wallace regularly urged Darwin to jettison the term selection as misleadingly
anthropomorphic, and substitute Herbert Spencer's survival of the fittest. Darwin went half wayin
later editions he added or Survival of the Fittest to Natural Selection in the title of chapter 4. As the
theory developed in the mid-20th century, the expression survival of the fittest was gradually
eliminated from any serious presentation of Darwinian selection theory. On the other hand, the concept
of fitness has played a prominent, and problematic, role. In the mathematical models used in
population genetics, fitness refers either to the abilities of the different genotypes in a population to
leave offspring, or to the measures of those abilities, represented by the variable W. Here is a rather
standard textbook presentation of the relevant concepts:
In the neo-Darwinian approach to natural selection that incorporates consideration of
genetics, fitness is attributed to particular genotypes. The genotype that leaves the most
descendants is ascribed the fitness value W=1, and all other genotypes have fitnesses,
relative to this, that are less than 1. Fitness measures the relative evolutionary advantage
of one genotype over another, but it is often important also to measure the relative penalties
incurred by different genotypes subject to natural selection. This relative penalty is the
corollary of fitness and is referred to by the term selection coefficient. It is given the
symbol s and is simply calculated by subtracting the fitness from 1, so that: s = 1 W.
(Skelton 1993 164)

The problem lies in the fact that the concept of fitness plays dual roles that are instructively conflated in
this quotation. For when fitnesses are viewed as measures of differential abilitiesof organisms with
different genotypes to leave different numbers of offspring, the language of fitness encourages us to
suppose that fitness refers to the relative selective advantages of genotypes. On the other hand, if
fitness simply refers to the measure of reproductive success, it is a quantitative representation of small
scale evolutionary change in a population, and leaves entirely open the question of the causes of the
change. But then the assumed connections among the concepts of fitness, adaptation and natural
selection are severed. Selection coefficients may have nothing to do with selection; what W represents
may have nothing to do with selective advantage.
There is, however, a way of formulating the theory in its modern guise which maintains an essentially
Darwinian character. Since there are a number of confirmed ways in which natural populations can
evolve in the absence of natural selection, and since balancing selection, i.e. countervailing selection
forces, may prevent a population from evolving in its presence, it is clear that establishing, by
measuring different reproductive rates among its members, that the genetic make-up of a population
has changed does not establish that natural selection was the source of that change; nor does the fact
that no change has been measured establish that natural selection is not operative. Population genetics
and its associated models should be treated as the kinematics, not the dynamics of evolutionary
processes. That is, it is a way of establishing that a population either is or is not in equilibrium, and it
provides sophisticated tools for measuring rates of change in a population across generations.
Moreover, like the kinematics of any physical theory, if it establishes cross-generational change, it also
tells us that there are causes to be foundthe detailed contours of those measures may even provide
suggestions as to where to look for those causes. What it cannot do on its own is provide knowledge of
the forces at work. To use language introduced by Elliott Sober, fitness, unlike natural selection,
is causally inert. (For further information, see the entry on: population genetics.)
That means that, as valuable as population genetics is, it should not be equated with the theory of
natural selection. Too often in both biological presentations of the theory and philosophical discussions
of it, this is forgotten. For example:
Most people are familiar with the basic theory of natural selection. Organisms vary in a
heritable fashion. Some variants leave more offspring than others; their characteristics,
therefore, are represented at a greater frequency in the next generation. (Wilson 1984, 273)

This is a presentation of the basic theory of natural selection that makes no reference to natural
selection at all!
Natural selection, if it is to resemble the Darwinian concept that bears that name, must be reserved for
reference to an interaction between a variable, heritable feature of an organic system and the
environment of that system. That interaction may or may not change the proportions of those features
across generations, and those proportions may change for reasons other than those interactions. But a
plausible natural selection hypothesis must posit some such interaction. On this issue I will give the last
word to Stephen Jay Gould:
when we consider natural selection as a causal process, we can only wonder why so
many people confused a need for measuring the results of natural selection by counting the
differential increase of some hereditary attribute (bookkeeping) with the mechanism that
produces relative reproductive success (causality). (Gould 2003, 619)

The concept of natural selection has to this point been presented broadly because of the other range of
critical questions surrounding the contemporary Darwinian concept of natural selection that I
mentioned earlierquestions having to do with possible limiting constraints on natural selection and
about the sorts of objects that can be viewed as appropriate organismic/environmental interactors in
the selection process.
If we suppose that for Darwin natural selection was almost exclusively thought of as an interaction
between individual organisms and their organic and inorganic environments, then we can see two
challenges to Darwinism today with respect to levels of selection. There are those, such as G. C.
Williams and Richard Dawkins, who argue that selection is always and only of genes. Here is a clear
statement:
These complications [those introduced by organism/environment interactions] are best
handled by regarding individual [organismic] selection, not as a level of selection in
addition to that of the gene, but as the primary mechanism of selection at the genic level.
(Williams 1993, 16)

Dawkins preferred mode for making the same point is to refer to organismsor interactors--as
the vehicles of their genes, in fact vehicles constructed by the genome for its own perpetuation.
The original impulse for this approach, especially clear in Williams classic Adaptation and Natural
Selection (1966) was philosophicalit was to use a sort of Ockham's razor strategy against Group
Selection hypotheses, showing that alleged group selection effects could be explained by explanations
operating at the level of the genome. Throughout that book selection is always said to be of individual
alleles, regardless of the role environments at various levels may play in the process.
This view has been extensively challenged by philosophers of biology on both methodological and
conceptual grounds, though there are, among philosophers, enthusiastic supporters (cf. Dennett 1995).
In all the give and take, it is seldom noticed that defenders of this view claim to be carrying the
Darwinian flag (Gayon 1998 and Gould 2003 are exceptions). Yet it is certainly not a position that
Darwin would recognize--and not merely because he lacked a coherent theory of the units of
inheritance. It is not a Darwinian view because for Darwin it was differences in the abilities of
organisms at various stages of development to respond to the challenges of life that had causal primacy
in the explanation of evolutionary change. Among evolutionary biologists from the neo-Darwinian
synthesis on, it is those who stress the role of organisms in populations interacting differentially to
ever-variable ecological conditions in causing changes in the gene pools of those populations who are
the card-carrying Darwinians.
Darwinism also has challenges from the opposite direction. In the 1970s a number of biologists
working in the fields of paleontology and systematics challenged the Neo-Darwinian dogma that you
could account for macro-evolution by simple, long term extrapolation from micro-evolution. Gould,
in particular, opens Part II of The Structure of Evolutionary Theory (Towards a Revised and Expanded
Evolutionary Theory), with a chapter entitled Species as Individuals in the Hierarchical Theory of
Selection. That chapter title combines two conceptually distinct theses: first, the thesis defended by
Michael Ghiselin (Ghiselin 1997) and championed and refined by David Hull (Hull 2001), that species
are in a robust sense of the term individuals; and second, that there may well be selection among
groups of organisms, qua groups. Gould's title exemplifies one approach to group selectionthe unit of
selection is always the individual, but there are individuals other than individual organisms that are
subject to selection. A very different result emerges if one assumes that groups of organisms such as
demes, kin-groups, or species, though not individuals, are nevertheless subject to selection. Adding to
the conceptual complexity, some researchers propose that the term group selection be restricted to the
process whereby group-level traits provide advantages to one group over another, in which case there
are strict conditions delimiting cases of group selection. Others define group selection primarily in
terms of group level effects. Thus a debate analogous to that earlier discussed regarding the definitions
of fitness emerges hereby group selection do we mean a distinct type of causal process that needs to
be conceptually distinguished from selection at the level of individual organism or gene, or do we
merely mean a tendency within certain populations for some well defined groups to displace others
over time? (For further discussion, see Sterelny and Griffiths 1999, 151179; Hull 2001, 4990; and
see the entry on: levels and units of selection.)

3.3 Selection, Adaptation and Teleology

Early in the introduction to On the Origin of Species, Darwin observes that the conclusion that each
species had descended from others even if well founded, would be unsatisfactory, until it could be
shown how the innumerable species inhabiting this world have been modified so as to acquire that
perfection of structure and co-adaptation which most justly excites our admiration (Darwin 1859, 3).
One might say this was the central promise of Darwinismto account for both phylogenic
continuity and adaptive differentiation by means of the same principles; or as Darwin puts it, to
integrate in one theory the supposed opposition between Unity of Type and Conditions of Existence.
But it is here that even the most sympathetic of Darwin's theistic supporters were forced to qualify their
support for the theory of descent with modification by means of natural selection. In Darwin's day the
reactions of Asa Gray and John Herschel are perhaps the most interesting in this respect. Both men saw
in Darwin's theory a way to account for that mystery of mysteries, the regular appearance of new
species by means of natural, or as they might say, intermediate causes. However both instinctively
recoiled from the irreducible and central role of chance in the theory. They did not, but easily could
have, said God does not play dice with the universe. But as Darwin stated repeatedly, if gently, to
Grayif God ordained that variations should be along beneficial lines, natural selection would be
redundant. Moreover, the evidence from the study of variation in domestic and natural populations put
the lie to any claim that God directs all or most variation along beneficial lines. Darwinian selection
theory is a two-step processthe production of variation unrelated to the adaptive requirements of the
organism, and differential perpetuation of those variations that serve adaptive needs. Again, a theory of
evolution that could not be so described would not be a Darwinian theory.
The nature of selection explanations is a topic to which much philosophical attention has been
devoted in recent years. Here I want to focus on only one important questionto what extent is the
teleological appearance of such explanations simply that, an appearance masking a causal process in
which goals play no role?
The appearance of teleology is certainly present in Darwinian explanations, and has been since Darwin
spoke of natural selection working solely for the good of each being. The appearance of teleology stems
from the ease with which both evolutionary biology and common sense take it for granted that animals
and plants have the adaptations they dobecause of some benefit or advantage to the organism provided
by those adaptations.
This is a hotly contested question, and I will here simply sketch a case that selective explanations of
adaptations are robustly teleological. The interested reader may want to refer to the literature on this
question referred to in the discussion and listed in the list of readings provided at the end of this entry.
A question I think not worth discussing is whether the word teleology should be replaced by
teleonomy. Etymologically, they come to the same thing; and the philosophical arguments given in
favor of the change all rest on an historically doubtful assumptionthat philosophical defenses of
teleology have always been either theistic or vitalistic. The serious philosophical issue can be put
simply and directly: in selection explanations of adaptations, are the functions served by adaptations a
central and irreducible feature of those explanations? If the answer is yes, the explanations are
teleological.[14]
A good place to begin is with a simple, yet realistic, example. In research carried out over many years
and combining painstaking field work and laboratory experimentation, John Endler was able to
demonstrate that the color patterns of males in the guppy populations he was studying in rivers feeding
into the southern Caribbean were a consequence of a balance between mate selection and predator
selection. To take one startling example, he was able to test and confirm a hypothesis that a group of
males, with a color pattern that matched that of the pebbles on the bottoms of the streams and ponds
they populated except for bright red spots, have that pattern because a common predator in those
populations, a prawn, is color blind for red. Red spots did not put their possessors at a selective
disadvantage, and were attractors for mates. (Endler 1983, 173190) We may refer to this pattern of
coloration as a complex adaptation that serves the functions of predator avoidance and mate attraction.
But what role do those functions play in explaining why it is that the males in this population have the
coloration they do?
This color pattern is an adaptation, as that term is used in Darwinism, only if it is a production of
natural selection (Williams 1966 261; Brandon 1985; Burian 1983). In order for it to be a product of
natural selection, there must be an array of color variation available in the genetic/developmental
resources of the species wider that this particular pattern but including this pattern. Which factors are
critical, then, in producing differential survival and reproduction of guppies with this particular pattern?
The answer would seem to be the value-consequences this pattern has compared to others available in
promoting viability and reproduction. In popular parlance (and the parlance favored by Darwin), this
color pattern isgood for the male guppies that have it, and for their male offspring, and that is why they
have it (Binswanger 1990; Brandon 1985; Lennox 2002). This answer strengthens the selected effects
or consequence etiology accounts of selection explanations by stressing that selection ranges
over value differences. The reason for one among a number of color patterns having a higher fitness
value has to do with the value of that pattern relative to the survival and reproductive success of its
possessors.
Selection explanations are, then, a particular kind of teleological explanation, an explanation in
which that for the sake of which a trait is possessed, its valuable consequence, accounts for the trait's
differential perpetuation and maintenance in the population.

3.4 Species and the Concept of Species

In listing the topics under which I would discuss neo-Darwinism, I distinguished the question of the
ontological status of species from the epistemological status of the species concept. Though they are
closely related questions, it is important to keep them distinct. As will become clear as we proceed, this
distinction is rarely honored. Moreover, it is equally important to distinguish the species concept from
the categories of features that belong in a definition of species. Advances in our theoretical
understanding may lead us to reconsider the sorts of attributes that are most important for determining
whether a group of organisms is a species, and thus whether it deserves to be assigned a name at that
taxonomic level. It should not be assumed that such changes constitute a change in the species concept,
though at least some such changes may lead us to restrict or expand the range of taxa that are
designated as species. In his contribution to the Neo-Darwinian Synthesis, Systematics and the Origin
of Species, Ernst Mayr titled chapter five The Systematic Categories and the New Species Concept.
Recall that Darwin made a point of treating the species category as continuous with well-marked
variety and sub-species, and made the radical suggestion that its boundaries would be just as fluid.
Without explicitly acknowledging Darwin, Mayr takes the same tack, discussing individual variants
and sub-species as a preliminary to discussing the species concept. Mayr notes that for someone
studying the evolutionary process, speciation is a critical juncture; his interpretation of the
speciation process depends largely on what he considers to be the final stage of this process, the
species. (Mayr 1942/1982, 113) With this in mind, he offers the following definition, the so-called
biological species concept (BSC):
Species are groups of actually or potentially interbreeding natural populations, which are
 reproductively isolated from other such groups (Mayr 1942/1982, 120; 1976 518)

Mayr was well aware of the limitations of this definition, and treated it somewhat as a regulative
ideal. Dobzhansky in 1937 gave what he claimed to be a definition of species, but which seems, as
Mayr noted (Mayr 1976 481), much more a definition of speciation:
that stage of evolutionary process at which the once actually or potentially interbreeding
array of forms becomes segregated in two or more separate arrays which are
physiologically incapable of interbreeding. (312)

Simpson (1943) and others built even more historicity into the concept. These are all, of course,
intended as definitions of the species category, and they attempt to provide a test (or a yardstick:
Mayr 1976 479) that in principle will permit a researcher to decide whether a group of individuals
should all be identified by a single species-level concept such as homo sapiens. The test for species
membership is the capacity to interbreed; the test distinguishing two species is incapacity to interbreed.
Dobzhansky makes the importance of this test transparentthe transition from a single interbreeding
population to two reproductively isolated ones is the process of speciation.
Now in each of these definitions, little attention is paid to the actual methods used by taxonomists and
systematists in differentiating between varieties of a species and distinct species, something to which
Darwin gave a great deal of attention. Darwins nominalism regarding the species concept likely
stemmed from his close attention to his own taxonomic practices and those of other specialists. But
nominalism typically combines a view about theontology of species with one about the epistemological
status of the species concept. On the first question, the nominalist insists that there are no species
there are more or less similar individuals. On the second question, the nominalist typically insists that
the species conceptis, at best, a useful or convenient grouping of similar individuals or, at worst,
an arbitrarygrouping of similar individuals.
In his work, Mayr relates different approaches to the species concept to the philosophical distinction
between essentialism and nominalism. He associates essentialism with the view that a
species concept refers to a universal or type. This view of the referent of the concept leads to the
Typological Species Concept, which he traces from Linnaeus back to Plato and Aristotle, and which he
claims is now universally abandoned. (1976 516; it is worth noting that serious doubt has been cast
both on the historical and the philosophical credentials of Mayrs Typological Species Concept (see,
e.g. Lennox, 1987; repr. in Lennox 2001b; Winsor 2001, 2006; Walsh 2006; Wilkins 2009) At the
opposite extreme is nominalism, which combines the view that only individuals exist in nature and that
species are concepts invented for the purpose of grouping these individuals collectively.
Mayr claims that his Biological Species Concept (BSC) is an advance on both; individual species
members are objectively related to one another not by a shared relation to a type but by causal and
historical relationships to one another. He can thus be understood as arguing for a new, objective way
of understanding the epistemological grounds for grouping individuals into species. This new way of
grouping stresses historical, genetic and various ecological relationships among the individuals as the
grounds for determining species membership. His claim is that this is more reliable and objective than
similarities of phenotypic characteristics. This makes sense of the importance he eventually places on
 the fact the BSC defines species relationally:
species are relationally defined. The word species corresponds very closely to other relational terms
such as, for instance, the word brother. To be a different species is not a matter of degree of
difference but of relational distinctness. (Mayr 1976 518)
Mayr has in mind that brothers may or may not look alike; the question of whether two people are
brothers is determined by their historical and genetic ties to a common ancestry. Notice, however, that
this is a claim about which characteristics, among the many that they have, should be taken most
seriously in determining the applicability to them of the concept brother. That is, it is a defense of a
sort of essentialism.
A number of critics have pointed out that essentialism need not be committed to types understood
as universalia in re; and on certain accounts of essences any species taxon that meets the standards of
BSC does so in virtue of certain essential (though relational and historical) properties. At one extreme,
Michael Ghiselin and David Hull have argued that this causal/historical structure of species provides
grounds, at least within evolutionary biology, for considering species to be individuals.[15] Organisms
are not members of a class or set, but parts of a phylogenetic unit. Taking a very different tack, Denis
Walsh has recently argued that a form of evolutionary essentialism, bearing a striking resemblance to
the essentialism of Aristotles zoological work, is implicit in the work of a number of evolutionary
developmental theorists. (Walsh, 2006)
A critical issue in this debate over the account of the species concept most appropriate for Darwinism is
the extent to which the process of biological classificationtaxonomyshould be informed by
advances in biological theory. Besides those already discussed, the moderate pluralism associated with
Robert Brandon and Brent Mischler or the more radical pluralism defended by Philip Kitcher, argues
that different explanatory aims within the biological sciences will require different criteria for
determining whether a group constitutes a species. Cladists, on the other hand, employ strictly defined
phylogenetic tests to determine species rank.
Unlike many of the other topics that define the history of Darwinism, there is no clear-cut position on
this question that can be identified as Darwinian or neo-Darwinian. In a recent collection of papers
defending most of the alternatives currently being advanced (Ereshefsky 1992), my suspicion is that
virtually every author in that collection would identify himself as Darwinian. This may be because, as
different as they are, a number of positions currently being defended have their roots in Darwins own
theory and practice (see Beatty 1985; reprinted in Ereshefsky 1992).

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 Further Reading

Charles Darwin's Life

Browne, E. J. 1995, Charles Darwin: A Biography. Vol. 1: Voyaging, Princeton: Princeton University
Press.
, 2000, Charles Darwin: A Biography. Vol. 2: The Power of Place, Princeton: Princeton University
Press.
Desmond, A. and Moore, J., 1992, Darwin: The Life of a Tormented Evolutionist, New York: Norton.
Herbert, S. 2005, Charles Darwin, Geologist, Ithaca: Cornell University Press.
Charles Darwin: Primary Sources
Barrett, P. H. (ed.), 1977, The Collected Papers of Charles Darwin, 2 Vols., Chicago: University of
Chicago Press.
Burkhardt, F. (ed.), 19852015, The Correspondence of Charles Darwin, Volumes 121, Cambridge:
Cambridge University Press.
Peckham, M. (ed.), 1959, The Origin of Species by Charles Darwin: A Variorum Text, Philadelphia:
University of Pennsylvania Press. [1st Paperback edition, 2006]
Weinshank, D. et al. (eds.), 1990, A Concordance to Charles Darwin's Notebooks, 18361844, Ithaca:
Cornell University Press.
Charles Darwin's Context
Hodge, J. and Radick, G. (eds.), 2003, The Cambridge Companion to Darwin, Cambridge: Cambridge
University Press.
Owen, R., 1837/1992, The Hunterian Lectures in Comparative Anatomy, 1837, Chicago: Chicago
University Press.
Rudwick, M., 1997, George Cuvier, Fossil Bones and Geological Catastrophes, Chicago: University of
Chicago Press.
Ruse, M., 1999, The Darwinian Revolution: Science Red in Tooth and Claw (Revised edition),
Cambridge: Cambridge University Press.
Ruse, M. and Richards, R. J. (eds.), 2009, The Cambridge Companion to the "Origin of Species",
Cambridge: Cambridge University Press.
Snyder, L., 2010, The Philosophical Breakfast Club, New York: Broadway Books.
The Evolution of Darwinism
Amundson, R., 2005, The Changing Role of the Embryo in Evolutionary Thought: Roots of Evo-Devo,
Cambridge: Cambridge University Press.
Depew, D. and Weber, B., 1995, Darwinism Evolving: Systems Dynamics and the Genealogy of
Natural Selection, Cambridge MA: MIT Press.
Kohn, D. (ed.), 1995, The Darwinian Heritage, Princeton: Princeton University Press.
Mayr, E., 1976, Evolution and the Diversity of Life, Cambridge MA: Harvard University Press.
Philosophy and Evolutionary Theory
Brandon, R. N., 1996, Concepts and Methods in Evolutionary Biology, Cambridge: Cambridge
University Press.
 Burian, R. M., 2005, The Epistemology of Development, Evolution, and Genetics, Cambridge:
Cambridge University Press.
Hull, D. and Ruse, M. (eds.), 1998, The Philosophy of Biology, Oxford: Oxford University Press.
Lloyd, E., 1994, The Structure and Confirmation of Evolutionary Theory, 2nd edition Princeton:
Princeton University Press.
Sober, E., 1984, The Nature of Selection: Evolutionary Theory in Philosophical Focus, Cambridge
MA: MIT Press.
, (ed.), 1994, Conceptual Issues in Evolutionary Biology, 2nd edition, Cambridge MA: MIT Press.
, 2008, Evidence and Evolution: The Logic Behind the Science, Cambridge: Cambridge University
Press.

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Other Internet Resources

By googling Darwinism, one quickly discovers that the world wide web has an abundance of web
sites and pages that do not meet high academic standards. However below are three useful sites. The
first is the official site for the publication of material in the extensive Darwin Archives at Cambridge
University, but has grown to become the default site for Darwin texts and related literature as well. The
second is the official site for on-line publication of Darwin's extensive correspondence. The third site is
a very good starting point for linking you to sites related to Charles Darwin's historical context.
Complete World of Charles Darwin Online
Darwin Correspondence Project
Victorian Science: An Overview, The Victorian Web (funded by the University Scholars
Program, National University of Singapore)

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adaptationism | biology: philosophy of | creationism | developmental biology: evolution and
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of | scientific explanation | Scottish Philosophy: in the 18th Century | Scottish Philosophy: in the 19th
century | species | teleology: teleological notions in biology |Whewell, William
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James Lennox <jglennox+@pitt.edu>